{"status":"ok","message-type":"work","message-version":"1.0.0","message":{"indexed":{"date-parts":[[2025,8,2]],"date-time":"2025-08-02T18:30:52Z","timestamp":1754159452257,"version":"3.41.2"},"reference-count":81,"publisher":"Frontiers Media SA","license":[{"start":{"date-parts":[[2025,7,25]],"date-time":"2025-07-25T00:00:00Z","timestamp":1753401600000},"content-version":"vor","delay-in-days":0,"URL":"https:\/\/creativecommons.org\/licenses\/by\/4.0\/"}],"content-domain":{"domain":["frontiersin.org"],"crossmark-restriction":true},"short-container-title":["Front. Microbiol."],"abstract":"<jats:p>The linkage between methane emissions and the metabolic activity of archaeal species is broadly established. However, the structural and functional dynamics of this phenomenon within the scarab larval gut and associated host soil environment have not been investigated. In this study, we used shotgun metagenome sequencing to explore the archaeal communities associated with the digestive tract of third instar Japanese beetle (<jats:italic>Popillia japonica<\/jats:italic> Newman; Coleoptera: Scarabaeidae) (JB) larvae and its host soil. Our findings showed that both the JB gut compartment (midgut <jats:italic>vs<\/jats:italic>. hindgut) and experimental conditions (field <jats:italic>vs<\/jats:italic>. manipulative laboratory studies) significantly affect the composition of archaeal taxa. Moreover, gut compartment affected the functional profile. Results revealed an increase of methane metabolism-related taxa and gene sequences in the larval hindgut, supporting the hypothesis that methanogenesis is primarily maintained in that gut compartment. Methane production associated with the JB larval gut takes place primarily via CO<jats:sub>2<\/jats:sub> reduction (~30%) and methanol methanation (4%) pathways. The presence of the same archaeal features in both soil and JB midgut suggests that the JB midgut archaeome may be environmentally sourced, with more tailored selection of the archaeome occurring in the JB hindgut. In turn, we found that JB larval infestation also increases the abundance of at least one methanogenic archaeon, <jats:italic>Methanobrevibacter<\/jats:italic>, in infested soil. Results underscore the potential impact of invasive root-feeding scarab larvae on the soil archaeome and highlight their potential contributions to climate change, especially in light of predicted global range expansion for this species.<\/jats:p>","DOI":"10.3389\/fmicb.2025.1609893","type":"journal-article","created":{"date-parts":[[2025,7,25]],"date-time":"2025-07-25T09:41:23Z","timestamp":1753436483000},"update-policy":"https:\/\/doi.org\/10.3389\/crossmark-policy","source":"Crossref","is-referenced-by-count":0,"title":["Metagenomic analysis reveals methanogenic and other archaeal genes in the digestive tract of invasive Japanese beetle larvae and associated soil"],"prefix":"10.3389","volume":"16","author":[{"given":"Helena","family":"Avila-Arias","sequence":"first","affiliation":[],"role":[{"role":"author","vocabulary":"crossref"}]},{"given":"Michael E.","family":"Scharf","sequence":"additional","affiliation":[],"role":[{"role":"author","vocabulary":"crossref"}]},{"given":"Ronald F.","family":"Turco","sequence":"additional","affiliation":[],"role":[{"role":"author","vocabulary":"crossref"}]},{"given":"Diego J.","family":"Jim\u00e9nez","sequence":"additional","affiliation":[],"role":[{"role":"author","vocabulary":"crossref"}]},{"given":"Audrey","family":"Simard","sequence":"additional","affiliation":[],"role":[{"role":"author","vocabulary":"crossref"}]},{"given":"Douglas S.","family":"Richmond","sequence":"additional","affiliation":[],"role":[{"role":"author","vocabulary":"crossref"}]}],"member":"1965","published-online":{"date-parts":[[2025,7,25]]},"reference":[{"key":"ref1","doi-asserted-by":"publisher","first-page":"e01202-20","DOI":"10.1128\/msphere.01202-01220","article-title":"Primer, pipelines, parameters: issues in 16S rRNA gene sequencing","volume":"6","author":"Abellan-Schneyder","year":"2021","journal-title":"mSphere"},{"key":"ref2","doi-asserted-by":"publisher","first-page":"2","DOI":"10.1093\/jipm\/pmab043","article-title":"Japanese beetle (Coleoptera: Scarabaeidae) invasion of North America: history, ecology, and management","volume":"13","author":"Althoff","year":"2022","journal-title":"J. Integr. Pest Manag."},{"key":"ref3","doi-asserted-by":"publisher","first-page":"854513","DOI":"10.3389\/fmicb.2022.854513","article-title":"Soil environments influence gut prokaryotic communities in the larvae of the invasive Japanese beetle Popillia japonica Newman","volume":"13","author":"Avila-Arias","year":"2022","journal-title":"Front. Microbiol."},{"key":"ref4","doi-asserted-by":"publisher","first-page":"1102523","DOI":"10.3389\/fmicb.2023.1102523","article-title":"Larvae of an invasive scarab increase greenhouse gas emissions from soils and recruit gut mycobiota involved in C and N transformations","volume":"14","author":"Avila-Arias","year":"2023","journal-title":"Front. Microbiol."},{"key":"ref5","doi-asserted-by":"publisher","first-page":"887","DOI":"10.1038\/s41564-020-0715-z","article-title":"Diversity, ecology and evolution of Archaea","volume":"5","author":"Baker","year":"2020","journal-title":"Nat. Microbiol."},{"key":"ref6","doi-asserted-by":"publisher","first-page":"852","DOI":"10.1038\/s41587-019-0209-9","article-title":"Reproducible, interactive, scalable and extensible microbiome data science using QIIME 2","volume":"37","author":"Bolyen","year":"2019","journal-title":"Nat. Biotechnol."},{"key":"ref7","doi-asserted-by":"publisher","first-page":"622","DOI":"10.1038\/s41579-020-0407-y","article-title":"The host-associated archaeome","volume":"18","author":"Borrel","year":"2020","journal-title":"Nat. Rev. Microbiol."},{"key":"ref8","doi-asserted-by":"publisher","first-page":"27","DOI":"10.1111\/j.1574-6941.2001.tb00785.x","article-title":"Molecular phylogenetic profiling of prokaryotic communities in guts of termites with different feeding habits","volume":"35","author":"Brauman","year":"2001","journal-title":"FEMS Microbiol. Ecol."},{"key":"ref9","doi-asserted-by":"crossref","first-page":"229","DOI":"10.1007\/978-3-319-78108-2_13","article-title":"Methanogenesis in the digestive tracts of insects and other arthropods","volume-title":"Biogenesis of Hydrocarbons","author":"Brune","year":"2019"},{"key":"ref10","doi-asserted-by":"publisher","first-page":"e0002422","DOI":"10.1128\/mmbr.00024-22","article-title":"Methyl-based methanogenesis: an ecological and genomic review","volume":"87","author":"Bueno De Mesquita","year":"2023","journal-title":"Microbiol. Mol. Biol. Rev."},{"key":"ref11","doi-asserted-by":"publisher","first-page":"135","DOI":"10.1023\/A:1023325817663","article-title":"Promicromonospora pachnodae sp. nov., a member of the (hemi)cellulolytic hindgut flora of larvae of the scarab beetle Pachnoda marginata","volume":"83","author":"Cazemier","year":"2003","journal-title":"Antonie Van Leeuwenhoek"},{"key":"ref12","doi-asserted-by":"publisher","first-page":"799","DOI":"10.1038\/s41596-019-0264-1","article-title":"Using MicrobiomeAnalyst for comprehensive statistical, functional, and meta-analysis of microbiome data","volume":"15","author":"Chong","year":"2020","journal-title":"Nat. Protoc."},{"key":"ref13","doi-asserted-by":"publisher","first-page":"4343","DOI":"10.1111\/1462-2920.14797","article-title":"Developmental stages and gut microenvironments influence gut microbiota dynamics in the invasive beetle Popillia japonica Newman (Coleoptera: Scarabaeidae)","volume":"21","author":"Chouaia","year":"2019","journal-title":"Environ. Microbiol."},{"key":"ref14","doi-asserted-by":"publisher","first-page":"108","DOI":"10.1016\/j.cbpa.2015.10.018","article-title":"Lignocellulose degradation mechanisms across the Tree of Life","volume":"29","author":"Cragg","year":"2015","journal-title":"Curr. Opin. Chem. Biol."},{"key":"ref15","doi-asserted-by":"publisher","first-page":"525","DOI":"10.1007\/s002030050606","article-title":"Acetyl-CoA decarbonylase\/synthase complex from Archaeoglobus fulgidus","volume":"169","author":"Dai","year":"1998","journal-title":"Arch. Microbiol."},{"key":"ref16","doi-asserted-by":"publisher","first-page":"291","DOI":"10.1111\/1758-2229.13152","article-title":"Effects of 5-year experimental warming in the Alpine belt on soil Archaea: Multi-omics approaches and prospects","volume":"15","author":"D'al\u00f2","year":"2023","journal-title":"Environ. Microbiol. Rep."},{"key":"ref17","doi-asserted-by":"publisher","first-page":"766","DOI":"10.3390\/app13020766","article-title":"The current state and future potential of microbial control of scarab pests","volume":"13","author":"Deans","year":"2023","journal-title":"Appl. Sci."},{"key":"ref18","doi-asserted-by":"publisher","first-page":"567","DOI":"10.3390\/land11040567","article-title":"The new dominator of the world: modeling the global distribution of the Japanese beetle under land use and climate change scenarios","volume":"11","author":"Della Rocca","year":"","journal-title":"Land"},{"key":"ref19","doi-asserted-by":"publisher","first-page":"658","DOI":"10.3390\/d14080658","article-title":"The spread of the Japanese beetle in a European human-dominated landscape: high anthropization favors colonization of Popillia japonica","volume":"14","author":"Della Rocca","year":"","journal-title":"Diversity"},{"key":"ref20","doi-asserted-by":"publisher","first-page":"W180","DOI":"10.1093\/nar\/gkx295","article-title":"MicrobiomeAnalyst: a web-based tool for comprehensive statistical, visual and meta-analysis of microbiome data","volume":"45","author":"Dhariwal","year":"2017","journal-title":"Nucleic Acids Res."},{"key":"ref21","doi-asserted-by":"publisher","first-page":"1348","DOI":"10.1111\/1462-2920.15434","article-title":"Anaerobic microbial methanol conversion in marine sediments","volume":"23","author":"Fischer","year":"2021","journal-title":"Environ. Microbiol."},{"key":"ref22","doi-asserted-by":"publisher","first-page":"321","DOI":"10.3389\/fpls.2016.00321","article-title":"Belowground ecology of scarabs feeding on grass roots: current knowledge and future directions for management in Australasia","volume":"7","author":"Frew","year":"2016","journal-title":"Front. Plant Sci."},{"key":"ref23","doi-asserted-by":"publisher","first-page":"1972","DOI":"10.3390\/microorganisms11081972","article-title":"Exploring Gut Microbiome Variations between Popillia japonica Populations of Azores","volume":"11","author":"Frias","year":"2023","journal-title":"Microorganisms"},{"key":"ref24","doi-asserted-by":"publisher","first-page":"16062","DOI":"10.3748\/wjg.v20.i43.16062","article-title":"Archaea and the human gut: new beginning of an old story","volume":"20","author":"Gaci","year":"2014","journal-title":"World J. Gastroenterol."},{"key":"ref25","doi-asserted-by":"publisher","first-page":"112","DOI":"10.1016\/j.rhisph.2018.06.003","article-title":"Root herbivores accelerate carbon inputs to soil and drive changes in biogeochemical processes","volume":"6","author":"Gan","year":"2018","journal-title":"Rhizosphere"},{"key":"ref26","doi-asserted-by":"publisher","first-page":"107972","DOI":"10.1016\/j.soilbio.2020.107972","article-title":"Root herbivory and soil carbon cycling: Shedding \u201cgreen\u201d light onto a \u201cbrown\u201d world","volume":"150","author":"Gan","year":"2020","journal-title":"Soil Biol. Biochem."},{"key":"ref27","doi-asserted-by":"publisher","first-page":"e0238057","DOI":"10.1371\/journal.pone.0238057","article-title":"First field estimation of greenhouse gas release from European soil-dwelling Scarabaeidae larvae targeting the genus Melolontha","volume":"15","author":"G\u00f6rres","year":"2020","journal-title":"PLoS One"},{"key":"ref28","doi-asserted-by":"publisher","first-page":"32","DOI":"10.1007\/s00203-005-0006-3","article-title":"A single operon-encoded form of the acetyl-CoA decarbonylase\/synthase multienzyme complex responsible for synthesis and cleavage of acetyl-CoA in Methanosarcina thermophila","volume":"184","author":"Grahame","year":"2005","journal-title":"Arch. Microbiol."},{"key":"ref29","doi-asserted-by":"publisher","first-page":"156","DOI":"10.1111\/eea.12768","article-title":"The microbiome of pest insects: it is not just bacteria","volume":"167","author":"Gurung","year":"2019","journal-title":"Entomol. Exp. Appl."},{"key":"ref30","doi-asserted-by":"publisher","first-page":"622824","DOI":"10.3389\/fmicb.2020.622824","article-title":"Genomic insights into the lifestyles of thaumarchaeota inside sponges","volume":"11","author":"Haber","year":"2021","journal-title":"Front. Microbiol."},{"key":"ref31","doi-asserted-by":"publisher","first-page":"5441","DOI":"10.1073\/pnas.91.12.5441","article-title":"Methane production in terrestrial arthropods","volume":"91","author":"Hackstein","year":"1994","journal-title":"Proc. Natl. Acad. Sci. USA"},{"key":"ref32","first-page":"121","article-title":"Methanogens in the Gastrointestinal Tract of Animals","volume-title":"Microbiology Monographs","author":"Hackstein","year":"2018"},{"key":"ref33","doi-asserted-by":"publisher","first-page":"175","DOI":"10.1111\/j.1744-7917.2010.01320.x","article-title":"The scarab gut: a potential bioreactor for bio-fuel production","volume":"17","author":"Huang","year":"2010","journal-title":"Insect Sci."},{"key":"ref34","doi-asserted-by":"publisher","first-page":"162","DOI":"10.1016\/j.mib.2018.01.013","article-title":"The soil microbiome\u2014from metagenomics to metaphenomics","volume":"43","author":"Jansson","year":"2018","journal-title":"Curr. Opin. Microbiol."},{"key":"ref35","doi-asserted-by":"publisher","first-page":"13845","DOI":"10.1038\/srep13845","article-title":"Unveiling the metabolic potential of two soil-derived microbial consortia selected on wheat straw","volume":"5","author":"Jim\u00e9nez","year":"2015","journal-title":"Sci. Rep."},{"key":"ref36","doi-asserted-by":"publisher","first-page":"517","DOI":"10.1146\/annurev-ento-010814-020608","article-title":"Root-feeding insects and their interactions with organisms in the rhizosphere","volume":"60","author":"Johnson","year":"2015","journal-title":"Annu. Rev. Entomol."},{"key":"ref37","doi-asserted-by":"publisher","first-page":"491","DOI":"10.1007\/s00374-017-1199-8","article-title":"Stimulation of methane oxidation by CH4-emitting rose chafer larvae in well-aerated grassland soil","volume":"53","author":"Kammann","year":"2017","journal-title":"Biol. Fertil. Soils"},{"key":"ref38","doi-asserted-by":"publisher","first-page":"D457","DOI":"10.1093\/nar\/gkv1070","article-title":"KEGG as a reference resource for gene and protein annotation","volume":"44","author":"Kanehisa","year":"2016","journal-title":"Nucleic Acids Res."},{"key":"ref39","article-title":"ARTool: aligned rank transform for nonparametric factorial ANOVAs","volume-title":"R package version 0.10.6","author":"Kay","year":"2019"},{"key":"ref40","doi-asserted-by":"crossref","first-page":"207","DOI":"10.1007\/978-1-4939-3369-3_13","article-title":"MG-RAST, a Metagenomics Service for Analysis of Microbial Community Structure and Function","volume-title":"Microbial Environmental Genomics (MEG)","author":"Keegan","year":"2016"},{"key":"ref41","doi-asserted-by":"publisher","first-page":"e1002541","DOI":"10.1371\/journal.pcbi.1002541","article-title":"A platform-independent method for detecting errors in metagenomic sequencing data: DRISEE","volume":"8","author":"Keegan","year":"2012","journal-title":"PLoS Comput. Biol."},{"key":"ref42","doi-asserted-by":"publisher","first-page":"656","DOI":"10.1101\/gr.229202","article-title":"BLAT--the BLAST-like alignment tool","volume":"12","author":"Kent","year":"2002","journal-title":"Genome Res."},{"key":"ref43","doi-asserted-by":"publisher","first-page":"1","DOI":"10.1186\/s40168-020-00894-x","article-title":"The human gut archaeome: identification of diverse haloarchaea in Korean subjects","volume":"8","author":"Kim","year":"2020","journal-title":"Microbiome"},{"key":"ref44","doi-asserted-by":"publisher","first-page":"1","DOI":"10.1093\/jisesa\/iez023","article-title":"The potential global distribution and voltinism of the Japanese Beetle (Coleoptera: Scarabaeidae) under current and future climates","volume":"19","author":"Kistner-Thomas","year":"2019","journal-title":"J. Insect Sci."},{"key":"ref45","doi-asserted-by":"publisher","first-page":"993","DOI":"10.3389\/fmicb.2020.00993","article-title":"The core gut microbiome of Black soldier fly (Hermetia illucens) larvae raised on low-bioburden diets","volume":"11","author":"Klammsteiner","year":"2020","journal-title":"Front. Microbiol."},{"key":"ref46","doi-asserted-by":"publisher","first-page":"357","DOI":"10.1038\/nmeth.1923","article-title":"Fast gapped-read alignment with Bowtie 2","volume":"9","author":"Langmead","year":"2012","journal-title":"Nat. Methods"},{"key":"ref47","doi-asserted-by":"publisher","first-page":"6650","DOI":"10.1128\/AEM.69.11.6650-6658.2003","article-title":"Physicochemical conditions and microbial activities in the highly alkaline gut of the humus-feeding larva of Pachnoda ephippiata (Coleoptera: Scarabaeidae)","volume":"69","author":"Lemke","year":"2003","journal-title":"Appl. Environ. Microbiol."},{"key":"ref48","doi-asserted-by":"publisher","first-page":"550","DOI":"10.1186\/s13059-014-0550-8","article-title":"Moderated estimation of fold change and dispersion for RNA-seq data with DESeq2","volume":"15","author":"Love","year":"2014","journal-title":"Genome Biol."},{"key":"ref49","doi-asserted-by":"crossref","first-page":"19","DOI":"10.1007\/978-3-319-78108-2_5","article-title":"Diversity and Taxonomy of Methanogens","volume-title":"Biogenesis of Hydrocarbons","author":"Lyu","year":"2019"},{"key":"ref50","doi-asserted-by":"publisher","first-page":"170687","DOI":"10.1016\/j.scitotenv.2024.170687","article-title":"Invasive Japanese beetle (Popillia japonica Newman) larvae alter structure and carbon distribution in infested surface soil","volume":"918","author":"Macleod","year":"2024","journal-title":"Sci. Total Environ."},{"key":"ref51","doi-asserted-by":"publisher","first-page":"37","DOI":"10.1016\/j.soilbio.2014.03.025","article-title":"Contribution of white grubs (Scarabaeidae: Coleoptera) to N2O emissions from tropical soils","volume":"75","author":"Majeed","year":"2014","journal-title":"Soil Biol. Biochem."},{"key":"ref52","doi-asserted-by":"publisher","first-page":"e00016","DOI":"10.1128\/msystems.00016-19","article-title":"A novel sparse compositional technique reveals microbial perturbations","volume":"4","author":"Martino","year":"2019","journal-title":"mSystems"},{"key":"ref53","first-page":"665","article-title":"Methanogens and methanogenesis in hypersaline environments","volume-title":"Biogenesis of Hydrocarbons, Handbook of Hydrocarbon and Lipid Microbiology","author":"Mcgenity","year":"2018"},{"key":"ref54","doi-asserted-by":"publisher","first-page":"386","DOI":"10.1186\/1471-2105-9-386","article-title":"The metagenomics RAST server \u2013 a public resource for the automatic phylogenetic and functional analysis of metagenomes","volume":"9","author":"Meyer","year":"2008","journal-title":"BMC Bioinformatics"},{"key":"ref55","doi-asserted-by":"publisher","first-page":"D733","DOI":"10.1093\/nar\/gkv1189","article-title":"Reference sequence (RefSeq) database at NCBI: current status, taxonomic expansion, and functional annotation","volume":"44","author":"O\u2019Leary","year":"2015","journal-title":"Nucleic Acids Res."},{"key":"ref56","doi-asserted-by":"crossref","first-page":"259","DOI":"10.1007\/978-3-642-38954-2_408","article-title":"The Family Methanosarcinaceae","volume-title":"The Prokaryotes: Other Major Lineages of Bacteria and The Archaea","author":"Oren","year":"2014"},{"key":"ref57","doi-asserted-by":"publisher","first-page":"6781","DOI":"10.1038\/s41598-021-86378-w","article-title":"Evaluation of 16S rRNA primer sets for characterisation of microbiota in paediatric patients with autism spectrum disorder","volume":"11","author":"Palkova","year":"2021","journal-title":"Sci. Rep."},{"key":"ref58","doi-asserted-by":"publisher","first-page":"1079756","DOI":"10.3389\/finsc.2022.1079756","article-title":"A nationwide pest risk analysis in the context of the ongoing Japanese beetle invasion in continental Europe: the case of metropolitan France","volume":"2","author":"Poggi","year":"2022","journal-title":"Front. Insect Sci."},{"key":"ref59","doi-asserted-by":"publisher","first-page":"175","DOI":"10.1146\/annurev.ento.47.091201.145153","article-title":"Biology and management of the Japanese beetle","volume":"47","author":"Potter","year":"2002","journal-title":"Annu. Rev. Entomol."},{"key":"ref60","doi-asserted-by":"crossref","first-page":"277","DOI":"10.1007\/978-981-10-7485-1_14","article-title":"Insect Gut Bacteria and Their Potential Application in Degradation of Lignocellulosic Biomass: A Review","volume-title":"Bioremediation: Applications for Environmental Protection and Management","author":"Prasad","year":"2018"},{"key":"ref61","doi-asserted-by":"publisher","first-page":"1281628","DOI":"10.3389\/fmicb.2023.1281628","article-title":"Diversity and taxonomic revision of methanogens and other archaea in the intestinal tract of terrestrial arthropods","volume":"14","author":"Protasov","year":"2023","journal-title":"Front. Microbiol."},{"key":"ref62","doi-asserted-by":"publisher","first-page":"fiae111","DOI":"10.1093\/femsec\/fiae111","article-title":"Genome reduction in novel, obligately methyl-reducing Methanosarcinales isolated from arthropod guts (Methanolapillus gen. nov. and Methanimicrococcus)","volume":"100","author":"Protasov","year":"2024","journal-title":"FEMS Microbiol. Ecol."},{"key":"ref63","doi-asserted-by":"publisher","first-page":"D61","DOI":"10.1093\/nar\/gkl842","article-title":"NCBI reference sequences (RefSeq): a curated non-redundant sequence database of genomes, transcripts and proteins","volume":"35","author":"Pruitt","year":"2007","journal-title":"Nucleic Acids Res."},{"key":"ref64","first-page":"63","article-title":"The distribution and diversity of Euryarchaeota in termite guts","volume-title":"Advances in applied microbiology","author":"Purdy","year":"2007"},{"key":"ref65","article-title":"Managing white grubs in turfgrass","author":"Richmond","year":"2022"},{"key":"ref66","doi-asserted-by":"publisher","first-page":"3","DOI":"10.1111\/gcb.14893","article-title":"Global change biology: A primer","volume":"26","author":"Sage","year":"2020","journal-title":"Glob. Chang. Biol."},{"key":"ref67","doi-asserted-by":"publisher","first-page":"23","DOI":"10.1146\/annurev-ento-022420-074746","article-title":"A century of synergy in termite symbiosis research: Linking the past with new genomic insights","volume":"66","author":"Scharf","year":"2021","journal-title":"Annu. Rev. Entomol."},{"key":"ref68","doi-asserted-by":"publisher","first-page":"1882","DOI":"10.1111\/1462-2920.12086","article-title":"Comparative metagenomic and rRNA microbial diversity characterization using archaeal and bacterial synthetic communities","volume":"15","author":"Shakya","year":"2013","journal-title":"Environ. Microbiol."},{"key":"ref69","doi-asserted-by":"publisher","first-page":"1","DOI":"10.1093\/jipm\/pmz009","article-title":"Biology and management of Japanese beetle (Coleoptera: Scarabaeidae) in corn and soybean","volume":"10","author":"Shanovich","year":"2019","journal-title":"J. Integr. Pest Manag."},{"key":"ref70","doi-asserted-by":"publisher","first-page":"690","DOI":"10.1186\/s12864-022-08906-w","article-title":"Genome-resolved analyses show an extensive diversification in key aerobic hydrocarbon-degrading enzymes across bacteria and archaea","volume":"23","author":"Somee","year":"2022","journal-title":"BMC Genomics"},{"key":"ref71","doi-asserted-by":"publisher","first-page":"104147","DOI":"10.1016\/j.jprot.2021.104147","article-title":"The structure and function of soil archaea across biomes","volume":"237","author":"Starke","year":"2021","journal-title":"J. Proteome"},{"key":"ref72","doi-asserted-by":"crossref","first-page":"496","DOI":"10.1093\/aesa\/24.3.496","article-title":"The influence of soil acidity on the pH value of the contents of the digestive tract of Japanese beetle larvae","volume":"24","author":"Swingle","year":"1931","journal-title":"Ann. Entomol. Soc. Am."},{"key":"ref73","doi-asserted-by":"publisher","first-page":"494","DOI":"10.3389\/fmicb.2017.00494","article-title":"Comparative evaluation of four bacteria-specific primer pairs for 16S rRNA gene surveys","volume":"8","author":"Thijs","year":"2017","journal-title":"Front. Microbiol."},{"key":"ref74","doi-asserted-by":"publisher","first-page":"3358","DOI":"10.1038\/s41467-022-31038-4","article-title":"Factors shaping the abundance and diversity of the gut archaeome across the animal kingdom","volume":"13","author":"Thomas","year":"2022","journal-title":"Nat. Commun."},{"key":"ref75","doi-asserted-by":"publisher","first-page":"203","DOI":"10.1016\/j.apsoil.2004.08.004","article-title":"Effects of root feeding, cranefly larvae on soil microorganisms and the composition of rhizosphere solutions collected from grassland plants","volume":"28","author":"Treonis","year":"2005","journal-title":"Appl. Soil Ecol."},{"key":"ref76","doi-asserted-by":"publisher","first-page":"1","DOI":"10.1186\/2047-217X-2-16","article-title":"EMPeror: a tool for visualizing high-throughput microbial community data","volume":"2","author":"V\u00e1zquez-Baeza","year":"2013","journal-title":"GigaScience"},{"key":"ref77","doi-asserted-by":"publisher","first-page":"33","DOI":"10.1111\/ens.12394","article-title":"Site-specific profiles of biochemical properties in the larval digestive tract of Japanese rhinoceros beetle, Trypoxylus dichotomus (Coleoptera: Scarabaeidae)","volume":"23","author":"Wada","year":"2020","journal-title":"Entomol. Sci."},{"key":"ref78","doi-asserted-by":"publisher","first-page":"27","DOI":"10.1186\/s40168-017-0237-y","article-title":"Normalization and microbial differential abundance strategies depend upon data characteristics","volume":"5","author":"Weiss","year":"2017","journal-title":"Microbiome"},{"key":"ref79","doi-asserted-by":"publisher","first-page":"141","DOI":"10.1186\/1471-2105-13-141","article-title":"The M5nr: a novel non-redundant database containing protein sequences and annotations from multiple sources and associated tools","volume":"13","author":"Wilke","year":"2012","journal-title":"BMC Bioinformatics"},{"key":"ref80","doi-asserted-by":"crossref","DOI":"10.1145\/1978942.1978963","article-title":"The aligned rank transform for nonparametric factorial analyses using only anova procedures","author":"Wobbrock","year":"2011"},{"key":"ref81","doi-asserted-by":"publisher","first-page":"e3905","DOI":"10.1002\/ecy.3905","article-title":"Species-level termite methane production rates","volume":"104","author":"Zhou","year":"2023","journal-title":"Ecology"}],"container-title":["Frontiers in Microbiology"],"original-title":[],"link":[{"URL":"https:\/\/www.frontiersin.org\/articles\/10.3389\/fmicb.2025.1609893\/full","content-type":"unspecified","content-version":"vor","intended-application":"similarity-checking"}],"deposited":{"date-parts":[[2025,7,25]],"date-time":"2025-07-25T09:41:26Z","timestamp":1753436486000},"score":1,"resource":{"primary":{"URL":"https:\/\/www.frontiersin.org\/articles\/10.3389\/fmicb.2025.1609893\/full"}},"subtitle":[],"short-title":[],"issued":{"date-parts":[[2025,7,25]]},"references-count":81,"alternative-id":["10.3389\/fmicb.2025.1609893"],"URL":"https:\/\/doi.org\/10.3389\/fmicb.2025.1609893","relation":{},"ISSN":["1664-302X"],"issn-type":[{"value":"1664-302X","type":"electronic"}],"subject":[],"published":{"date-parts":[[2025,7,25]]},"article-number":"1609893"}}