Ecology

Ebner, J., Babbitt, C., Winer, M., Hilton, B. & Williamson, A. Life cycle greenhouse gas (GHG) impacts of a novel process for converting food waste to ethanol and co-products. Appl. Energy 130, 86–93 (2014).CAS 

Google Scholar 
Tonini, D., Albizzati, P. F. & Astrup, T. F. Environmental impacts of food waste: Learnings and challenges from a case study on UK. Waste Manag. 76, 744 (2018).PubMed 

Google Scholar 
Sze, E., Yau, Y. H. & Wu, K. C. Application of anaerobic bacterial ammonification pretreatment to microalgal food waste leachate cultivation and biofuel production. Mar. Pollut. Bull. 153, 111007 (2020).PubMed 

Google Scholar 
Winkel, T. D., Wahlen, S. & Jensen, T. in Nordic Conference on Consumer Research.Wang, P. et al. Effects of graphite, graphene, and graphene oxide on the anaerobic co-digestion of sewage sludge and food waste: attention to methane production and the fate of antibiotic resistance genes. Bioresour. Technol. 339, 125585 (2021).CAS 
PubMed 

Google Scholar 
Gianico, A., Gallipoli, A., Pagliaccia, P. & Braguglia, C. M. Anaerobic bioconversion of food waste into energy: A critical review (2013).Smetana, S., Ites, S., Parniakov, O., Aganovic, K. & Heinz, V. in 71st Annual Meeting of the European Federation of Animal Science.Ojha, S., Buler, S. & Schlüter, O. Food waste valorisation and circular economy concepts in insect production and processing. Waste Manag. 118, 600–609 (2020).CAS 
PubMed 

Google Scholar 
Scala, A. et al. Rearing substrate impacts growth and macronutrient composition of Hermetia illucens (L.) (Diptera: Stratiomyidae) larvae produced at an industrial scale. Sci. Rep. 10, 1–8 (2020).ADS 

Google Scholar 
McDonald, C., Campbell, K. A., Benson, C., Davis, M. J. & Frost, C. J. Workforce development and multiagency collaborations: a presentation of two case studies in child welfare. Sustainability 13, 10190 (2021).
Google Scholar 
Kim, C.-H. et al. Use of black soldier fly larvae for food waste treatment and energy production in asian countries: a review. Processes 9, 161 (2021).CAS 

Google Scholar 
Julita, U., Fitri, L., Putra, R. & Permana, A. Mating success and reproductive behavior of black soldier fly Hermetia illucens L. (diptera, stra-tiomyidae) in tropics. J. Ento-mol. 17, 117–127 (2020).CAS 

Google Scholar 
Rehman, K. U. et al. Conversion of mixtures of dairy manure and soybean curd residue by black soldier fly larvae (Hermetia illucens L.). J. Clean. Prod. 154, 366–373 (2017).CAS 

Google Scholar 
Li, Q., Zheng, L., Hao, C., Garza, E. & Zhou, S. From organic waste to biodiesel: Black soldier fly, Hermetia illucens, makes it feasible. Fuel 90, 1545–1548 (2011).CAS 

Google Scholar 
Köhler, R., Kariuki, L., Lambert, C. & Biesalski, H. K. Protein, amino acid and mineral composition of some edible insects from Thailand. J. Asia Pac. Entomol. 22, 372–378 (2019).
Google Scholar 
Belghit, I. et al. Black soldier fly larvae meal can replace fish meal in diets of sea-water phase Atlantic salmon (Salmo salar). Aquaculture (2018).Moretta, A. et al. Antimicrobial peptides: A new hope in biomedical and pharmaceutical fields. Front. Cell. Infect. Microbiol. 11, 453 (2021).
Google Scholar 
Manniello, M. et al. Insect antimicrobial peptides: potential weapons to counteract the antibiotic resistance. Cell. Mol. Life Sci. 89, 1–24 (2021).
Google Scholar 
Henriques, B. S., Garcia, E. S., Azambuja, P. & Genta, F. A. Determination of chitin content in insects: an alternate method based on calcofluor staining. Front. Physiol. 11, ARTN 11710.3389/fphys.2020.00117 (2020).Hbl, M., Mráz, P., Ipo, J., Hotiková, I. & Kopec, T. Polyphenols as food supplement improved food consumption and longevity of honey bees (Apis mellifera) intoxicated by pesticide thiacloprid. Insects 12 (2021).Li, H., Dai, C., Zhu, Y. & Hu, Y. Larvae crowding increases development rate, improves disease resistance, and induces expression of antioxidant enzymes and heat shock proteins in Mythimna separata (Lepidoptera: Noetuidae). J. Econ. Entomol. 4 (2021).Hao, et al. Effects of enzymatic hydrolysis assisted by high hydrostatic pressure processing on the hydrolysis and allergenicity of proteins from ginkgo seeds. Food Bioprocess Technol. 9, 839–848 (2016).
Google Scholar 
Nadeem, M., Mumtaz, M. W., Danish, M., Rashid, U. & Raza, S. A. Calotropis procera: UHPLC-QTOF-MS/MS based profiling of bioactives, antioxidant and anti-diabetic potential of leaf extracts and an insight into molecular docking. J. Food Meas. Charact. 13, 3206–3220 (2019).
Google Scholar 
Altomare, A. A., Baron, G., Aldini, G., Carini, M. & D’Amato, A. Silkworm pupae as source of high-value edible proteins and of bioactive peptides. Food Sci. Nutr. 8, 2652–2661 (2020).CAS 
PubMed 
PubMed Central 

Google Scholar 
Zielińska, E., Baraniak, B. & Karaś, M. Identification of antioxidant and anti-inflammatory peptides obtained by simulated gastrointestinal digestion of three edible insects species (Gryllodes sigillatus, Tenebrio molitor, Schistocerca gragaria). Int. J. Food Sci. Technol. 53, 2542–2551 (2018).
Google Scholar 
Sousa, P., Borges, S. & Pintado, M. Enzymatic hydrolysis of insect Alphitobius diaperinus towards the development of bioactive peptide hydrolysates. Food Funct. 11, 3539–3548 (2020).CAS 
PubMed 

Google Scholar 
Jakubczyk, A., Karaś, M., Rybczyńska-Tkaczyk, K., Zielińska, E. & Zieliński, D. Current trends of bioactive peptides—New sources and therapeutic effect. Foods 9, 846 (2020).CAS 
PubMed Central 

Google Scholar 
Li, K., Li, X.-M., Ji, N.-Y. & Wang, B.-G. Natural bromophenols from the marine red alga Polysiphonia urceolata (Rhodomelaceae): structural elucidation and DPPH radical-scavenging activity. Bioorg. Med. Chem. 15, 6627–6631 (2007).CAS 
PubMed 

Google Scholar 
He, R., Girgih, A. T., Malomo, S. A., Ju, X. R. & Aluko, R. E. Antioxidant activities of enzymatic rapeseed protein hydrolysates and the membrane ultrafiltration fractions. J. Funct. Foods 5, 219–227. https://doi.org/10.1016/j.jff.2012.10.008 (2013).CAS 
Article 

Google Scholar 
Cui, Q., Sun, Y. X., Cheng, J. J. & Guo, M. R. Effect of two-step enzymatic hydrolysis on the antioxidant properties and proteomics of hydrolysates of milk protein concentrate. Food Chem. 366, 10. https://doi.org/10.1016/j.foodchem.2021.130711 (2022).CAS 
Article 

Google Scholar 
Liu, Y., Wan, S., Liu, J., Zou, Y. & Liao, S. Antioxidant activity and stability study of peptides from enzymatically hydrolyzed male silkmoth. J. Food Process. Preserv. 41 (2017).Carrasco-Castilla, J. et al. Antioxidant and metal chelating activities of peptide fractions from phaseolin and bean protein hydrolysates. Food Chem. 135, 1789–1795 (2012).CAS 
PubMed 

Google Scholar 
Phongthai, S., D’Amico, S., Schoenlechner, R., Homthawornchoo, W. & Rawdkuen, S. Fractionation and antioxidant properties of rice bran protein hydrolysates stimulated by in vitro gastrointestinal digestion. Food Chem. 240, 156 (2018).CAS 
PubMed 

Google Scholar 
Lee, S. J. et al. Antioxidant activity of a novel synthetic hexa-peptide derived from an enzymatic hydrolysate of duck skin by-products. Food Chem. Toxicol. 62, 276–280 (2013).CAS 
PubMed 

Google Scholar 
Collin, F. Chemical basis of reactive oxygen species reactivity and involvement in neurodegenerative diseases. Int. J. Mol. Sci. 20, 2407 (2019).PubMed Central 

Google Scholar 
Xiang, Q., Yu, J. & Wong, P. K. Quantitative characterization of hydroxyl radicals produced by various photocatalysts. J. Colloid Interface Sci. 357, 163–167 (2011).ADS 
CAS 
PubMed 

Google Scholar 
Wang, Y. et al. Optimizing oxygen functional groups in graphene quantum dots for improved antioxidant mechanism. Phys. Chem. Chem. Phys. 21, 1336–1343 (2019).CAS 
PubMed 

Google Scholar 
Arise, A. K. et al. Antioxidant activities of bambara groundnut (Vigna subterranea) protein hydrolysates and their membrane ultrafiltration fractions. Food Funct. 7, 2431–2437. https://doi.org/10.1039/c6fo00057f (2016).CAS 
Article 
PubMed 

Google Scholar 
Ren, J. et al. Purification and identification of antioxidant peptides from grass carp muscle hydrolysates by consecutive chromatography and electrospray ionization-mass spectrometry. Food Chem. 108, 727–736. https://doi.org/10.1016/j.foodchem.2007.11.010 (2008).CAS 
Article 
PubMed 

Google Scholar 
Zhang, C., Wei, X., Omenn, G. S. & Zhang, Y. Structure and protein interaction-based gene ontology annotations reveal likely functions of uncharacterized proteins on human chromosome 17. 17 (2018).Long, C. N. et al. High-level production of Monascus pigments in Monascus ruber CICC41233 through ATP-citrate lyase overexpression. Biochem. Eng. J. 146, 160–169. https://doi.org/10.1016/j.bej.2019.03.007 (2019).CAS 
Article 

Google Scholar 
Brito Querido, J. et al. The cryo-EM structure of a novel 40S kinetoplastid-specific ribosomal protein. Structure 25, 1785–1794 e1783. https://doi.org/10.1016/j.str.2017.09.014 (2017).Hamey, J. J. & Wilkins, M. R. Methylation of elongation factor 1A: Where, who, and why?. Trends Biochem. Sci. 43, 211–223. https://doi.org/10.1016/j.tibs.2018.01.004 (2018).CAS 
Article 
PubMed 

Google Scholar 
Kuo, C. P. et al. Analysis of the immune response of human dendritic cells to Mycobacterium tuberculosis by quantitative proteomics. Proteome Sci. 14, 1–11 (2016).
Google Scholar 
Zhu, J. et al. Expression and RNA interference of ribosomal protein L5 gene in Nilaparvata lugens (Hemiptera: Delphacidae). J. Insect. Sci. 3 (2017).Teng, T., Mercer, C. A., Hexley, P., Thomas, G. & Fumagalli, S. Loss of tumor suppressor RPL5/RPL11 does not induce cell cycle arrest but impedes proliferation due to reduced ribosome content and translation capacity. Mol. Cell. Biol. 33, 4660–4671. https://doi.org/10.1128/mcb.01174-13 (2013).CAS 
Article 
PubMed 
PubMed Central 

Google Scholar 
Rittschof, C. C. & Schirmeier, S. Insect models of central nervous system energy metabolism and its links to behavior. Glia (2017).Alar, A. F., Akr, B. & Gülseren, I. LC-Q-TOF/MS based identification and in silico verification of ACE-inhibitory peptides in Giresun (Turkey) hazelnut cakes. Eur. Food Res. Technol. (2021).Shang, W. H. et al. In silico assessment and structural characterization of antioxidant peptides from major yolk protein of sea urchin Strongylocentrotus nudus. Food Funct. 9, 6435–6443 (2018).CAS 
PubMed 

Google Scholar 
Ajibola, C. F., Fashakin, J. B., Fagbemi, T. N. & Aluko, R. E. Effect of peptide size on antioxidant properties of African yam bean seed (Sphenostylis stenocarpa) protein hydrolysate fractions. Int. J. Mol. Sci. 12, 6685–6702 (2011).CAS 
PubMed 
PubMed Central 

Google Scholar 
Liu, H. et al. Enhancing the antioxidative effects of foods containing rutin and α-amino acids via the Maillard reaction: A model study focusing on rutin-lysine system. J. Food Biochem. 44, e13086 (2020).PubMed 

Google Scholar 
Tsopmo, A. et al. Tryptophan released from mother’s milk has antioxidant properties. Pediatr. Res. 66, 614–618 (2009).CAS 
PubMed 

Google Scholar 
Yu, Z. et al. Identification and molecular docking study of fish roe-derived peptides as potent BACE 1, AChE, and BChE inhibitors. Food Funct. 11 (2020).Li, C. et al. Preliminary study on a potential antibacterial peptide derived from histone H2A in hemocytes of scallop Chlamys farreri. Fish Shellf. Immunol. 22, 663–672 (2007).
Google Scholar 
Arockiaraj, J. et al. An unconventional antimicrobial protein histone from freshwater prawn Macrobrachium rosenbergii: analysis of immune properties. Fish Shellfish Immunol. 35, 1511–1522 (2013).CAS 
PubMed 

Google Scholar 
Ju, J. et al. Major components in Lilac and Litsea cubeba essential oils kill Penicillium roqueforti through mitochondrial apoptosis pathway. Ind. Crops Products 149, 112349 (2020).CAS 

Google Scholar 
Al-Dhafri, K., Chai, L. C. & Karsani, S. A. Purification and characterization of antimicrobial peptide fractions of Junipers seravschanica. Biocatal. Agric. Biotechnol. 28, 101554 (2020).
Google Scholar 
Ratnakomala, S., Ridwan, R., Lisdiyanti, P., Abinawanto, A. & Andi, U. Screening of actinomycetes producing an ATPase inhibitor of japanese encephalitis virus RNA helicase from soil and leaf litter samples. Microbiol. Indonesia 5, 15–20 (2011).
Google Scholar 
Zhao, X., Zhang, J. & Zhu, K. Y. Chito-protein matrices in arthropod exoskeletons and peritrophic matrices (2019).Pustylnikov, S., Sagar, D., Jain, P. & Khan, Z. K. Targeting the C-type lectins-mediated host-pathogen interactions with dextran. J. Pharm. Pharm. Sci. 17 (2014).Kiew, P. L. & Don, M. M. Jewel of the seabed: sea cucumbers as nutritional and drug candidates. Int. J. Food Sci. Nutr. 63, 616–636 (2012).CAS 
PubMed 

Google Scholar 
Liu, Z., Su, Y. & Zeng, M. Amino acid composition and functional properties of giant red sea cucumber (Parastichopus californicus) collagen hydrolysates. J. Ocean Univ. China 10, 80–84 (2011).ADS 
CAS 

Google Scholar 
Zaky, A. A., Liu, Y., Han, P., Chen, Z. & Jia, Y. Effect of pepsin–trypsin in vitro gastro-intestinal digestion on the antioxidant capacities of ultra-filtrated rice bran protein hydrolysates (molecular weight > 10 kDa; 3–10 kDa, and< 3 kDa). Int. J. Pept. Res. Ther. 1–7 (2019).Kim, S.-B., Yoon, N. Y., Shim, K.-B. & Lim, C.-W. Antioxidant and angiotensin I-converting enzyme inhibitory activities of northern shrimp (Pandalus borealis) by-products hydrolysate by enzymatic hydrolysis. Fish. Aquat. Sci. 19, 1–6 (2016). Google Scholar  Chung, Y. C., Chang, C. T., Chao, W. W., Lin, C. F. & Chou, S. T. Antioxidative activity and safety of the 50 ethanolic extract from red bean fermented by Bacillus subtilis IMR-NK1. J. Agric. Food Chem. 50, 2454–2458. https://doi.org/10.1021/jf011369q (2002).CAS  Article  PubMed  Google Scholar  Zielińska, E., BaRaniak, B. & Karaś, M. Antioxidant and anti-inflammatory activities of hydrolysates and peptide fractions obtained by enzymatic hydrolysis of selected heat-treated edible insects. Nutrients 9, 1–14 (2017). Google Scholar  Rahman, M. M., Byanju, B., Grewell, D. & Lamsal, B. P. High-power sonication of soy proteins: Hydroxyl radicals and their effects on protein structure. Ultrason. Sonochem. 64, 105019 (2020).CAS  PubMed  Google Scholar  More

Wu, X. et al. Effects of bio-organic fertiliser fortified by Bacillus cereus QJ-1 on tobacco bacterial wilt control and soil quality improvement. Biocontrol Sci. Technol. 30, 351–369 (2020).
Google Scholar 
Hu, W. et al. Flue-cured tobacco (Nicotiana tabacum L.) leaf quality can be improved by grafting with potassium-efficient rootstock. Field Crop. Res. 274, 108305 (2021).
Google Scholar 
Wu, X. et al. Bioaugmentation of Bacillus amyloliquefaciens-Bacillus kochii co-cultivation to improve sensory quality of flue-cured tobacco. Arch. Microbiol. 203, 5723–5733 (2021).CAS 
PubMed 

Google Scholar 
Jiang, C. et al. Optimal lime application rates for ameliorating acidic soils and improving the yield and quality of tobacco leaves. Appl. Ecol. Environ. Res. 18, 5411–5423 (2020).
Google Scholar 
Yin, Q. et al. Investigation of associations between rhizosphere microorganisms and the chemical composition of flue-cured tobacco leaves using canonical correlation analysis. Commun. Soil Sci. Plant 44, 1524–1539 (2013).CAS 

Google Scholar 
Shen, H. et al. Promotion of lateral root growth and leaf quality of flue-cured tobacco by the combined application of humic acids and npk chemical fertilizers. Exp. Agric. 53, 59–70 (2017).CAS 

Google Scholar 
Hu, W. Q. et al. Grafting alleviates potassium stress and improves growth in tobacco. BMC Plant Biol. 19, 130 (2019).PubMed 
PubMed Central 

Google Scholar 
Zhong, J. Study of K+ uptake kinetics of flue-cured tobacco in K+-enriched and conventional tobacco genotypes. J. Plant Nutr. 42(7), 1–7 (2019).
Google Scholar 
Tang, Z. et al. Climatic factors determine the yield and quality of Honghe flue-cured tobacco. Sci. Rep. 10, 19868 (2020).ADS 
CAS 
PubMed 
PubMed Central 

Google Scholar 
Yan, S. et al. Correlation between soil microbial communities and tobacco aroma in the presence of different fertilizers. Ind. Crop. Prod. 151, 112454 (2020).CAS 

Google Scholar 
Tabaxi, I. Effect of organic fertilization on quality and yield of oriental tobacco (Nicotiana tabacum L.) under Mediterranean conditions. Asian J. Agric. Biol. https://doi.org/10.35495/ajab.2020.05.274 (2021).Article 

Google Scholar 
Chen, Y. L. et al. Distinct microbial communities in the active and permafrost layers on the Tibetan Plateau. Mol. Ecol. 26, 6608–6620 (2017).CAS 
PubMed 

Google Scholar 
Wagg, C. et al. Soil biodiversity and soil community composition determine ecosystem multifunctionality. Proc. Natl. Acad. Sci. U.S.A. 111, 5266–5270 (2014).ADS 
CAS 
PubMed 
PubMed Central 

Google Scholar 
French, E. et al. Emerging strategies for precision microbiome management in diverse agroecosystems. Nat. Plants 7, 256–267 (2021).PubMed 

Google Scholar 
Cui, Y. et al. Diversity patterns of the rhizosphere and bulk soil microbial communities along an altitudinal gradient in an alpine ecosystem of the eastern Tibetan Plateau. Geoderma 338, 118–127 (2019).ADS 
CAS 

Google Scholar 
Zheng, J. et al. The effects of tetracycline residues on the microbial community structure of tobacco soil in pot experiment. Sci. Rep. 10, 8804 (2020).ADS 
CAS 
PubMed 
PubMed Central 

Google Scholar 
Zhang, J. Effects of tobacco planting systems on rates of soil N transformation and soil microbial community. Int. J. Agric. Biol. 19, 992–998 (2017).CAS 

Google Scholar 
Yang, Y. et al. Metagenomic insights into effects of wheat straw compost fertiliser application on microbial community composition and function in tobacco rhizosphere soil. Sci. Rep. 9, 6168 (2019).ADS 
PubMed 
PubMed Central 

Google Scholar 
Wang, S. et al. Response of soil fungal communities to continuous cropping of flue-cured tobacco. Sci. Rep. 10, 19911 (2020).ADS 
CAS 
PubMed 
PubMed Central 

Google Scholar 
Finlay, B. J. Global dispersal of free-living microbial eukaryote species. Science 296, 1061–1063 (2002).ADS 
CAS 
PubMed 

Google Scholar 
Ehrmann, J. & Ritz, K. Plant: soil interactions in temperate multi-cropping production systems. Plant Soil 376, 1–29 (2013).
Google Scholar 
Liu, H. et al. Response of soil fungal community structure to long-term continuous soybean cropping. Front. Microbiol. 9, 3316 (2018).PubMed 

Google Scholar 
Gao, Z. et al. Effects of continuous cropping of sweet potato on the fungal community structure in rhizospheric soil. Front. Microbiol. 10, 2269 (2019).PubMed 
PubMed Central 

Google Scholar 
Li, J. Analysis on the method selection of tobacco disease control. South China Agric. 15, 49–50 (2021).
Google Scholar 
Dai, C. et al. Comprehensive evaluation of soil fertility status of tobacco-planting district in Bijie area. Acta Agric. Jiangxi 23, 9–11 (2011).
Google Scholar 
Wang, Z. et al. Time-course relationship between environmental factors and microbial diversity in tobacco soil. Sci. Rep. 9, 19969 (2019).ADS 
CAS 
PubMed 
PubMed Central 

Google Scholar 
Wang, X. et al. Study on the primary chemical components, sensory quality of flue-cured tobacco and their correlativity in Bijie. J. Henan Agric. Sci. 41, 58–61 (2012).
Google Scholar 
Zhang, S. et al. Analysis of variation characteristics and coordination of conventional chemical components of flue-cured tobacco in Youyang County, Chongqing City. Acta Agric. Jiangxi 32, 75–86 (2020).
Google Scholar 
Lakatos, L. et al. The influence of meteorological variables on sour cherry quality parameters. In Vi International Cherry Symposium (Int Soc Horticultural Science, Leuven 1), Vol. 1020, 287–292 (2014).Zhao, Z. et al. Why does potassium concentration in flue-cured tobacco leaves decrease after apex excision? Field Crop. Res. 116, 86–91 (2010).
Google Scholar 
Travlos, I. S. et al. Green manure and pendimethalin impact on oriental sun-cured tobacco. Agron. J. 106, 1225–1230 (2014).
Google Scholar 
Bilalis, D. et al. Effect of organic fertilization on soil characteristics, yield and quality of Virginia Tobacco in Mediterranean area. Emir. J. Food Agric. https://doi.org/10.9755/ejfa.2020.v32.i8.2138 (2020).Article 

Google Scholar 
Henry, J. B., Vann, M. C. & Lewis, R. S. Agronomic practices affecting nicotine concentration in flue-cured tobacco: A review. Agron. J. 111, 3067–3075 (2019).CAS 

Google Scholar 
Lamarre, M. & Payette, S. Influence of nitrogen-fertilization on Quebec flue-cured tobacco production. Can. J. Plant Sci. 72, 411–419 (1992).CAS 

Google Scholar 
Zhang, L. et al. Dynamic changes of nutrients in tobacco-planting soils in Bijie City during 2014–2016. Guizhou Agric. Sci. 45, 51–55 (2017).CAS 

Google Scholar 
Lisuma, J. B., Mbega, E. R. & Ndakidemi, P. A. Dynamics of nicotine across the soil–tobacco plant interface is dependent on agro-ecology, nitrogen source, and rooting depth. Rhizosphere 12, 100175 (2019).
Google Scholar 
Cosme, M. & Wurst, S. Interactions between arbuscular mycorrhizal fungi, rhizobacteria, soil phosphorus and plant cytokinin deficiency change the root morphology, yield and quality of tobacco. Soil Biol. Biochem. 57, 436–443 (2013).CAS 

Google Scholar 
Chandanie, W. A., Kubota, M. & Hyakumachi, M. Interactions between the arbuscular mycorrhizal fungus Glomus mosseae and plant growth-promoting fungi and their significance for enhancing plant growth and suppressing damping-off of cucumber (Cucumis sativus L.). Appl. Soil Ecol. 41, 336–341 (2009).
Google Scholar 
Liu, D. et al. Geographic distance and soil microbial biomass carbon drive biogeographical distribution of fungal communities in Chinese Loess Plateau soils. Sci. Total Environ. 660, 1058–1069 (2019).ADS 
CAS 
PubMed 

Google Scholar 
Li, S. & Wu, F. Diversity and co-occurrence patterns of soil bacterial and fungal communities in seven intercropping systems. Front. Microbiol. https://doi.org/10.3389/fmicb.2018.01521 (2018).Article 
PubMed 
PubMed Central 

Google Scholar 
Li, H. et al. Chaetosemins A-E, new chromones isolated from an Ascomycete Chaetomium seminudum and their biological activities. RSC Adv. 5, 29185–29192 (2015).ADS 
CAS 

Google Scholar 
Gorte, O., Kugel, M. & Ochsenreither, K. Optimization of carbon source efficiency for lipid production with the oleaginous yeast Saitozyma podzolica DSM 27192 applying automated continuous feeding. Biotechnol. Biofuels 13, 181 (2020).CAS 
PubMed 
PubMed Central 

Google Scholar 
Zhang, J. et al. Biochar applied to consolidated land increased the quality of an acid surface soil and tobacco crop in Southern China. J. Soil. Sediment. 20, 3091–3102 (2019).
Google Scholar 
Zong, J. et al. Effect of two drying methods on chemical transformations in flue-cured tobacco. Dry Technol. https://doi.org/10.1080/07373937.2020.1779287 (2020).Article 

Google Scholar 
Ismail, E. et al. Evaluation of in vitro antifungal activity of potassium bicarbonate on Rhizoctonia solani AG 4 HG-I, Sclerotinia sclerotiorum and Trichoderma sp.. Afr. J. Biotechnol. 10, 8605–8612 (2011).
Google Scholar 
d’Aquino, L. et al. Effect of some rare earth elements on the growth and lanthanide accumulation in different Trichoderma strains. Soil Biol. Biochem. 41, 2406–2413 (2009).CAS 

Google Scholar 
Bijie Municipal Government. Statistical Yearbooks of Bijie City (2016)He, R., Liu, S. & Liu, Y. Application of SD model in analyzing the cultivated land carrying capacity: A case study in Bijie Prefecture, Guizhou Province, China. Procedia Environ. Sci. 10, 1985–1991 (2011).
Google Scholar 
Wang, M. et al. Spatial variation and fractionation of fluoride in tobacco-planted soils and leaf fluoride concentration in tobacco in Bijie City, Southwest China. Environ. Sci. Pollut. Res. 28, 26112–26123 (2021).CAS 

Google Scholar 
Wang, J. T. et al. Altitudinal distribution patterns of soil bacterial and archaeal communities along Mt. Shegyla on the Tibetan Plateau. Microb. Ecol. 69, 135–145 (2015).PubMed 

Google Scholar 
Zhang, Q. et al. Soil available phosphorus content drives the spatial distribution of archaeal communities along elevation in acidic terrace paddy soils. Sci. Total Environ. 658, 723–731 (2019).ADS 
CAS 
PubMed 

Google Scholar 
Cui, Q. et al. Sulfur application improved leaf yield and quality of flue-cured tobacco by maintaining soil sulfur balance. Int. J. Agric. Biol. 23, 357–363 (2020).CAS 

Google Scholar 
Cao, X. et al. Distribution, availability and translocation of heavy metals in soil-oilseed rape (Brassica napus L.) system related to soil properties. Environ. Pollut. 252, 733–741 (2019).CAS 
PubMed 

Google Scholar 
Wang, C. et al. Prevalence of antibiotic resistance genes and bacterial pathogens along the soil-mangrove root continuum. J. Hazard. Mater. 408, 124985 (2021).CAS 
PubMed 

Google Scholar 
Magoc, T. & Salzberg, S. L. FLASH: Fast length adjustment of short reads to improve genome assemblies. Bioinformatics 27, 2957–2963 (2011).CAS 
PubMed 
PubMed Central 

Google Scholar 
Bolger, A. M., Lohse, M. & Usadel, B. Trimmomatic: A flexible trimmer for Illumina sequence data. Bioinformatics 30, 2114–2120 (2014).CAS 
PubMed 
PubMed Central 

Google Scholar 
Edgar, R. C. et al. UCHIME improves sensitivity and speed of chimera detection. Bioinformatics 27, 2194–2200 (2011).CAS 
PubMed 
PubMed Central 

Google Scholar 
Edgar, R. C. UPARSE: Highly accurate OTU sequences from microbial amplicon reads. Nat. Methods 10, 996–998 (2013).CAS 
PubMed 

Google Scholar 
Bokulich, N. A. et al. Quality-filtering vastly improves diversity estimates from Illumina amplicon sequencing. Nat. Methods 10, 57–59 (2013).CAS 
PubMed 

Google Scholar 
Wang, Q. et al. Naive Bayesian classifier for rapid assignment of rRNA sequences into the new bacterial taxonomy. Appl. Environ. Microbiol. 73, 5261–5267 (2007).ADS 
CAS 
PubMed 
PubMed Central 

Google Scholar 
Koljalg, U. et al. Towards a unified paradigm for sequence-based identification of fungi. Mol. Ecol. 22, 5271–5277 (2013).CAS 
PubMed 

Google Scholar 
Bolyen, E. et al. Reproducible, interactive, scalable and extensible microbiome data science using QIIME 2. Nat. Biotechnol. 37, 852–857 (2019).CAS 
PubMed 
PubMed Central 

Google Scholar 
Lindstrom, E. S. et al. Distribution of typical freshwater bacterial groups is associated with pH, temperature, and lake water retention time. Appl. Environ. Microbiol. 71, 8201–8206 (2005).ADS 
PubMed 
PubMed Central 

Google Scholar  More

Phylogenetic analysesA total of 21 sequences were newly generated and deposited in GenBank (Supplementary Table 1). The concatenated sequence alignment of the three loci comprised 100 sequences (38 for ITS, 30 for rpb2 and 32 for mtSSU) from 43 collections (Supplementary Table 1). The alignment was 2,004 characters long, including gaps. Multi-locus trees generated from ML and BI analyses showed similar topologies without any supported topological conflict. The multi-locus phylogeny (Fig. 1) confirmed placement of all Thai collections within the well-supported R. subsect. Amoeninae (ML = 99, BI = 1.0). Five collections from northeastern Thailand and two collections from northern Thailand form two strongly supported clades and are described below as the new species R. bellissima sp. nov. and R. luteonana sp. nov. The new species are not resolved as sister. The first species, R. bellissima, is strongly supported as sister to a clade of Australian sequestrate species that includes R. variispora T. Lebel and an undescribed Russula sp. labeled as Macowanites sp. The Indian species R. intervenosa S. Paloi, A.K. Dutta & K. Acharya is placed as sister to them with bootstrap support of 77. The second species, R. luteonana, is placed with moderate support as sister to the sequestrate European species R. andaluciana T.F. Elliott & Trappe.Figure 1ML phylogenetic tree inferred from the three-gene dataset (ITS, rpb2, mtSSU) of Russula subsection Amoeninae species, using ML and BI analyses. Three members of R. subg. Heterophyllidiae are used as outgroup. Species in boldface are new species in this study. Bootstrap support values (BS ≥ 50%) and posterior probabilities (PP ≥ 0.90) are shown at the supported branches.Full size imageThe ITS tree (Fig. 2) shows a similar topology and relationships for the studied specimens. In addition, R. intervenosa received good support (ML = 84, BI = 0.99) as sister to the clade of R. bellissima and R. variispora. Five additional ITS sequences that are grouped with strong support within R. bellissima species clade were recovered, three from Thailand, one from Laos, and one from Singapore. We did not recover any other Amoeninae ITS sequences from Thailand.Figure 2ML phylogenetic tree inferred from the ITS region of Russula subsection Amoeninae species and allied groups, using ML and BI methods. Samples in boldface are new species in this study. Bootstrap support values (BS ≥ 50%) and posterior probabilities (PP ≥ 0.90) are shown at the supported branches.Full size imageTaxonomy
Russula bellissima Manz & F. Hampe sp. nov.
Mycobank: MB 840549Holotype THAILAND, Theong district, Chiang Rai, 19°36′45”N 100°4′00”E, alt. 500 m, dry dipterocarpus forest in small groups on loamy soil, 12 July 2012, F. Hampe (Holotype: GENT FH 12-127; Isotype: MFLU12-0619).Etymology ’bellus’ = latin for beautiful, pretty, lovely; ’bellissima’ = the most beautiful. Resembling the species Russula bella which is also belonging to Russula subsection Amoeninae.Diagnosis Pileus small to medium-sized; cuticle dry, smooth, matt and pruinose, red; stipe white or with a red flush; spore ornamentation of moderately distant to dense amyloid spines or warts, frequently fused into short crests or even long wings; suprahilar spot inamyloid; hymenial cystidia and pileocystidia absent.Pileus (Fig. 3) small to medium sized, 10–50 mm diam., young hemispherical or convex, becoming plane and depressed at the centre; margin first even, when old distinctly tuberculate-striate up to 10 mm from the margin, often radially cracking; cuticle hardly peeling, radially disrupted into small patches, pruinose when young, later dry, smooth, matt and pruinose in the centre, colour near the margin when young varnish red (9C8), later red to coral red (9B6-7); near the centre deep red, blood red, dark red (10C7-8), raspberry red (10D7), strawberry red (10D8) or purple brown (10E-F8). Lamellae: 3–5 mm deep, thin, moderately dense, 6–8 at 1 cm near the pileus margin, adnexed, white, slightly anastomosing at the base; lamellulae absent, occasionally forked near the stipe; edges concolorous, entire but pruinose under lens. Stipe: 10–30 × 3–7 mm, usually narrowed towards the base, sometimes cylindrical, surface smooth, white and mainly with a distinct pastel red to red flush, occasionally completely white or sometimes also almost completely red, interior stuffed. Context: white, fragile, unchanging when damaged, reaction with guaiac after 5 s negative on both stipe and lamellae surfaces, reaction to FeSO4 and sulfovanillin negative; taste mild; odour inconspicuous. Spore print: not observed.Figure 3Basidiomata of Russula bellissima. (A) FH12-127 (Holotype). (B) FH12-158. Scale bar = 1 cm. Photos by Felix Hampe.Full size imageSpores (Figs. 4, 5) (6.9–)7.3–7.8–8.3(–8.9) × (6.1–)6.8–7.2–7.6(–8.4) µm, subglobose to broadly ellipsoid, Q = 1.01–1.1–1.2(–1.29); ornamentation of moderately distant [(4–)5–6(–7) in a 3 µm diam. circle] amyloid spines or warts, (1.1–)1.2–1.4–1.6(–1.7) µm high, fused or connected by fine line connections into often long crests or wings, [(0–)1–3(–4) fusions and the same number of line connections in a 3 µm diam. circle], crests and wings frequently branched and occasionally form closed loops, isolated elements dispersed, edge of crests and wings irregularly wavy; suprahilar spot moderately large, inamyloid. Basidia: (30.5–)34.5–44.1–53.5(–65.0) × (10.5–)11.5–12.6–14.0(–16.0) µm, broadly clavate or obpyriform, 4-spored; basidiola cylindrical, ellipsoid or broadly clavate, ca. 5–10 µm wide. Hymenial cystidia on lamellae sides: absent. Lamellae edges: covered by densely arranged or fasciculate marginal cells. Marginal cells: (27.0–)38.5–46.4–54.5(–61.0) × (5.0–)5.5–6.7–7.5(–9.0) µm; subulate or narrowly lageniform, apically attenuated and constricted to ca. 1–2 µm, sometimes slightly moniliform or flexuous. Pileipellis: (Fig. 6) orthochromatic in Cresyl Blue, gradually passing to the underlying context, 200–300 µm deep; suprapellis 60–130 µm deep, composed of erect or ascending hyphal terminations forming a dry trichoderm, well delimited from 140 to 210 µm deep subpellis composed of horizontally oriented, strongly gelatinized narrow hyphae. Subpellis not well delimited from the underlying context, elongate hyphae gradually changing to sphaerocytes. Acid- resistant incrustations: absent. Hyphal terminations near the pileus margin: composed of long apically attenuated terminal cell and a chain of 1–4 ovoid to barrel shaped, short unbranched cells with one distinctly longer apical cell; constricted on septa, usually not flexuous, oriented towards the pileus surface, usually thin-walled, sometimes slightly thick-walled (up to 1 µm thick); terminal cells mainly subulate or lageniform, apically attenuated and acute, measuring (19–)27.5–38.3–49.0(–66.5) × (3.3–)4.5–5.8–7.0(–9.0) µm, rarely with a forked apex, mixed with dispersed, cylindrical or ellipsoid, distinctly shorter, obtuse terminal cells measuring (7.5–)11.5–17.8–29.5(–42.5) × (3.0–)4.0–4.5–5.0 µm; subterminal cells measuring (4.5–)5.5–8.3–11.5(–16.0) × 4.5–5.3–6.0(–7.0) µm. Hyphal terminations near the pileus centre: similar in shape and also with a mixture of long acute and short obtuse terminal cells, acute ones measuring (12.0–)22.0–35.2–48.5(–79.0) × (2.5–)3.5–4.9–6.5(–8.0) µm, obtuse ones more frequent, measuring (6.5–)8.5–12.0–15.5(–22.0) × (3.5–)4.0–4.9–6.0(–7.5) µm. Primordial hyphae or pileocystidia: absent. Cystidioid hyphae and oleipherous hyphae not observed.Figure 4Hymenial elements of Russula bellissima (holotype, FH 12-127). (A) Basidia and basidiolae. (B) Marginal cells. (C) Spores as seen in Melzer’s reagent. Scale bar = 10 µm, but only 5 µm for spores.Full size imageFigure 5Scanning electron microscope photo of spore ornamentation. Russula bellissima (holotype, FH 12-127). Scale bar = 2 μm.Full size imageFigure 6Elements of the pileipellis of Russula bellissima (holotype, FH 12-127). (A) Hyphal terminations near the pileus margin. (B) Hyphal terminations near the pileus centre. Scale bar = 10 μm.Full size imageAdditional material studied THAILAND, Chiang Mai Province, Mae On District, about 3 km from Tharnthong lodges, 18° 51′ 55″ N 99° 17′ 23″ E, alt. 725 m, Dipterocarpaceae dominated forest with the presence of some Castanopsis trees, in small groups on loamy soil, 17 July 2012, F. Hampe (GENT FH 12-158, duplicate: MFLU12-0648).Note Russula bellissima is a small species with a bright red pileus and pink colour on the stipe. This colour is distinctive and resembles North American R. mariae, Indian R. intervenosa and Asian R. bella. It is very unlikely that the distribution of any European or North American species is overlapping with the Thai species. However, little is known about the distributional ranges and the ecological niches of other Asian Russula species. Therefore discussing the morphological distinguishing characters between Asian species and R. bellissima is more relevant. Russula bellissima is not closely related to R. bella and it differs from this species by larger spores with a more prominent spore ornamentation, absence of hymenial cystidia on lamellae sides, and subterminally short, ellipsoid cells in the suprapellis arranged in unbranched chains of up to four7. The Thai species resembles and is closely related to the Indian R. intervenosa, but it has a more prominent spore ornamentation, hymenial cystidia (on lamellae sides) are absent, and hyphal terminations in the pileipellis are wider22.
Russula luteonana M. Pobkwamsuk & K. Wisitrassameewong sp. nov.
Mycobank: MB 840550Holotype: THAILAND, Amnat Charoen province, Hua Taphan district, Junction near Watbochaneng , dry dipterocarp forest, alt. 145 m, 15° 41′ 28″ N 104° 31′ 41″ E, 13 July 2016, Thitiya Boonpratuang, Rattaket Choeyklin, Prapapan Sawhasan, Maneerat Pobkwamsuk, Pattrachai Juthamas, Nattawut Wiriyathanawudhiwong, Patcharee Patangwesa (BBH41120).Etymology ‘Luteolus’ = yellow colour, ‘Nanus’ = small. Refer to pileus color and size of the species.Diagnosis Pileus medium-sized, dry, usually yellow, spores with subreticulate amyloid ornamentation and inamyloid suprahilar spot, hymenial cystidia on lamellae sides large, lamellae edges with combination of subulate, clavate and pyriform marginal cells.Pileus (Fig. 7) medium-sized, 28‒53 mm diam., plano-convex with depressed centre, infundibuliform when mature; margin striated and radially cracking in dry condition; cuticle dry, peeling to almost ½ of radius, smooth to minutely wrinkled, dull in dry condition, color very variable, some collections pale cream and with darker pale brownish-yellow centre, other yellow brownish and with darker orange-brown centre, sometimes also bright red-brown and with discolored centre, always with rusty-brown spots especially when near the centre. Lamellae: 3‒5 mm deep, moderately distant, intervenose, forking near the stipe, white to cream, edges even, concolorous. Stipe: 26‒40 × 6‒9 mm, cylindrical or narrowed at the base, surface dry, longitudinally wrinkled, white, turning brown when bruised. Context: 2‒4 mm in at the half pileus radius, soft, solid, becoming partially hollow when mature, white, unchanging when cut. Taste mild; odour rather strong, fishy. Spore print: not observed.Figure 7Basidiomata of Russula luteonana. (A) BBH41120 (Holotype). (B) BBH41121. (C) BBH41122. (D) BBH42510. Scale bar = 1 cm. Photos by Thitiya Boonpratuang.Full size imageSpores (Figs. 8, 9) (7.4‒)8.1‒8.6‒9(‒10.1) × (6.1‒)7.4‒7.5‒7.9(‒9.1) μm, subglobose to broadly ellipsoid, Q = (1.03‒)1.09‒1.15‒1.20(‒1.30), ornamentation of moderately distant, obtuse, (0.7‒)1.1‒1.3‒1.5(‒1.9) μm high spines, connected by abundant line connections [(0‒)3‒6(‒8) in in a 3 µm diam. circle], branched, forming an incomplete reticulum, crest irregularly wavy and occasionally fused [(0‒)1‒2(‒5) fusions in the circle], isolated elements rare; suprahilar spot inamyloid. Basidia: (29‒)34.5‒39.1‒44(‒51.5) × (10‒)12‒13.2‒14.5(‒16.5) μm, clavate, 4-spored, rarely 2-spored, basidiola subcylindrical to subclavate, (25.5‒)30‒35.4‒41(‒47) × (9‒)11‒12.2‒14 (‒16) μm. Hymenial cystidia on lamellae sides: usually protruding over other elements of hymenium, widely dispersed ( More

Lincoln, G. A. Teeth, horns and antlers: the weapons of sex. In The Differences between the Sexes (eds R. V. Short & E. Balaban) 131–158 (Cambridge Univ. Press, 1994).Lundrigan, B. Morphology of horns and fighting behavior in the family bovidae. J. Mammal. 77, 462–475 (1996).Article 

Google Scholar 
Bro-Jorgensen, J. The intensity of sexual selection predicts weapon size in male bovids. Evolution 61, 1316–1326 (2007).Article 

Google Scholar 
Plard, F., Bonenfant, C. & Gaillard, J. M. Revisiting the allometry of antlers among deer species: male-male sexual competition as a driver. Oikos 120, 601–606 (2011).Article 

Google Scholar 
Okada, K. & Miyatake, T. Sexual dimorphism in mandibles and male aggressive behavior in the presence and absence of females in the beetle Librodor japonicus (Coleoptera: Nitidulidae). Ann. Entomol. Soc. Am. 97, 1342–1346 (2004).Article 

Google Scholar 
Emlen, D. J., Marangelo, J., Ball, B. & Cunningham, C. W. Diversity in the weapons of sexual selection: Horn evolution in the beetle genus Onthophagus (Coleoptera: Scarabaeidae). Evolution 59, 1060–1084 (2005).CAS 
Article 

Google Scholar 
Pomfret, J. C. & Knell, R. J. Sexual selection and horn allometry in the dung beetle Euoniticellus intermedius. Anim. Behav. 71, 567–576 (2006).Article 

Google Scholar 
McCullough, E. L., Weingarden, P. R. & Emlen, D. J. Costs of elaborate weapons in a rhinoceros beetle: how difficult is it to fly with a big horn?. Behav. Ecol. 23, 1042–1048 (2012).Article 

Google Scholar 
David, P., Bjorksten, T., Fowler, K. & Pomiankowski, A. Condition-dependent signalling of genetic variation in stalk-eyes flies. Nature 406, 186–188 (2000).ADS 
CAS 
Article 

Google Scholar 
Baker, R. H. & Wilkinson, G. S. Phylogenetic analysis of sexual dimorphism and eye-span allometry in stalk-eyed flies (Diopsidae). Evolution 55, 1373–1385 (2001).CAS 
Article 

Google Scholar 
Stankowich, T. Armed and dangerous: predicting the presence and function of defensive weaponry in mammals. Adapt. Behav. 20, 32–43 (2012).Article 

Google Scholar 
Hashimoto, K. & Hayashi, F. Structure and function of the large pronotal horn of the sand-living anthicid beetle Mecynotarsus tenuipes. Entomol. Sci. 15, 274–279 (2012).Article 

Google Scholar 
Hayashi, M. & Ohba, S. Y. Mouth morphology of the diving beetle Hyphydrus japonicus (Dytiscidae: Hydroporinae) is specialized for predation on seed shrimps. Biol. J. Linn. Soc. 125, 315–320 (2018).Article 

Google Scholar 
Stocker, R. F. The organization of the chemosensory system in Drosophila melanogaster: a review. Cell Tissue Res. 275, 3–26 (1994).CAS 
Article 

Google Scholar 
Dweck, H. K. M. Antennal sensory receptors of Pteromalus puparum female (Hymenoptera: Pteromalidae), a gregarious pupal endoparasitoid of Pieris rapae. Micron 40, 769–774 (2009).Article 

Google Scholar 
Crespo, J. G. A review of chemosensation and related behavior in aquatic insects. J. Insect Sci. 11, 1–39 (2011).Article 

Google Scholar 
Stoffolano, J. G. Jr., Rice, M. & Murphy, W. L. The importance of antennal mechanosensilla of Sepedon fuscipennis (Diptera: Sciomyzidae). Can. Entomol. 145, 265–272 (2013).Article 

Google Scholar 
Gabel, B. et al. Floral volatiles of Tanacetum vulgare L. attractive to Lobesia botrana Den. et Schiff. females. J. Chem. Ecol. 18, 693–701 (1992).CAS 
Article 

Google Scholar 
Fox, H. Barbels and barbel-like tentacular structures in sub-mammalian vertebrates: a review. Hydrobiologia 403, 153–193 (1999).Article 

Google Scholar 
Plepys, D., Ibarra, F., Francke, W. & Lofstedt, C. Odour-mediated nectar foraging in the silver Y moth, Autographa gamma (Lepidoptera: Noctuidae): behavioural and electrophysiological responses to floral volatiles. Oikos 99, 75–82 (2002).CAS 
Article 

Google Scholar 
Stankowich, T. & Caro, T. Evolution of weaponry in female bovids. Proc. R. Soc. Lond. Ser. B Biol. Sci. 276, 4329–4334 (2009).Bergmann, P. J. & Berk, C. P. The Evolution of Positive Allometry of Weaponry in Horned Lizards (Phrynosoma). Evol. Biol. 39, 311–323 (2012).Article 

Google Scholar 
Damman, H. The osmaterial glands of the swallowtail butterfly Eurytide marcellus as a defense against natural enemies. Ecol. Entomol. 11, 261–265 (1986).Article 

Google Scholar 
Berenbaum, M. R., Moreno, B. & Green, E. Soldier bug predation on swallowtail caterpillars (Lepidoptera, Papilionidae): circumvention of defensive chemistry. J. Insect Behav. 5, 547–553 (1992).Article 

Google Scholar 
Juma, G. et al. Distribution of chemo- and mechanoreceptors on the antennae and maxillae of Busseola fusca larvae. Entomol. Exp. Appl. 128, 93–98 (2008).Article 

Google Scholar 
Liu, Z., Hua, B.-Z. & Liu, L. Ultrastructure of the sensilla on larval antennae and mouthparts in the peach fruit moth, Carposina sasakii Matsumura (Lepidoptera: Carposinidae). Micron 42, 478–483 (2011).Article 

Google Scholar 
Kandori, I., Tsuchihara, K., Suzuki, T. A., Yokoi, T. & Papaj, D. R. Long frontal projections help Battus philenor (Lepidoptera: Papilionidae) larvae find host plants. PLoS ONE 10, e0131596 (2015).Article 

Google Scholar 
Greeney, H. F., Dyer, L. A. & Smilanich, A. M. Feeding by lepidopteran larvae is dangerous: A review of caterpillars’ chemical, physiological, morphological, and behavioral defenses against natural enemies. ISJ Invert. Surviv. J. 9, 7–34 (2012).
Google Scholar 
Sugiura, S. Predators as drivers of insect defenses. Entomol. Sci. 23, 316–337 (2020).Article 

Google Scholar 
Martin, W. R. & Nordlund, D. A. Ovipositional behavior of the parasitoid Palexorista laxa (Diptera, Tachinidae) on Heliothis zea (Lepidoptera, Noctuidae) larvae. J. Entomol. Sci. 24, 460–464 (1989).Article 

Google Scholar 
Constantino, L. M. Notes on Haetera from Colombia, with description of the immature stages of Haetera piera (Lepidoptera:Nymphalidae: Satyrinae). Trop. Lepid. 4(1), 13–15 (1993).
Google Scholar 
Devries, P. J., Kitching, I. J. & Vanewright, R. I. The systematic position of Antirrhea and Caerois, with comments on the classification of the Nymphalidae (Lepidoptera). Syst. Entomol. 10, 11–32. https://doi.org/10.1111/j.1365-3113.1985.tb00561.x (1985).Article 

Google Scholar 
Dias, F. M. S., Casagrande, M. M. & Mielke, O. H. H. Biology and external morphology of immature stages of Memphis appias (Hubner) (Lepidoptera: Nymphalidae: Charaxinae). Zootaxa, 21–32 (2010).Dias, F. M. S., Casagrande, M. M. & Mielke, O. H. H. Biology and external morphology of the immature stages of the butterfly Callicore pygas eucale, with comments on the taxonomy of the genus Callicore (Nymphalidae: Biblidinae). J. Insect Sci. 14, doi:https://doi.org/10.1093/jis/14.1.91 (2014).Dias, F. M. S., Casagrande, M. M. & Mielke, O. H. H. Immature stages of the turquoise-banded shoemaker Archaeoprepona amphimachus pseudomeander (Fruhstorfer, 1906) and a comparative review of the Preponini (Lepidoptera: Nymphalidae). Aust. Entomol. 58, 451–462. https://doi.org/10.1111/aen.12339 (2019).Article 

Google Scholar 
Dias, F. M. S., de Oliveira-Neto, J. F., Casagrande, M. M. & Mielke, O. H. H. External morphology of immature stages of Zaretis strigosus (Gmelin) and Siderone galanthis catarina Dottax and Pierre comb. nov., with taxonomic notes on Siderone (Lepidoptera: Nymphalidae: Charaxinae). Rev. Bras. Entomol. 59, 307–319, doi:https://doi.org/10.1016/j.rbe.2015.07.007 (2015).Dias, F. M. S. et al. An integrative approach elucidates the systematics of Sea Hayward and Cybdelis Boisduval (Lepidoptera: Nymphalidae: Biblidinae). Syst. Entomol. 44, 226–250. https://doi.org/10.1111/syen.12327 (2019).Article 

Google Scholar 
Freitas, A. V. L., Barbosa, E. P. & Marin, M. A. Immature Stages and Natural History of the Neotropical Satyrine Pareuptychia ocirrhoe Interjecta (Nymphalidae: Euptychiina). J. Lepid. Soc. 70, 271–276. https://doi.org/10.18473/lepi.70i4.a4 (2016).Article 

Google Scholar 
Freitas, A. V. L., Kaminski, L. A., Mielke, O. H. H., Barbosa, E. P. & Silva-Brandao, K. L. A new species of Yphthimoides (Lepidoptera: Nymphalidae: Satyrinae) from the southern Atlantic forest region. Zootaxa, 31–44 (2012).Furtado, E. & Campos-Neto, F. C. Caligopsis seleucida (Hewitson) and its immature stages (Lepidoptera, Nymphalidae, Brassolinae). Rev. Bras. Zool. 21(3), 593–597 (2004).Article 

Google Scholar 
Greeney, H. F. et al. The early stages and natural history of Antirrhea adoptiva porphyrosticta (Watkins, 1928) in eastern Ecuador (Lepidoptera: Nymphalidae: Morphinae). J. Insect Sci. 9 (2009).Greeney, H. F. et al. Early stages and natural history of Perisama oppelii (Nymphalidae, Lepidoptera) in eastern Ecuador. Kempffiana 6(1), 16–30 (2010).
Google Scholar 
Greeney, H. F., Dyer, L. A. & Pyrcz, T. W. First description of the early stage biology of the genus Mygona: The natural history of the satyrine butterfly, Mygona irmina in eastern Ecuador. J. Insect Sci. 11, doi:https://doi.org/10.1673/031.011.0105 (2011).Greeney, H. F., Pyrcz, T. W., DeVries, P. J. & Dyer, L. A. The early stages of Pedaliodes poesia (Hewitson, 1862) in eastern Ecuador (Lepidoptera: Satyrinae: Pronophilina). J. Insect Sci. 9 (2009).Greeney, H. F., Whitfield, J. B., Stireman, J. O., Penz, C. M. & Dyer, L. A. Natural history of Eryphanis greeneyi (Lepidoptera: Nymphalidae) and its enemies, with a description of a new species of Braconid parasitoid and notes on its Tachinid parasitoid. Ann. Entomol. Soc. Am. 104, 1078–1090. https://doi.org/10.1603/an10064 (2011).Article 

Google Scholar 
Kaminski, L. A. & Freitas, A. V. L. Immature stages of the butterfly Magneuptychia libye (L.) (Lepidoptera : Nymphalidae, Satyrinae). Neotrop. Entomol. 37, 169–172, doi:https://doi.org/10.1590/s1519-566×2008000200010 (2008).Lambkin, T. & Kendall, R. The status of Yoma algina (boisduval, 1832) & Y. sabina (cramer, 1780) (Lepidoptera: Nymphalidae: Nymphalinae) in Australia. Aust. Entomol. 43 (4), 211–234 (2016).Leite, L. A. R., Casagrande, M. M., Mielke, O. H. H. & Freitas, A. V. L. Immature stages of the Neotropical butterfly, Dynamine agacles agacles. J. Insect Sci. 12 (2012).Leite, L. A. R., Dias, F. M. S., Carneiro, E., Casagrande, M. M. & Mielke, O. H. H. Immature stages of the Neotropical cracker butterfly, Hamadryas epinome. J. Insect Sci. 12 (2012).Murillo, L. R. & Nishida, K. Life history of Manataria maculata (Lepidoptera : Satyrinae) from Costa Rica. Rev. Biol. Trop. 51, 463–469 (2003).PubMed 

Google Scholar 
Nakahara, S., Janzen, D. H., Hallwachs, W. & Espeland, M. Description of a new genus for Euptychia hilara (C. Felder & R. Felder, 1867) (Lepidoptera: Nymphalidae: Satyrinae). Zootaxa 4012, 525-541, doi:https://doi.org/10.11646/zootaxa.4012.3.7 (2015).Penz, C. M., Freitas, A. V. L., Kaminski, L. A., Casagrande, M. M. & Devries, P. J. Adult and early-stage characters of Brassolini contain conflicting phylogenetic signal (Lepidoptera, Nymphalidae). Syst. Entomol. 38, 316–333. https://doi.org/10.1111/syen.12000 (2013).Article 

Google Scholar 
Pyrcz, T. W. et al. Uncovered diversity of a predominantly Andean butterfly clade in the Brazilian Atlantic forest: a revision of the genus Praepedaliodes Forster (Lepidoptera: Nymphalidae, Satyrinae, Satyrini). Neotrop. Entomol. 47, 211–255. https://doi.org/10.1007/s13744-017-0543-x (2018).CAS 
Article 
PubMed 

Google Scholar 
Shirai, L. T. et al. Natural history of Selenophanes cassiope guarany (Lepidoptera: Nymphalidae: Brassolini): an integrative approach, from molecules to ecology. Ann. Entomol. Soc. Am. 110, 145–159. https://doi.org/10.1093/aesa/saw068 (2017).Article 

Google Scholar 
Silva, P. L. et al. Immature Stages of the Brazilian Crescent Butterfly Ortilia liriope (Cramer) (Lepidoptera: Nymphalidae). Neotrop. Entomol. 40, 322–327. https://doi.org/10.1590/s1519-566×2011000300006 (2011).CAS 
Article 
PubMed 

Google Scholar 
Song-yun, L. Immature stages of Faunis aerope (Leech, 1890) (Lepidoptera, Nymphalidae). Atalanta 42, 221–222 (2011).
Google Scholar 
Steiner, H. Life history of Melanocyma faunula in Malaysia (Lepidoptera: Nymphalidae: Morphinae). Trop. Lepid. Res. 16, 23–26 (2005).
Google Scholar 
Velez, P. D., Montoya, H. H. V. & Wolff, M. Immature stages and natural history of the Andean butterfly Altinote ozomene (Nymphalidae: Heliconiinae: Acraeini). Zoologia 28, 593–602. https://doi.org/10.1590/s1984-46702011000500007 (2011).Article 

Google Scholar 
Wahlberg, N. et al. Nymphalid butterflies diversify following near demise at the Cretaceous/Tertiary boundary. Proc. R. Soc. Lond. Ser. B Biol. Sci. 276, 4295–4302, doi:https://doi.org/10.1098/rspb.2009.1303 (2009).Willmott, K. R., Elias, M. & Sourakov, A. Two possible caterpillar mimicry complexes in neotropical Danaine butterflies (Lepidoptera: Nymphalidae). Ann. Entomol. Soc. Am. 104, 1108–1118. https://doi.org/10.1603/an10086 (2011).Article 

Google Scholar 
Willmott, K. R. & Freitas, A. V. L. Higher-level phylogeny of the Ithomiinae (Lepidoptera : Nymphalidae): classification, patterns of larval hostplant colonization and diversification. Cladistics 22, 297–368. https://doi.org/10.1111/j.1096-0031.2006.00108.x (2006).Article 
PubMed 

Google Scholar 
Zacca, T. et al. Revision of Godartiana Forster (Lepidoptera: Nymphalidae), with the description of a new species from northeastern Brazil. Aust. Entomol. 56, 169–190. https://doi.org/10.1111/aen.12223 (2017).Article 

Google Scholar 
Bossart, J.L., Fetzner Jr., J.F. & Rawlins, J.E. Ghana Butterfly Biodiversity Project website. https://www.invertebratezoology.org/GhanaBfly/default.asp (2007).Butterflies and Moths of North America project. Butterflies and Moths of North America website. https://www.butterfliesandmoths.org/ (2021).Dauphin, D. & Dauphin, J. The Rio Grande Valley’s Nature Site website. http://www.thedauphins.net (2021).Eeles, P. UK Butterflies website. https://www.ukbutterflies.co.uk/index.php. (2021).Florida Museum of Natural History. Florida Museum website. https://www.floridamuseum.ufl.edu/ (2021).Khew, S. K. et al. Butterflies of Singapore website. https://butterflycircle.blogspot.com/ (2021).Kunte, K., Sondhi, S. & Roy, P. Butterflies of India, v. 3.24. Indian Foundation for Butterflies website. https://www.ifoundbutterflies.org (2021).Miller, S. & Morrison, C. Parasitoid-Caterpillar-Plant Interactions in the Americas website. https://caterpillars.myspecies.info/ (2021).National Biodiversity Network Trust. iNaturalistUK website. https://uk.inaturalist.org/ (2021).Nature Picture Library Limited. Nature Picture Library website. https://www.naturepl.com/blog/ (2021).Project Noah Team. Project Noah website. https://www.projectnoah.org/ (2021).Shiraiwa, K. Pteron World. The encyclopedia website of the butterflies. https://www.pteron-world.com/index.html (2021).Wagner, W. Lepidoptera and Their Ecology website. http://www.pyrgus.de/ (2021).Wahlberg, N. & Peña, C. Nymphalidae.net. website. http://www.nymphalidae.net/ (2021).Wikimedia Foundation, Inc. Wikimedia Commons website. https://commons.wikimedia.org/ (2021).Matsuura, M. Social Wasps of Japan in Color. (in Japanese) (Hokkaido university press 2015).IBM SPSS. SPSS Base 25.0 User’s Guide. (SPSS Inc., 2017). More

Phyla Bdellovibrionota, Fusobacteriota, and Myxococcota were present in the green microbial mat but in negligible quantities in the brown mat. The unique phyla detected in the brown mat, but not in the green microbial mat, included Caldatribacteriota, Thermodesulfobacteriota, Dictyoglomota, Elusimicrobiota, Thermotogota, Candidatus Calescamantes, Fervidibacteria, Hydrothermae, GAL15 and TA06. The Candidatus Caldatribacterium (phyla Caldatribacteriota), earlier named OP9 was also detected in this work. Using single-cell and metagenome sequencing, data elucidated that Ca. Caldatribacterium conducts anaerobic sugar fermentation and exhibited diverse glycosyl hydrolases, including endoglucanase37.Cyanobacteria and Chloroflexota were the main identified phyla in the green microbial mat. Because the hot spring is almost stagnant, undisturbed, and the water surface temperature ( More

Variability of N2O concentrations and fluxesAll sampled streams and rivers were supersaturated on all dates (117.9–242.5%, n = 342 samples from 114 site visits) in N2O with respect to the atmosphere. Dissolved N2O concentrations fluctuated between 10.2 and 18.9 nmol L−1 with an average of 12.4 ± 1.7 nmol L−1, which is one-third of the global average3 (37.5 nmol L−1; Supplementary Table 3). Significantly higher N2O concentrations were observed in spring (P  More

Theory of changeAt the heart of the of CBRLM’s theory of change is the assumption that improvements in the ecological sub-system provide a sustainable resource base for increased livestock production and marketing34. The ecological sub-system, however, depends on a functioning economic sub-system because herd owners must be able to destock quickly in response to adverse ecological circumstances. The theory holds that the most important constraint on the economic sub-system is unproductive herds and low-quality cattle because farmers are unwilling to sell their cattle when they command low market prices. Therefore, improvements in rangeland grazing management need to be complemented by improvements in information and access to livestock markets, herd structures, and animal husbandry practices.Crucially, changes to the ecological, economic, and livestock sub-systems rely on effective community governance and collective-action capacity in CBRLM communities. This is because rangeland grazing management practices can be easily undermined by non-participating herd owners inside or outside the GA. The theory therefore calls for investments at multiple levels of the social-ecological system to ensure that improvements in certain program areas are not undermined by failures in others34. The CBRLM implementers believed that previous rangeland development programs were undermined by a failure to account for the linkages among sub-systems, which motivated them to design a more holistic intervention34.Intervention componentsCBRLM was a multi-faceted package of administrative, educational, financial, and technical support. Implementation of the package was designed as an experimental treatment to assist in project assessment. To select study areas for evaluation, GOPA identified 38 RIAs with sufficiently low density of people, livestock, and bush cover to enable the implementation of new group-grazing plans, one of the core treatment components. The evaluation team randomly assigned 19 RIAs to treatment and 19 RIAs to control (see Randomization for details). GOPA implemented CBRLM in up to seven GAs within each treatment RIA.MobilizationGOPA conducted pre-mobilization meetings with TAs and other stakeholders in the second half of 2010 to identify GA communities most likely to participate in CBRLM34. Early mobilization efforts focused on soliciting community buy-in for the cornerstone principles of CBRLM, including community-planned grazing, combined herding of cattle, and efficient livestock management. There is also substantial evidence from qualitative surveys that some community members were motivated to participate in the CBRLM by prospects for water infrastructure development by GOPA34.While almost 100 GAs were initially mobilized for the project, by 2014 GOPA was targeting resources and support towards 58 GAs based on community receptivity and the discretion of CBRLM management. In each GA, GOPA worked principally with households owning 10 or more cattle, although other community members benefitted from participation in a “Small Stock Pass-on Scheme” and a variety of training activities, which are described below.Rangeland grazing managementThe core aim of CBRLM was to shift how communities approached livestock grazing, forage conservation, and risk management by encouraging two key practices: planned grazing and combined herding. Planned grazing entails rotating a community’s cattle to a new pasture on a regular basis in accordance with a written plan. The goal was to preserve grass for the dry season and allow grazed pastures more time to recover. Combined herding entails grouping many owners’ cattle into one large herd and herding them in a tight bunch. This practice is meant to concentrate animal impact on rangeland, minimize cattle losses, and increase the likelihood that cows are exposed to bulls, thus increasing the pregnancy and calving rates of the entire herd. The scientific and practical rationale behind these practices is reviewed in Supplementary Note 2.GOPA staff developed grazing plans with each participating community and taught them planned grazing and combined herding via field-based training sessions. These followed a “training of trainers” approach in which GOPA recruited field facilitators from each community, taught them the principles of CBRLM, and tasked them with training their fellow participating pastoralists.Livestock managementGOPA taught participants some best practices in animal husbandry, including structuring herds to maximize productivity (by increasing the proportion of bulls and reducing the proportion of oxen and cattle over the age of 10 years), providing vaccinations and supplements, and deworming34. Additionally, to support the introduction of more bulls into herds, the project implemented a “bull scheme” in which participating communities were given the opportunity to collectively buy certified breeding bulls at a subsidized price. Communities were meant to repay the cost of the bulls either with cash or in-kind trades of goats. Goats collected in this repayment process fed into the small stock pass-on scheme under which participating community members nominated households to receive goats from GOPA. GOPA requested that communities nominate households that owned few or no livestock and were led by youth and/or women. When GOPA received goats as payment for loaned bulls, they would pass them on to nominated households. The recipients were then expected to pass on the offspring of the goats they received to other disadvantaged households.Cattle marketingCBRLM also sought to increase participants’ marketing of cattle to generate revenue from livestock raising and encourage offtake of unproductive animals34. Community facilitators and project experts provided participating herd owners with information about market opportunities and ideal herd composition, and encouraged flexible offtake in response to forage shortages. In 2013, GOPA invested in the development of regional livestock cooperatives that held local auctions and helped farmers transport their animals to markets. Finally, GOPA invested in identifying international export opportunities for CBRLM farmers to Zimbabwe and Angola, although these were generally not successful31.Community developmentThe project sought to institutionalize community-level governance to organize and enforce collective activities like planned grazing, water point maintenance, and financing of livestock inputs. The central management unit of each GA was a new Grazing Area Committee consisting of five to 10 elected community members. The project encouraged participating communities to collectively cover operational expenses in their GA through a GA fund managed by the committee. Among these expenses were the payments to herders, costs of diesel for water pumps and maintenance of water infrastructure, financing collective livestock vaccination campaigns, and any other collective expenses that would support operation of the GA. CBRLM supported every GA fund with a 1:1 matched subsidy. The matched subsidy was limited by a ceiling amount determined by the estimated number of cattle in a GA. GOPA also instructed committees to maintain “GA record books” to track grazing plans, record meeting minutes, and keep logs of community members’ participation and financial contributions.Water infrastructureGOPA upgraded water infrastructure at a total of 84 sites throughout the NCAs to facilitate planned grazing and combined herding. Water infrastructure improvement included minor upgrades like water tanks and drinking troughs, and larger investments such as the installation of diesel and solar pump systems, the drilling and installation of boreholes, and the construction of pipelines, deep wells, and a large earthen dam31.Intervention timelineThe timeline for major components of the research process and CBRLM roll-out is illustrated in Supplementary Fig. 1. The research team conducted the random assignments and the implementation team began community mobilization in early 2010. Formal enrollment in CBRLM began in early 2011. The program implementer conducted mobilization in two waves: they mobilized 11 of 19 RIAs in 2010 and the remaining 8 RIAs in 2011. The evaluation team conducted qualitative data collection to inform the design of social and cattle surveys prior to project end 2014; social surveys in 2014 and 2016; rangeland surveys in the wet and dry seasons of 2016; a cattle survey in 2016; and a household economic survey in 2017.Cumulative GA-level implementation is illustrated in Supplementary Fig. 2. The project implementer first formally reported enrollment and field visits in April 2011. The implementer achieved nearly full targeted enrollment (50 GAs) by November 11, although some grazing areas were added or subtracted thereafter. Mobilization exceeded enrollment because some grazing area communities chose not to participate in the program and some enrolled in the program and then dropped out. The program averaged between 25 and 50 field visits per month over the project period. A field visit consisted of a week-long community meeting about grazing-plan development and implementation, animal husbandry and budget training, and marketing opportunities.RandomizationThe unit of randomization is the RIA, an intervention zone with a locally recognized boundary. Each RIA falls under the jurisdiction of a single local governing body, known as a Traditional Authority (TA). As noted above, RIAs contain five to 15 GAs where a community of producers share water and forage resources. Grazing areas do not have legally defined boundaries. A herd owner’s ability to move among GAs is variable.GOPA mapped 41 RIAs prior to randomization. Three contiguous RIAs in the north-central region, composed of two treatment RIAs and one control RIA, were omitted from the study post-randomization because reexamination of baseline density of bushland vegetation deemed them unviable for CBRLM implementation. These are the three RIAs without sampled GAs in Fig. 1. The other 38 RIAs were randomly assigned to either receive the CBRLM treatment (19 RIAs) or serve as controls (19 RIAs).The randomization was stratified by TA to ensure that at least one RIA was assigned to the treatment in each TA. The research team then re-randomized the sample units until seven variables were balanced (a p value of 0.33 or higher for an omnibus f test of all seven variables) between treatment and control: (1) Presence of forest; (2) number of households; (3) number of cattle; (4) cattle density per unit area; (5) quality of water sources; (6) presence of community-based organizations (CBOs); and (7) overlap with complementary interventions (see Supplementary Table 1). For future researchers, we recommend re-randomizing a set number of times and choosing the re-randomization with the highest balance35. These variables and indicator variables for TA are included as covariates in all analyses.Sample selectionIn the original sampling strategy, the project implementer was asked to predict the GAs where they would implement the project if the RIA were assigned to treatment. However, there was limited overlap between the GAs that the implementer predicted and the GAs where CBRLM was ultimately implemented. Therefore, the evaluation team devised a revised sampling strategy in 2013, which proceeded in four steps:

1.

Map GAs in sampled RIAs: The evaluation team traveled to all 38 RIAs and worked with TAs and Namibian Agricultural Extension (AE) officers to map all the GAs in each RIA. The team mapped 171 GAs in control RIAs and 213 GAs in treatment RIAs.

2.

Collect pre-program data on GAs: The evaluation team collected information on pre-program characteristics of each GA from interviews with TAs and AE staff, the Namibian national census36, and the Namibian Atlas37. The latter has a geo-referenced database on climate, ecology, and livestock for the nation.

3.

Predict CBRLM enrollment for treatment GAs: The researchers used these data in a logistic regression to predict the probability that each GA would enroll in CBRLM and would adopt the CBRLM interventions based on pre-program characteristics. For example, the model found that GAs with more existing water infrastructure, strong social cohesion, and adequate cell phone service were more likely to be enrolled in the program. The variables used to predict CBRLM adoption were: (1) Presence of water installations (yes/no); (2) carrying capacity of the land (above/below the regional median); (3) community’s readiness to change (high/very high); (4) community’s social cohesion (high/very high); (5) spillover effects from neighbors; (6) quality of herders and herder turnover; (7) presence of members of the Himba ethnic group; (8) the TA’s readiness to change; (9) cell phone coverage; and (10) primary housing material (mud, clay, or brick).

4.

Generate sample of GAs in treatment and control RIAs: The evaluation team applied the statistical model (above) to all GAs in the sample and set a cut-off point to separate GAs that were likely to adopt the CBRLM program vs. those that were unlikely to do so. In treatment RIAs, the model predicted 52 GAs, of which 37 were formally enrolled in CBRLM and 15 were not. In control RIAs, 71 GAs met or exceeded the cutoff; they offer the best counter-factual estimate of which GAs would have enrolled in the program had their RIA received treatment.

Data collectionThe names, survey questions, and variable constructions for all outcomes included in the analysis are available at the AEA RCT Registry (ID number: AEARCTR-0002723). See Supplementary Methods for a list of definitions of variables depicted in Fig. 2 and 3.Social surveysSocial surveys were intended to assess the effect of CBRLM on community behaviors, community dynamics, knowledge, and attitudes. All data were collected using electronic tablets with the SurveyCTO software38.The primary unit of analysis for household respondents is the manager of the cattle kraal (holding pen). Researchers conducted surveys with kraal managers, rather than heads of households, for three reasons. First, many kraals contain cattle owned by multiple households, and decisions about grazing practices, cattle treatment, and participation in grazing groups are generally made at the kraal level. Second, many cattle-owning households do not directly oversee the day-to-day activities of their cattle (many live outside the GA), and so would be unable to answer questions about key outcomes, such as livestock management behaviors and community dynamics39. Finally, enrollment in CBRLM occurred at the kraal, rather than household, level.In 2014, the research team worked with local headmen and other community members to generate a complete census of kraals in every sampled Grazing Area (GA) that contained 10 or more cattle at the start of the program (an eligibility requirement for enrollment in CBRLM). The research team randomly sampled up to 11 community members for participation in the 2014 kraal manager survey. Surveys were conducted in the manager’s local language and lasted ~45 min. Alongside the 2014 survey, teams of two surveyors visited all grazing areas where at least one respondent reported participating in a community grazing group or community combined herd to corroborate reported behaviors through direct observation.To assess the persistence of CBRLM’s effects on behaviors, community dynamics, knowledge, and attitudes, the research team conducted a follow-up survey of kraal managers in 2016, two years after program end. The survey team randomly sampled two additional kraals in each grazing area to account for the possibility of attrition. The 2016 survey lasted approximately one hour on average, and included an expanded list of questions about governance, social conflict, and collective action as well as new survey modules on cattle marketing, cattle movement, and livestock management. In 2017, the research team randomly sampled three kraals in each grazing area to conduct direct observation audits of key rangeland grazing-management behaviors.To assess the effects of CBRLM on economic outcomes, the research team conducted a household-level survey in 2017, three years after program end. The survey instrument asked detailed questions on topics that could not be answered by kraal managers, such as household consumption, income, food security, and savings. To select households for this survey, during the 2016 survey the research team asked kraal managers to list all households that owned cattle in the manager’s kraal, then randomly selected one household from each kraal. Alongside the 2017 survey, the research team conducted an in-depth survey with the local headman of all 123 GAs in the sample. The headman survey focused on historical background about the grazing area, as well as the headman’s perceptions of rangeland and livestock issues.Cattle dataThe cattle component was intended to assess effects of CBRLM on cattle numbers, body condition, and productivity. The variables of key interest involved the average liveweight and body condition, calving rates, and average market value of cattle, as well as overall herd structures.The data collection protocols closely followed standards from livestock assessments elsewhere in Sub-Saharan Africa40. The research team randomly selected up to six kraals in each GA to participate in the cattle survey. The survey team mobilized selected herds during multiple community visits to ensure all herds were accounted for. Herd owners were compensated for the costs of rounding up animals and weighed cattle received anti-parasite treatment (“dipping”)41. A total of 19,875 cattle from 669 herds were weighed.The data-collection process for each herd proceeded in six steps. First, surveyors worked with herd managers to round up all cattle that regularly stayed in the selected cattle kraal. Once cattle had been brought to the designated location for data collection, they were passed through a mobile crush pen and scale. As each animal passed through the crush pen, a survey team member recorded the animal type (i.e., bull, ox, cow, calf) and used a SurveyCTO randomizer to calculate whether the animal was randomly selected for assessment. The random number generator was set to randomly select approximately 30 cattle from each herd for weighing. If the animal was selected, the survey team kept the animal on the scale and recorded its weight and body condition. A semi-subjective 1–5 scale, commonly used by livestock buyers in the NCAs (see Supplementary Fig. 3), was adjusted to a 0–4 scale used to determine formal market pricing. The team then placed the animal in a neck clamp and estimated the animal’s age by dentition (but extremely young calves were aged visually). Each animal was marked as it moved through the crush pen to ensure that it was assessed only once. In addition to assessing randomly selected animals, the survey team weighed and aged all bulls in the herd. The cattle survey yielded average cattle weight, age, and body condition for 19,875 animals across all treatment and control GAs, as well as estimates of calving rates, ratios of bulls to cows, and ratios of productive to unproductive animals.Rangeland dataThe rangeland ecology research was intended to assess treatment effects on vegetation and soil surface conditions. Full research details, including field technician training protocols, are available elsewhere42. The data collection approach followed methods commonly used in Africa43,44. Extended definitions of variables depicted in Fig. 3 and Table 2 are available in the “Supplementary Methods” section.The rationale for how the ecological variables presented in Fig. 3 translate into assessments of rangeland condition or health is based on forage and soil characteristics from a livestock production perspective25. The highest quality forages for cattle on rangelands are perennial grasses, since annual grasses are more ephemeral in terms of nutritive value and productivity. Herbaceous forbs often have the poorest forage quality for large grazers because of their low fiber content and risks of containing toxic chemicals. When rangelands are degraded by over-grazing, perennial grasses are reduced and replaced by annual grasses and forbs. This trend reflects animal diet selectivity that favors consumption of the perennial plants. Reversing such trends via management interventions can be difficult. The main option is to reduce grazing pressure and hope that perennial grasses can outcompete annuals and become reestablished over time. Another option is to implement a grazing rotation that allows perennial grasses to recover after a grazing period.Increases in annual grasses are documented to occur as one outcome of chronic overgrazing in Namibia45,46. In 2016, annual grasses were 5-times more abundant than perennial grasses in our study area. When over-grazing occurs, most plant material is harvested and less is available for the pool of organic matter (OM) for the topsoil. Less OM (e.g., plant litter) on the soil surface means that more soil is also exposed to wind and rain, accelerating erosion. The GAs in our research occur on various soil types and landscapes, some of which are more susceptible to erosion than others. Silty soils on slopes are vulnerable to erosion, for example, while sandy soils on level sites are less vulnerable25.On-the-ground sampling was conducted in all 123 selected GAs along an 800-km zone running West to East. Elevations ranged from 750 to 1700 masl (West) and 1050 to 1120 masl (East). Within each sampled GA, up to 12 1-ha (square) sampling sites were initially chosen using coordinates generated randomly from latitude and longitude coordinates in a satellite image of the GA47. About 17% of sites were later removed from the sample based on their close proximity to landscape disturbances or inaccessibility by field technicians. Overall, 972 sites were analyzed in the wet season and 885 in the dry season of 2016, two years after the implementation phase of CBRLM had ended.The geographic center-point for a sampling site was generated using a spatially constrained random distribution algorithm applied to the satellite image, and the field team navigated to the center-point coordinates using GPS technology. The team took photographs and recorded descriptive information including elevation, slope, aspect, other landscape features, vegetation type, dominant plant species, soil type, soil erosion, and degree of grazing or browsing pressure, and proximity to high impact areas such as trails, water points, and villages.At the center point, the survey team then established two perpendicular transects, each 100 m in length and crossing at the middle. The resulting four, 50-m transect lines ran according to each cardinal direction (N, S, E, W) as determined with a compass. Technicians then placed 1-m notched sampling sticks at randomized locations along each transect line and recorded what plants or other materials (i.e., stone, wood, leaf litter, animal dung, etc.) were located under or above the notches of the sampling sticks. These data points were tabulated to calculate percent cover for various categories of vegetation; there were n = 200 data points per site based on 40 stick placements and 5 notches per stick. This method enabled precise calculation of cover values for herbaceous (i.e., grass, forb) and diminutive woody plants (i.e., small shrubs, seedlings, saplings, etc.). Tree cover was estimated from point data collected via a small adjustment in the approach42. Herbaceous species were identified in wet seasons but not in dry seasons due to senescence during the latter.Quadrat sampling supplemented the notched stick approach. Random placements of a 1-m2 quadrat frame within the sampling site allowed for 20 estimates of a soil surface condition score ranging from 1 (poor) to 2 (moderate) or 3 (good)42. Poor was indicated by smooth soil surfaces, absence of litter, having poor infiltration and signs of erosion such as rills, pedestals, or terracettes; good was indicated by rough soil surfaces, abundant litter, seedlings evident, and lack of evidence of erosion. Herbaceous biomass was estimated in the quadrats and weighed to estimate herbaceous biomass.StatisticsIndex creationIndex construction for socioeconomic variables was composed of several steps48. For each response variable we first signed all component variables such that a higher sign is a positive outcome, i.e., in line with CBRLM’s intended impacts. Then we standardized each component by subtracting its control group mean and dividing by its control group standard deviation. We computed the mean of the standardized components of the index and standardized the sum once again by the control group sum’s mean and standard deviation. When the value of one component in an index was missing, we computed the index average from the remaining components. See Tables 3–6 for index components.Calculation of average treatment effectsThe estimate of interest is the Average Treatment Effect (ATE), or the average change in an outcome generated by assignment to CBRLM. We estimate the ATE using standard Ordinary Least Squares regression and control for variables used in stratification. Regressions for rangeland outcome variables include a unique set of controls, including rainfall over the project period, rainfall in the year of data collection, grazing area cattle density, grazing area ecological zones, and a remote-sensing estimate of pre-project biomass. The core model takes the form:$$hat{Y}=alpha +{beta }_{1}T+{{{{{boldsymbol{beta }}}}}}{{{{{bf{X}}}}}}$$
(1)
where T represents treatment assignment and X represents pre-treatment covariates used to test for balance during re-randomizations. The results capture the intention-to-treat (ITT) effect rather than the effect of treatment-on-treated (TOT). ITT is more appropriate than TOT in this context for two principal reasons. First, it is more relevant for policymakers – the effect of policies should account for imperfect compliance. Second, “uptake” is not well-defined, and certainly not a binary concept, for CBRLM since many communities and community members complied partially, complied with some but not all components, and complied for some but not all of the time.Standard errors and p valuesWe report two-tailed p values for all analyses. For each outcome, we show the two-tailed p value from a standard Ordinary Least Squares (OLS) regression with standard errors clustered at the level of the RIA, the unit of randomization49. We also calculate two-tailed p values using Randomization Inference (RI). To calculate RI p values, we re-run the randomization procedure (described above) 10,000 times and generate an Average Treatment Effect (ATE) under each hypothetical randomization. The p value is the percent of re-randomizations that generate a treatment effect that is either equal to, or larger in absolute value than, the true ATE.Multiple hypotheses correctionWe calculate q values to account for families of outcome indices with multiple hypotheses50. The q value represents the minimum false discovery rate at which the null hypothesis would be rejected for a given test. We pre-specified five families of indices:

1.

Behavioral outcomes (all in 2014): Grazing planning, Grazing-plan adherence, Herding practices, and Herder management.

2.

Behavioral outcomes (all in 2016): Grazing planning, Grazing-plan adherence, Herding practices, and Herder management.

3.

Primary material outcomes: Cattle herd value (2016), Herd productivity (2016), Household income (2017), Household expenditures (2017), Household livestock wealth (2017).

4.

Secondary material outcomes: Time use (2017), Resilience (2017), Female empowerment (2017), Diet (2017), and Herd structure (2016).

5.

Mechanisms: Collective Action (2014, 2016), Community Governance (2014, 2016), Community disputes (2014, 2016), Trust (2014), Self and community efficacy (2014, 2017), and Knowledge (2016).

Heterogeneous treatment effects analysisWe are interested in whether the effect of CBRLM was impacted by lower rainfall in some grazing areas during the project period. We evaluated heterogeneous treatment effects by rainfall in grazing areas using a variety of measures of rainfall, including aggregate rainfall during the project period and deviation in aggregate rainfall from the ten-year mean during the project period.For simplicity, Supplementary Tables 5 and 6 present the results of analysis of the interaction between treatment and a binary indicator of low rainfall. To construct this indicator, for each GA we first compute the absolute difference between mean rainfall during the project and mean rainfall during the 10 years prior (2000–2010). We divide the absolute difference by mean rainfall during the 10 years prior to produce a relative (%) difference. We then determine the median relative difference over all GAs. For each GA, we assign the value 1 to the low rainfall indicator if the relative difference for the GA is less than the median relative difference over all GAs; we assign 0 otherwise. The results are consistent when we use alternative rainfall measures.Spillovers analysisBecause CBRLM grazing areas were more likely to experience external incursions by cattle herds from outside the community, we test for spillovers. Specifically, we are interested in whether control grazing areas near treatment areas were affected by having a treatment grazing area nearby. We conducted the spillovers analysis only on control group grazing areas. For each control group grazing area, we measured the distance to the border of the nearest treatment grazing area. We created a binary measure taking the value 1 if the distance between the control group grazing area and nearest treatment group grazing area is below the median distance, and 0 otherwise. We find no evidence of spillover effects. The results are presented in Supplementary Table 7.Ethical considerationsApproval for this study was obtained from the Institutional Review Boards at Yale University (1103008148), Innovations for Poverty Action (253.11March-001), and Northwestern University (STU00205556-CR0001). The program was conceived, designed, and implemented by the Millennium Challenge Account compact between the Millennium Challenge Corporation and the Government of Namibia. The research team did not participate in program design or implementation. Communities and individual farmers were informed that they were free to withdraw from participation in evaluation activities at any time. The random assignment of the program was appropriate given the uncertainty around the program’s effect, and the Government of Namibia committed to implementing the program in control areas if the evaluation showed positive results.The research team took a number of steps to ensure the autonomy and well-being of study participants. First, we designed the survey and data collection protocols after considerable qualitative field work to ensure that questions about sensitive issues (e.g., cattle wealth, cattle losses, attitudes towards the Traditional Authority) were phrased appropriately and did not engender adverse emotional or social consequences. Second, all survey activities were reviewed and approved by the MCA compact, Regional Governors, and Traditional Authorities. Third, surveys were conducted with informed consent and in private to ensure that information remained private and respondents were as comfortable as possible during the survey. Finally, the research team disseminated findings on market prices and rangeland condition to communities and regional Agriculture Extension Officers.We received no negative reports about the community reception of the survey from surveyors during the evaluation. Two cows were injured during the cattle weighing exercise, and the owner was financially compensated in line with a compensation agreement made with all farmers prior to the cattle weighing exercise.Reporting summaryFurther information on research design is available in the Nature Research Reporting Summary linked to this article. More

Xin, F. et al. Large increases of paddy rice area, gross primary production, and grain production in Northeast China during 2000–2017. Sci. Total Environ. 711, 135–183. https://doi.org/10.1016/j.scitotenv.2019.135183 (2020).CAS 
Article 

Google Scholar 
Du, B. et al. Deep fertilizer placement improves rice growth and yield in zero tillage. Appl. Ecol. Environ. Res. 16, 8045–8054. https://doi.org/10.15666/aeer/1606_80458054 (2018).Article 

Google Scholar 
Ni, B., Liu, M., Lü, S., Xie, L. & Wang, Y. Environmentally friendly slow-release nitrogen fertilizer. J. Agric. Food Chem. 59, 10169–10175. https://doi.org/10.1021/jf202131z (2011).CAS 
Article 
PubMed 

Google Scholar 
Zhu, C. et al. Mechanized transplanting with side deep fertilization increases yield and nitrogen use efficiency of rice in Eastern China. Sci. Rep. 9, 5653. https://doi.org/10.1038/s41598-019-42039-7 (2019).ADS 
CAS 
Article 
PubMed 
PubMed Central 

Google Scholar 
Tilman, D., Cassman, K. G., Matson, P. A., Naylor, R. & Polasky, S. Agricultural sustainability and intensive production practices. Nature 418, 671–677. https://doi.org/10.1038/nature01014 (2002).ADS 
CAS 
Article 
PubMed 

Google Scholar 
Sharma, B. et al. Recycling of organic wastes in agriculture: An environmental perspective. Int. J. Environ. Res. 13, 409–429. https://doi.org/10.1007/s41742-019-00175-y (2019).CAS 
Article 

Google Scholar 
Pan, S. et al. Benefits of mechanized deep placement of nitrogen fertilizer in direct-seeded rice in South China. Field Crops Res. 203, 139–149. https://doi.org/10.1016/j.fcr.2016.12.011 (2017).Article 

Google Scholar 
Shahena, S., Rajan, M., Chandran, V. & Mathew, L. Conventional methods of fertilizer release. In Controlled Release Fertilizers for Sustainable Agriculture (eds Lewu, F. B. et al.) 1–24 (Academic Press, 2021). https://doi.org/10.1016/B978-0-12-819555-0.00001-7.Chapter 

Google Scholar 
Wang, C. et al. Effects of different fertilization methods on ammonia volatilization from rice paddies. J. Clean. Prod. 295, 126299. https://doi.org/10.1016/j.jclepro.2021.126299 (2021).CAS 
Article 

Google Scholar 
Wu, Q. et al. Effects of different types of slow- and controlled-release fertilizers on rice yield. J. Integr. Agric. 20, 1503–1514. https://doi.org/10.1016/S2095-3119(20)63406-2 (2021).CAS 
Article 

Google Scholar 
Mahajan, G., Kumar, V. & Chauhan, B. S. Rice production in India. In Rice production worldwide (eds Chauhan, B. et al.) 53–91 (Springer International Publishing, 2017). https://doi.org/10.1007/978-3-319-47516-5_3.Chapter 

Google Scholar 
Opoku-Kwanowaa, Y., Furaha, R. K., Yan, L. & Wei, D. Effects of planting field on groundwater and surface water pollution in China. Clean-Soil Air Water 48, 1900452. https://doi.org/10.1002/clen.201900452 (2020).CAS 
Article 

Google Scholar 
Lin, W. et al. The effects of chemical and organic fertilizer usage on rhizosphere soil in tea orchards. PLoS ONE 14, e0217018. https://doi.org/10.1371/journal.pone.0217018 (2019).CAS 
Article 
PubMed 
PubMed Central 

Google Scholar 
Sempeho, S. I., Kim, H. T., Mubofu, E. & Hilonga, A. Meticulous overview on the controlled release fertilizers. Adv. Chem. 1–16, 2014. https://doi.org/10.1155/2014/363071 (2014).Article 

Google Scholar 
Trenkel, M. E. Controlled-Release and Stabilized Fertilizers in Agriculture 1–156 (International Fertilizer Industry Association, 1997).
Google Scholar 
Lawrencia, D. et al. Controlled release fertilizers: A review on coating materials and mechanism of release. Plants 10, 238. https://doi.org/10.3390/plants10020238 (2021).CAS 
Article 
PubMed 
PubMed Central 

Google Scholar 
Tang, S. et al. Studies on the mechanism of single basal application of controlled-release fertilizers for increasing yield of rice (Oryza safiva L.). Agric. Sci. China 6, 586–596. https://doi.org/10.1016/S1671-2927(07)60087-X (2007).CAS 
Article 

Google Scholar 
Zheng, Y. et al. Effects of mixed controlled release nitrogen fertilizer with rice straw biochar on rice yield and nitrogen balance in northeast china. Sci. Rep. 10, 9452. https://doi.org/10.1038/s41598-020-66300-6 (2020).ADS 
CAS 
Article 
PubMed 
PubMed Central 

Google Scholar 
Ransom, C. J., Jolley, V. D., Blair, T. A., Sutton, L. E. & Hopkins, B. G. Nitrogen release rates from slow- and controlled-release fertilizers influenced by placement and temperature. PLoS ONE 15, e0234544. https://doi.org/10.1371/journal.pone.0234544 (2020).CAS 
Article 
PubMed 
PubMed Central 

Google Scholar 
Soni, R., Kumar, V., Suyal, D. C., Jain, L. & Goel, R. Metagenomics of plant rhizosphere microbiome. In Understanding host-microbiome interactions—an omics approach (eds Singh, R. et al.) 193–205 (Springer, 2017). https://doi.org/10.1007/978-981-10-5050-3_12.Chapter 

Google Scholar 
Kumar, A. Phosphate solubilizing bacteria in agriculture biotechnology: Diversity, mechanism and their role in plant growth and crop yield. Int. J. Adv. Res. 4, 116–124. https://doi.org/10.21474/IJAR01/111 (2016).Article 

Google Scholar 
Arjun, J. K. Metagenomic analysis of bacterial diversity in the rice rhizosphere soil microbiome. Biotechnol. Bioinf. Bioeng 1, 361–367 (2011).
Google Scholar 
Zhao, J. et al. Responses of bacterial communities in arable soils in a rice-wheat cropping system to different fertilizer regimes and sampling times. PLoS ONE 9, e85301. https://doi.org/10.1371/journal.pone.0085301 (2014).ADS 
CAS 
Article 
PubMed 
PubMed Central 

Google Scholar 
Huang, M. et al. Soil bacterial communities in three rice-based cropping systems differing in productivity. Sci. Rep. 10, 9867. https://doi.org/10.1038/s41598-020-66924-8 (2020).ADS 
CAS 
Article 
PubMed 
PubMed Central 

Google Scholar 
Hayatsu, M. A novel function of controlled-release nitrogen fertilizers. Microbes Environ. 29, 121–122. https://doi.org/10.1264/jsme2.ME2902rh (2014).Article 
PubMed 
PubMed Central 

Google Scholar 
Aslam, Z., Yasir, M., Yoon, H. S., Jeon, C. O. & Chung, Y. R. Diversity of the bacterial community in the rice rhizosphere managed under conventional and no-tillage practices. J. Microbiol. 51, 747–756. https://doi.org/10.1007/s12275-013-2528-8 (2013).Article 
PubMed 

Google Scholar 
Min, J. et al. Mechanical side-deep fertilization mitigates ammonia volatilization and nitrogen runoff and increases profitability in rice production independent of fertilizer type and split ratio. J. Clean. Prod. 316, 128370. https://doi.org/10.1016/j.jclepro.2021.128370 (2021).CAS 
Article 

Google Scholar 
Ke, J. et al. Combined controlled-released nitrogen fertilizers and deep placement effects of N leaching, rice yield and N recovery in machine-transplanted rice. Agr. Ecosyst. Environ. 265, 402–412. https://doi.org/10.1016/j.agee.2018.06.023 (2018).CAS 
Article 

Google Scholar 
Cardinale, B. J. et al. Effects of biodiversity on the functioning of trophic groups and ecosystems. Nature 443, 989–992. https://doi.org/10.1038/nature05202 (2006).ADS 
CAS 
Article 
PubMed 

Google Scholar 
Caporaso, J. G. et al. QIIME allows analysis of high-throughput community sequencing data. Nat. Methods 7, 335–336. https://doi.org/10.1038/nmeth.f.303 (2010).CAS 
Article 
PubMed 
PubMed Central 

Google Scholar 
Li, P. et al. Different regulation of soil structure and resource chemistry under animal- and plant-derived organic fertilizers changed soil bacterial communities. Appl. Soil. Ecol. 165, 104020. https://doi.org/10.1016/j.apsoil.2021.104020 (2021).Article 

Google Scholar 
Wang, J. et al. Wheat and rice growth stages and fertilization regimes alter soil bacterial community structure, but not diversity. Front. Microbiol. https://doi.org/10.3389/fmicb.2016.01207 (2016).Article 
PubMed 
PubMed Central 

Google Scholar 
Gu, Y., Zhang, X., Tu, S. & Lindström, K. Soil microbial biomass, crop yields, and bacterial community structure as affected by long-term fertilizer treatments under wheat-rice cropping. Eur. J. Soil Biol. 45, 239–246. https://doi.org/10.1016/j.ejsobi.2009.02.005 (2009).CAS 
Article 

Google Scholar 
Niu, J. et al. Insight into the effects of different cropping systems on soil bacterial community and tobacco bacterial wilt rate: Effects of different copping systems. J. Basic Microbiol. 57, 3–11. https://doi.org/10.1002/jobm.201600222 (2017).CAS 
Article 
PubMed 

Google Scholar 
Wu, T., Qin, Y. & Li, M. Intercropping of tea (Camellia sinensis L.) and Chinese chestnut: Variation in the structure of rhizosphere bacterial communities. J. Soil Sci. Plant Nutr. 21, 2178–2190. https://doi.org/10.1007/s42729-021-00513-0 (2021).CAS 
Article 

Google Scholar 
Li, Y. C. et al. Variations of rhizosphere bacterial communities in tea (Camellia sinensis L.) continuous cropping soil by high-throughput pyrosequencing approach. J. Appl. Microbiol. 121, 787–799. https://doi.org/10.1111/jam.13225 (2016).CAS 
Article 
PubMed 

Google Scholar 
Bei, Q., Moser, G., Müller, C. & Liesack, W. Seasonality affects function and complexity but not diversity of the rhizosphere microbiome in European temperate grassland. Sci. Total Environ. 784, 147036. https://doi.org/10.1016/j.scitotenv.2021.147036 (2021).ADS 
CAS 
Article 
PubMed 

Google Scholar 
You, J., Das, A., Dolan, E. M. & Hu, Z. Ammonia-oxidizing archaea involved in nitrogen removal. Water Res. 43, 1801–1809. https://doi.org/10.1016/j.watres.2009.01.016 (2009).CAS 
Article 
PubMed 

Google Scholar 
Chuang, S. et al. Potential effects of Rhodococcus qingshengii strain djl-6 on the bioremediation of carbendazim-contaminated soil and the assembly of its microbiome. J. Hazard. Mater. 414, 125496. https://doi.org/10.1016/j.jhazmat.2021.125496 (2021).CAS 
Article 
PubMed 

Google Scholar 
Luo, D. et al. The anaerobic oxidation of methane in paddy soil by ferric iron and nitrate, and the microbial communities involved. Sci. Total Environ. 788, 147773. https://doi.org/10.1016/j.scitotenv.2021.147773 (2021).ADS 
CAS 
Article 
PubMed 

Google Scholar 
Premnath, N. et al. A crucial review on polycyclic aromatic hydrocarbons—Environmental occurrence and strategies for microbial degradation. Chemosphere 280, 130608. https://doi.org/10.1016/j.chemosphere.2021.130608 (2021).ADS 
CAS 
Article 
PubMed 

Google Scholar 
Makino, A. Photosynthesis, grain yield, and nitrogen utilization in rice and wheat. Plant Physiol. 155, 125–129. https://doi.org/10.1104/pp.110.165076 (2011).CAS 
Article 
PubMed 

Google Scholar 
Sun, L., Lu, Y., Yu, F., Kronzucker, H. J. & Shi, W. Biological nitrification inhibition by rice root exudates and its relationship with nitrogen-use efficiency. New Phytol. 212, 646–656. https://doi.org/10.1111/nph.14057 (2016).CAS 
Article 
PubMed 

Google Scholar 
Coskun, D., Britto, D. T., Shi, W. & Kronzucker, H. J. How plant root exudates shape the nitrogen cycle. Trends Plant Sci. 22, 661–673. https://doi.org/10.1016/j.tplants.2017.05.004 (2017).CAS 
Article 
PubMed 

Google Scholar 
Qiang, S. et al. Deep placement of mixed controlled-release and conventional urea improves grain yield, nitrogen use efficiency of rainfed spring maize. Arch. Agronomy Soil Sci. 67, 1848–1858. https://doi.org/10.1080/03650340.2020.1817396 (2021).CAS 
Article 

Google Scholar 
Hou, P. et al. Deep fertilization with controlled-release fertilizer for higher cereal yield and N utilization in paddies: The optimal fertilization depth. Agronomy J. https://doi.org/10.1002/agj2.20772 (2021).Article 

Google Scholar 
Zhu, S., Vivanco, J. M. & Manter, D. K. Nitrogen fertilizer rate affects root exudation, the rhizosphere microbiome and nitrogen-use-efficiency of maize. Appl. Soil. Ecol. 107, 324–333. https://doi.org/10.1016/j.apsoil.2016.07.009 (2016).Article 

Google Scholar  More