{"status":"ok","message-type":"work","message-version":"1.0.0","message":{"indexed":{"date-parts":[[2026,9,7]],"date-time":"2026-09-07T16:08:06Z","timestamp":1788797286816,"version":"build-2803163510"},"reference-count":104,"publisher":"Oxford University Press (OUP)","license":[{"start":{"date-parts":[[2026,8,18]],"date-time":"2026-08-18T00:00:00Z","timestamp":1787011200000},"content-version":"am","delay-in-days":0,"URL":"https:\/\/academic.oup.com\/pages\/standard-publication-reuse-rights"}],"funder":[{"DOI":"10.13039\/501100012166","name":"National Key Research and Development Program of China","doi-asserted-by":"publisher","award":["2024YFC2607800"],"award-info":[{"award-number":["2024YFC2607800"]}],"id":[{"id":"10.13039\/501100012166","id-type":"DOI","asserted-by":"publisher"}]},{"DOI":"10.13039\/501100012166","name":"National Key Research and Development Program of China","doi-asserted-by":"publisher","award":["2024YFC2607804"],"award-info":[{"award-number":["2024YFC2607804"]}],"id":[{"id":"10.13039\/501100012166","id-type":"DOI","asserted-by":"publisher"}]},{"DOI":"10.13039\/501100001809","name":"National Natural Science Foundation of China","doi-asserted-by":"publisher","award":["31730086"],"award-info":[{"award-number":["31730086"]}],"id":[{"id":"10.13039\/501100001809","id-type":"DOI","asserted-by":"publisher"}]},{"DOI":"10.13039\/501100001809","name":"National Natural Science Foundation of China","doi-asserted-by":"publisher","award":["32100357"],"award-info":[{"award-number":["32100357"]}],"id":[{"id":"10.13039\/501100001809","id-type":"DOI","asserted-by":"publisher"}]},{"name":"Northeast Asia Biodiversity Research Center","award":["2572022DS09"],"award-info":[{"award-number":["2572022DS09"]}]},{"name":"Expert Workstation in Zhaotong, Yunnan","award":["2021ZTYX05"],"award-info":[{"award-number":["2021ZTYX05"]}]},{"DOI":"10.13039\/501100002365","name":"China Agricultural University","doi-asserted-by":"publisher","id":[{"id":"10.13039\/501100002365","id-type":"DOI","asserted-by":"publisher"}]}],"content-domain":{"domain":[],"crossmark-restriction":false},"short-container-title":[],"abstract":"<jats:title>Abstract<\/jats:title>\n                  <jats:p>Dietary adaptations are fundamental drivers of evolutionary innovations. Still, the morphological and molecular mechanisms underlying these transitions remain poorly understood. True bugs (Heteroptera), one of the most diverse clades of hemimetabolous insects, have evolved a vast array of trophic niches. They exhibit remarkable structural and functional diversity in their salivary glands, providing an ideal system for investigating how dietary variation shapes evolutionary trajectories. Using morphometric, proteo-transcriptomic, and comparative phylogenetic approaches, we examined the evolutionary patterns of salivary glands across 220 hemipteran species, encompassing all seven infraorders and six dietary categories of Heteroptera. Our results reveal strong correlations between dietary shifts, salivary gland size, and allometric scaling. Ancestral state estimation analysis demonstrates multiple evolutionary transitions in salivary gland morphology, highlighting structural adaptations associated with different diets. Furthermore, we found that distantly related species with similar diets share dominant salivary protein groups, implying convergent molecular solutions to shared feeding challenges. Notably, we identify venom protein family 2 as a key adaptation. This family originated through lineage-specific co-option of an ancestral protein into heteropteran salivary systems, followed by extensive expansion that facilitated dietary diversification. These findings provide novel insights into the morphological, physiological, and molecular adaptations of salivary glands, emphasizing the dynamic interplay between feeding ecology and macroevolution.<\/jats:p>","DOI":"10.1093\/sysbio\/syag067","type":"journal-article","created":{"date-parts":[[2026,8,17]],"date-time":"2026-08-17T11:19:26Z","timestamp":1786965566000},"source":"Crossref","is-referenced-by-count":0,"title":["Coordinated Evolution of Salivary Glands and Sialome in True Bugs: Dietary Adaptation and Venom Protein Co-option"],"prefix":"10.1093","author":[{"given":"Xinyu","family":"Li","sequence":"first","affiliation":[{"name":"State Key Laboratory of Agricultural and Forestry Biosecurity, MOA Key Lab of Pest Monitoring and Green Management, College of Plant Protection, China Agricultural University , 2 Yuanmingyuan West Road, Haidian District, Beijing 100193 ,","place":["China"]},{"name":"Beijing Key Laboratory for Forest Pest Control, College of Forestry, Beijing Forestry University , 35 Qinghua East Road, Haidian District, Beijing 100083 ,","place":["China"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Tianfang","family":"Wang","sequence":"additional","affiliation":[{"name":"Centre for Bioinnovation, University of the Sunshine Coast , 90 Sippy Downs Drive, Sippy Downs, Queensland 4556 ,","place":["Australia"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Fanding","family":"Gao","sequence":"additional","affiliation":[{"name":"State Key Laboratory of Agricultural and Forestry Biosecurity, MOA Key Lab of Pest Monitoring and Green Management, College of Plant Protection, China Agricultural University , 2 Yuanmingyuan West Road, Haidian District, Beijing 100193 ,","place":["China"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0003-2311-9859","authenticated-orcid":false,"given":"Yuange","family":"Duan","sequence":"additional","affiliation":[{"name":"State Key Laboratory of Agricultural and Forestry Biosecurity, MOA Key Lab of Pest Monitoring and Green Management, College of Plant Protection, China Agricultural University , 2 Yuanmingyuan West Road, Haidian District, Beijing 100193 ,","place":["China"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0009-0000-3752-8173","authenticated-orcid":false,"given":"Ling","family":"Ma","sequence":"additional","affiliation":[{"name":"State Key Laboratory of Agricultural and Forestry Biosecurity, MOA Key Lab of Pest Monitoring and Green Management, College of Plant Protection, China Agricultural University , 2 Yuanmingyuan West Road, Haidian District, Beijing 100193 ,","place":["China"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0003-3393-0338","authenticated-orcid":false,"given":"Zhuo","family":"Chen","sequence":"additional","affiliation":[{"name":"State Key Laboratory of Agricultural and Forestry Biosecurity, MOA Key Lab of Pest Monitoring and Green Management, College of Plant Protection, China Agricultural University , 2 Yuanmingyuan West Road, Haidian District, Beijing 100193 ,","place":["China"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Zhiyao","family":"Wang","sequence":"additional","affiliation":[{"name":"State Key Laboratory of Agricultural and Forestry Biosecurity, MOA Key Lab of Pest Monitoring and Green Management, College of Plant Protection, China Agricultural University , 2 Yuanmingyuan West Road, Haidian District, Beijing 100193 ,","place":["China"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0002-7288-9676","authenticated-orcid":false,"given":"Li","family":"Tian","sequence":"additional","affiliation":[{"name":"State Key Laboratory of Agricultural and Forestry Biosecurity, MOA Key Lab of Pest Monitoring and Green Management, College of Plant Protection, China Agricultural University , 2 Yuanmingyuan West Road, Haidian District, Beijing 100193 ,","place":["China"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0002-2900-4174","authenticated-orcid":false,"given":"Fan","family":"Song","sequence":"additional","affiliation":[{"name":"State Key Laboratory of Agricultural and Forestry Biosecurity, MOA Key Lab of Pest Monitoring and Green Management, College of Plant Protection, China Agricultural University , 2 Yuanmingyuan West Road, Haidian District, Beijing 100193 ,","place":["China"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0002-0400-9147","authenticated-orcid":false,"given":"Teiji","family":"Sota","sequence":"additional","affiliation":[{"name":"Kyoto University Department of Zoology, Graduate School of Science, , Kitashirakawa Oiwake-cho, Sakyo-ku, Kyoto 606-8502 ,","place":["Japan"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0002-8620-0446","authenticated-orcid":false,"given":"Wanzhi","family":"Cai","sequence":"additional","affiliation":[{"name":"State Key Laboratory of Agricultural and Forestry Biosecurity, MOA Key Lab of Pest Monitoring and Green Management, College of Plant Protection, China Agricultural University , 2 Yuanmingyuan West Road, Haidian District, Beijing 100193 ,","place":["China"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0001-8590-1753","authenticated-orcid":false,"given":"Hu","family":"Li","sequence":"additional","affiliation":[{"name":"State Key Laboratory of Agricultural and Forestry Biosecurity, MOA Key Lab of Pest Monitoring and Green Management, College of Plant Protection, China Agricultural University , 2 Yuanmingyuan West Road, Haidian District, Beijing 100193 ,","place":["China"]}],"role":[{"vocabulary":"crossref","role":"author"}]}],"member":"286","published-online":{"date-parts":[[2026,8,18]]},"reference":[{"key":"2026090711121380500_bib1","doi-asserted-by":"crossref","first-page":"103621","DOI":"10.1016\/j.gloplacha.2021.103621","article-title":"End-Permian terrestrial disturbance followed by the complete plant devastation, and the vegetation proto-recovery in the earliest-Triassic recorded in coastal sea sediments","volume":"205","author":"Aftabuzzaman","year":"2021","journal-title":"Glob. Planet. Change"},{"key":"2026090711121380500_bib2","doi-asserted-by":"crossref","first-page":"420","DOI":"10.1038\/s41587-019-0036-z","article-title":"SignalP 5.0 improves signal peptide predictions using deep neural networks","volume":"37","author":"Almagro\u00a0Armenteros","year":"2019","journal-title":"Nat. Biotechnol."},{"key":"2026090711121380500_bib3","first-page":"34","article-title":"The arroyo willow stem-galling sawfly","volume":"66","author":"Alpert","year":"2020","journal-title":"Euura lasiolepis (Hymenoptera: Tenthredinidae): Am. Entomol"},{"key":"2026090711121380500_bib4","doi-asserted-by":"crossref","first-page":"6207","DOI":"10.1074\/jbc.M109874200","article-title":"Trialysin, a novel pore-forming protein from saliva of hematophagous insects activated by limited proteolysis","volume":"277","author":"Amino","year":"2002","journal-title":"J. Biol. Chem."},{"key":"2026090711121380500_bib5","doi-asserted-by":"crossref","first-page":"2473","DOI":"10.1139\/z93-343","article-title":"Classification, phylogeny, and zoogeography of the pond skater genus Gerris Fabricius (Hemiptera: gerridae)","volume":"71","author":"Andersen","year":"1993","journal-title":"Can. J. Zool."},{"key":"2026090711121380500_bib6","article-title":"FastQC: a quality control tool for high throughput sequence data. 2010","author":"Andrews","year":"2010","journal-title":"Babraham Bioinformatics"},{"key":"2026090711121380500_bib7","doi-asserted-by":"crossref","first-page":"833","DOI":"10.1016\/0022-1910(83)90149-X","article-title":"Haemolytic factor from the crop of Rhodnius prolixus: evidence and partial characterization","volume":"29","author":"Azambuja","year":"1983","journal-title":"J. Insect Physiol."},{"key":"2026090711121380500_bib8","doi-asserted-by":"crossref","first-page":"91","DOI":"10.1242\/jcs.s2-83.329.91","article-title":"The morphology and physiology of the salivary glands of Hemiptera-Heteroptera","volume":"s2-83","author":"Baptist","year":"1941","journal-title":"J. Cell Sci."},{"key":"2026090711121380500_bib9","doi-asserted-by":"crossref","first-page":"2503","DOI":"10.1038\/s41467-021-22792-y","article-title":"Size, microhabitat, and loss of larval feeding drive cranial diversification in frogs","volume":"12","author":"Bardua","year":"2021","journal-title":"Nat. Commun."},{"key":"2026090711121380500_bib10","doi-asserted-by":"crossref","first-page":"1964","DOI":"10.1093\/molbev\/msz125","article-title":"Many options, few solutions: over 60 My snakes converged on a few optimal venom formulations","volume":"36","author":"Barua","year":"2019","journal-title":"Mol. Biol. Evol."},{"key":"2026090711121380500_bib11","doi-asserted-by":"crossref","first-page":"5138","DOI":"10.1016\/j.cub.2021.09.028","article-title":"Feeding ecology has shaped the evolution of modern sharks","volume":"31","author":"Bazzi","year":"2021","journal-title":"Curr. Biol."},{"key":"2026090711121380500_bib12","doi-asserted-by":"crossref","first-page":"2021","DOI":"10.1111\/brv.12431","article-title":"Pterosaur dietary hypotheses: a review of ideas and approaches","volume":"93","author":"Bestwick","year":"2018","journal-title":"Biol. Rev."},{"key":"2026090711121380500_bib13","doi-asserted-by":"crossref","first-page":"717","DOI":"10.1111\/j.0014-3820.2003.tb00285.x","article-title":"Testing for phylogenetic signal in comparative data: behavioral traits are more labile","volume":"57","author":"Blomberg","year":"2003","journal-title":"Evolution"},{"key":"2026090711121380500_bib14","doi-asserted-by":"crossref","first-page":"e1006650","DOI":"10.1371\/journal.pcbi.1006650","article-title":"BEAST 2.5: an advanced software platform for Bayesian evolutionary analysis","volume":"15","author":"Bouckaert","year":"2019","journal-title":"PLoS Comput. Biol."},{"key":"2026090711121380500_bib15","doi-asserted-by":"crossref","first-page":"366","DOI":"10.1038\/s41592-021-01101-x","article-title":"Sensitive protein alignments at tree-of-life scale using DIAMOND","volume":"18","author":"Buchfink","year":"2021","journal-title":"Nat. Methods"},{"key":"2026090711121380500_bib16","doi-asserted-by":"crossref","first-page":"164","DOI":"10.1016\/j.ibmb.2006.11.005","article-title":"An insight into the sialome of Anopheles funestus reveals an emerging pattern in anopheline salivary protein families","volume":"37","author":"Calvo","year":"2007","journal-title":"Insect Biochem. Mol. Biol."},{"key":"2026090711121380500_bib17","doi-asserted-by":"crossref","first-page":"1735","DOI":"10.1038\/s41559-024-02485-7","article-title":"Sugar assimilation underlying dietary evolution of Neotropical bats","volume":"8","author":"Camacho","year":"2024","journal-title":"Nat. Ecol. Evol."},{"key":"2026090711121380500_bib18","doi-asserted-by":"crossref","first-page":"86","DOI":"10.1016\/j.cois.2020.07.006","article-title":"Extraoral digestion: outsourcing the role of the hemipteran midgut","volume":"41","author":"Cant\u00f3n","year":"2020","journal-title":"Curr. Opin. Insect Sci."},{"key":"2026090711121380500_bib19","doi-asserted-by":"crossref","first-page":"R647","DOI":"10.1016\/j.cub.2017.05.091","article-title":"Evolution: gene co-option underpins venom protein evolution","volume":"27","author":"Casewell","year":"2017","journal-title":"Curr. Biol."},{"key":"2026090711121380500_bib20","doi-asserted-by":"crossref","first-page":"884","DOI":"10.1093\/bioinformatics\/bty560","article-title":"Fastp: an ultra-fast all-in-one FASTQ preprocessor","volume":"34","author":"Chen","year":"2018","journal-title":"Bioinformatics"},{"key":"2026090711121380500_bib21","doi-asserted-by":"crossref","first-page":"58","DOI":"10.1038\/s41586-019-1302-4","article-title":"Insect egg size and shape evolve with ecology but not developmental rate","volume":"571","author":"Church","year":"2019","journal-title":"Nature"},{"key":"2026090711121380500_bib22","doi-asserted-by":"crossref","first-page":"1367","DOI":"10.1038\/nbt.1511","article-title":"MaxQuant enables high peptide identification rates, individualized p.p.b.-range mass accuracies and proteome-wide protein quantification","volume":"26","author":"Cox","year":"2008","journal-title":"Nat. Biotechnol."},{"key":"2026090711121380500_bib23","doi-asserted-by":"crossref","first-page":"345","DOI":"10.1186\/s12915-025-02446-x","article-title":"A foul odor from the mesozoic: early evolution of scent gland defense in pentatomomorphan true bugs (Hemiptera: heteroptera)","volume":"23","author":"Dai","year":"2025","journal-title":"BMC Biol."},{"key":"2026090711121380500_bib24","doi-asserted-by":"crossref","first-page":"425","DOI":"10.3390\/toxins11070425","article-title":"The dual \u03b1-amidation system in scorpion venom glands","volume":"11","author":"Delgado-Prudencio","year":"2019","journal-title":"Toxins"},{"key":"2026090711121380500_bib25","doi-asserted-by":"crossref","unstructured":"Drukewitz S.H., Bokelmann L., Undheim E.A.B., von\u00a0Reumont B.M. \u00a02019. Toxins from scratch? Diverse, multimodal gene origins in the predatory robber fly Dasypogon diadema indicate a dynamic venom evolution in dipteran insects. Gigascience. 8:1\u201313.","DOI":"10.1093\/gigascience\/giz081"},{"key":"2026090711121380500_bib26","doi-asserted-by":"crossref","first-page":"699","DOI":"10.1371\/journal.pbio.0040088","article-title":"Relaxed phylogenetics and dating with confidence","volume":"4","author":"Drummond","year":"2006","journal-title":"PLoS Biol."},{"key":"2026090711121380500_bib27","doi-asserted-by":"crossref","first-page":"20202888","DOI":"10.1098\/rspb.2020.2888","article-title":"Mammalian intestinal allometry, phylogeny, trophic level and climate","volume":"288","author":"Duque-Correa","year":"2021","journal-title":"Proc. R. Soc. B: Biol. Sci."},{"key":"2026090711121380500_bib28","first-page":"253","article-title":"Birds as predators of true bugs (Heteroptera) in different habitats","volume":"58","author":"Exnerov\u00e1","year":"2003","journal-title":"Biol. Sect. Zool."},{"key":"2026090711121380500_bib29","doi-asserted-by":"crossref","first-page":"65","DOI":"10.1146\/annurev-ento-011624-013339","article-title":"How insect exocrine glands work","volume":"70","author":"Foster","year":"2025","journal-title":"Annu. Rev. Entomol."},{"key":"2026090711121380500_bib30","doi-asserted-by":"crossref","first-page":"1042","DOI":"10.1111\/j.1523-1739.2010.01455.x","article-title":"Selectivity in mammalian extinction risk and threat types: a new measure of phylogenetic signal strength in binary traits","volume":"24","author":"Fritz","year":"2010","journal-title":"Conserv. Biol."},{"key":"2026090711121380500_bib31","doi-asserted-by":"crossref","first-page":"3150","DOI":"10.1093\/bioinformatics\/bts565","article-title":"CD-HIT: accelerated for clustering the next-generation sequencing data","volume":"28","author":"Fu","year":"2012","journal-title":"Bioinformatics"},{"key":"2026090711121380500_bib32","first-page":"464","article-title":"Proteotranscriptomic analysis and toxicity assay suggest the functional distinction between venom gland chambers in twin-spotted assassin bug","volume":"11","author":"Gao","year":"2022","journal-title":"Platymeris Biguttatus. Biology"},{"key":"2026090711121380500_bib33","doi-asserted-by":"crossref","first-page":"313","DOI":"10.1046\/j.1365-313X.1997.11020313.x","article-title":"The C-terminal HDEL sequence is sufficient for retention of secretory proteins in the endoplasmic reticulum (ER) but promotes vacuolar targeting of proteins that escape the ER","volume":"11","author":"Gomord","year":"1997","journal-title":"Plant J."},{"key":"2026090711121380500_bib34","doi-asserted-by":"crossref","first-page":"1494","DOI":"10.1038\/nprot.2013.084","article-title":"De novo transcript sequence reconstruction from RNA-seq using the Trinity platform for reference generation and analysis","volume":"8","author":"Haas","year":"2013","journal-title":"Nat. Protoc."},{"key":"2026090711121380500_bib35","doi-asserted-by":"crossref","first-page":"129","DOI":"10.1093\/bioinformatics\/btm538","article-title":"GEIGER: investigating evolutionary radiations","volume":"24","author":"Harmon","year":"2008","journal-title":"Bioinformatics"},{"key":"2026090711121380500_bib36","doi-asserted-by":"crossref","DOI":"10.1073\/pnas.2015579118","article-title":"Phylogenetically diverse diets favor more complex venoms in North American pitvipers","volume":"118","author":"Holding","year":"2021","journal-title":"Proc. Natl Acad. Sci. USA"},{"key":"2026090711121380500_bib37","doi-asserted-by":"crossref","first-page":"257","DOI":"10.1038\/s42003-024-05961-9","article-title":"Salivary proteins potentially derived from horizontal gene transfer are critical for salivary sheath formation and other feeding processes","volume":"7","author":"Huang","year":"2024","journal-title":"Commun. Biol."},{"key":"2026090711121380500_bib38","doi-asserted-by":"crossref","first-page":"737","DOI":"10.1038\/s41467-023-36403-5","article-title":"Planthopper salivary sheath protein LsSP1 contributes to manipulation of rice plant defenses","volume":"14","author":"Huang","year":"2023","journal-title":"Nat. Commun."},{"key":"2026090711121380500_bib39","doi-asserted-by":"crossref","first-page":"12775","DOI":"10.1073\/pnas.1815820115","article-title":"Phylogenomics and the evolution of hemipteroid insects","volume":"115","author":"Johnson","year":"2018","journal-title":"Proc. Natl. Acad. Sci. USA"},{"key":"2026090711121380500_bib40","doi-asserted-by":"crossref","first-page":"275","DOI":"10.1093\/bioinformatics\/8.3.275","article-title":"The rapid generation of mutation data matrices from protein sequences","volume":"8","author":"Jones","year":"1992","journal-title":"Bioinformatics"},{"key":"2026090711121380500_bib41","doi-asserted-by":"crossref","first-page":"1236","DOI":"10.1093\/bioinformatics\/btu031","article-title":"InterProScan 5: genome-scale protein function classification","volume":"30","author":"Jones","year":"2014","journal-title":"Bioinformatics"},{"key":"2026090711121380500_bib42","doi-asserted-by":"crossref","first-page":"50","DOI":"10.1111\/j.1096-0031.2011.00365.x","article-title":"Molecular phylogeny of the plant bugs (Heteroptera: miridae) and the evolution of feeding habits","volume":"28","author":"Jung","year":"2012","journal-title":"Cladistics"},{"key":"2026090711121380500_bib43","doi-asserted-by":"crossref","first-page":"1160","DOI":"10.1093\/bib\/bbx108","article-title":"MAFFT online service: multiple sequence alignment, interactive sequence choice and visualization","volume":"20","author":"Katoh","year":"2019","journal-title":"Brief. Bioinform."},{"key":"2026090711121380500_bib44","doi-asserted-by":"crossref","first-page":"386","DOI":"10.1126\/science.abb9303","article-title":"Convergent evolution of pain-inducing defensive venom components in spitting cobras","volume":"371","author":"Kazandjian","year":"2021","journal-title":"Science"},{"key":"2026090711121380500_bib45","doi-asserted-by":"crossref","first-page":"1","DOI":"10.1111\/syen.12512","article-title":"Molecular phylogeny of Nabidae (Hemiptera: heteroptera: cimicomorpha): insight into relationships and reclassification with the proposal of the new tribe Stenonabini","volume":"47","author":"Kim","year":"2022","journal-title":"Syst. Entomol."},{"key":"2026090711121380500_bib46","doi-asserted-by":"crossref","first-page":"248","DOI":"10.1111\/cla.12435","article-title":"Phylogenetic relationships and revised classification of the true bug infraorder Dipsocoromorpha (Insecta: hemiptera: heteroptera)","volume":"37","author":"Knyshov","year":"2021","journal-title":"Cladistics"},{"key":"2026090711121380500_bib47","doi-asserted-by":"crossref","first-page":"229","DOI":"10.1186\/s12915-023-01656-5","article-title":"Prevalent bee venom genes evolved before the aculeate stinger and eusociality","volume":"21","author":"Koludarov","year":"2023","journal-title":"BMC Biol."},{"key":"2026090711121380500_bib48","doi-asserted-by":"crossref","first-page":"6001","DOI":"10.1038\/s41467-021-26285-w","article-title":"Multiple evolutionary origins and losses of tooth complexity in squamates","volume":"12","author":"Lafuma","year":"2021","journal-title":"Nat. Commun."},{"key":"2026090711121380500_bib49","doi-asserted-by":"crossref","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":"2026090711121380500_bib50","doi-asserted-by":"crossref","first-page":"323","DOI":"10.1186\/1471-2105-12-323","article-title":"RSEM: accurate transcript quantification from RNA-Seq data with or without a reference genome","volume":"12","author":"Li","year":"2011","journal-title":"BMC Bioinf."},{"key":"2026090711121380500_bib51","doi-asserted-by":"crossref","first-page":"20171223","DOI":"10.1098\/rspb.2017.1223","article-title":"Mitochondrial phylogenomics of Hemiptera reveals adaptive innovations driving the diversification of true bugs","volume":"284","author":"Li","year":"2017","journal-title":"Proc. R. Soc. B Biol. Sci."},{"key":"2026090711121380500_bib52","doi-asserted-by":"crossref","first-page":"810","DOI":"10.1111\/syen.12357","article-title":"Higher-level phylogeny and evolutionary history of Pentatomomorpha (Hemiptera: heteroptera) inferred from mitochondrial genome sequences","volume":"44","author":"Liu","year":"2019","journal-title":"Syst. Entomol."},{"key":"2026090711121380500_bib53","doi-asserted-by":"crossref","first-page":"msac082","DOI":"10.1093\/molbev\/msac082","article-title":"Venom gene sequence diversity and expression jointly shape diet adaptation in pitvipers","volume":"39","author":"Mason","year":"2022","journal-title":"Mol. Biol. Evol."},{"key":"2026090711121380500_bib54","doi-asserted-by":"crossref","first-page":"335","DOI":"10.1016\/0022-1910(82)90045-2","article-title":"Absorption of fluid from the anterior midgut in Rhodnius","volume":"28","author":"Michael\u00a0Barrett","year":"1982","journal-title":"J. Insect Physiol."},{"key":"2026090711121380500_bib55","first-page":"183","article-title":"The saliva of Hemiptera","volume":"9","author":"Miles","year":"1972","journal-title":"Adv. In Insect Phys."},{"key":"2026090711121380500_bib56","doi-asserted-by":"crossref","first-page":"1011","DOI":"10.1007\/BF01933942","article-title":"The accessory salivary gland as the source of water in the saliva of Hemiptera: heteroptera","volume":"32","author":"Miles","year":"1976","journal-title":"Experientia"},{"key":"2026090711121380500_bib57","doi-asserted-by":"crossref","first-page":"1188","DOI":"10.1093\/molbev\/mst024","article-title":"Ultrafast approximation for phylogenetic bootstrap","volume":"30","author":"Minh","year":"2013","journal-title":"Mol. Biol. Evol."},{"key":"2026090711121380500_bib58","doi-asserted-by":"crossref","first-page":"763","DOI":"10.1126\/science.1257570","article-title":"Phylogenomics resolves the timing and pattern of insect evolution","volume":"346","author":"Misof","year":"2014","journal-title":"Science"},{"key":"2026090711121380500_bib59","first-page":"1","article-title":"Comparative morphology of alimentary organs of Heteroptera, with the phylogenetic consideration","volume":"2","author":"Miyamoto","year":"1961","journal-title":"Sieboldia"},{"key":"2026090711121380500_bib60","article-title":"Adaptive evolution of distinct prey-specific toxin genes in rear-fanged snake venom","volume":"285","author":"Modahl","year":"2018","journal-title":"Proc. R. Soc. B: Biol. Sci."},{"key":"2026090711121380500_bib61","doi-asserted-by":"crossref","first-page":"646","DOI":"10.1093\/bioinformatics\/17.7.646","article-title":"Evaluation of methods for the prediction of membrane spanning regions","volume":"17","author":"M\u00f6ller","year":"2001","journal-title":"Bioinformatics"},{"key":"2026090711121380500_bib62","doi-asserted-by":"crossref","first-page":"599","DOI":"10.1038\/416599a","article-title":"Caterpillar saliva beats plant defences","volume":"416","author":"Musser","year":"2002","journal-title":"Nature"},{"key":"2026090711121380500_bib63","doi-asserted-by":"crossref","first-page":"268","DOI":"10.1093\/molbev\/msu300","article-title":"IQ-TREE: a fast and effective stochastic algorithm for estimating maximum-likelihood phylogenies","volume":"32","author":"Nguyen","year":"2015","journal-title":"Mol. Biol. Evol."},{"key":"2026090711121380500_bib64","article-title":"Caper: comparative analyses of phylogenetics and evolution in R","author":"Orme","year":"2018"},{"key":"2026090711121380500_bib65","doi-asserted-by":"crossref","first-page":"877","DOI":"10.1038\/44766","article-title":"Inferring the historical patterns of biological evolution","volume":"401","author":"Pagel","year":"1999","journal-title":"Nature"},{"key":"2026090711121380500_bib66","doi-asserted-by":"crossref","first-page":"289","DOI":"10.1093\/bioinformatics\/btg412","article-title":"APE: analyses of phylogenetics and evolution in R language","volume":"20","author":"Paradis","year":"2004","journal-title":"Bioinformatics"},{"key":"2026090711121380500_bib67","doi-asserted-by":"crossref","first-page":"4257","DOI":"10.1111\/mec.14859","article-title":"Venom gland size and venom complexity-essential trophic adaptations of venomous predators: a case study using spiders","volume":"27","author":"Pek\u00e1r","year":"2018","journal-title":"Mol. Ecol."},{"key":"2026090711121380500_bib68","doi-asserted-by":"crossref","first-page":"483","DOI":"10.1016\/0968-0004(90)90303-S","article-title":"The retention signal for soluble proteins of the endoplasmic reticulum","volume":"15","author":"Pelham","year":"1990","journal-title":"Trends Biochem. Sci"},{"key":"2026090711121380500_bib69","doi-asserted-by":"crossref","first-page":"401","DOI":"10.1186\/s12864-016-2755-6","article-title":"Dietary breadth is positively correlated with venom complexity in cone snails","volume":"17","author":"Phuong","year":"2016","journal-title":"Bmc Genom [Electron Res]"},{"key":"2026090711121380500_bib70","doi-asserted-by":"crossref","first-page":"375","DOI":"10.1007\/s11829-021-09817-5","article-title":"The localization of phytohormones within the gall-inducing insect Eurosta solidaginis (Diptera: tephritidae)","volume":"15","author":"Ponce","year":"2021","journal-title":"Arthropod. Plant. Interact."},{"key":"2026090711121380500_bib71","article-title":"FigTree v1.4.4. A graphical viewer of phylogenetic trees","author":"Rambaut","year":"2018"},{"key":"2026090711121380500_bib72","doi-asserted-by":"crossref","first-page":"217","DOI":"10.1111\/j.2041-210X.2011.00169.x","article-title":"phytools: an R package for phylogenetic comparative biology (and other things)","volume":"3","author":"Revell","year":"2012","journal-title":"Methods Ecol. Evol."},{"key":"2026090711121380500_bib73","first-page":"1","article-title":"An insight into the sialomes of bloodsucking Heteroptera","volume":"2012","author":"Ribeiro","year":"2012","journal-title":"Psyche"},{"key":"2026090711121380500_bib74","doi-asserted-by":"crossref","first-page":"61","DOI":"10.1016\/j.cois.2017.01.002","article-title":"Genomics of Lepidoptera saliva reveals function in herbivory","volume":"19","author":"Rivera-Vega","year":"2017","journal-title":"Curr. Opin. Insect Sci."},{"key":"2026090711121380500_bib75","doi-asserted-by":"crossref","first-page":"36","DOI":"10.1111\/syen.12514","article-title":"Opening Pandora\u2019s box: molecular phylogeny of the stink bugs (Hemiptera: heteroptera: pentatomidae) reveals great incongruences in the current classification","volume":"47","author":"Roca-Cusachs","year":"2022","journal-title":"Syst. Entomol."},{"key":"2026090711121380500_bib76","doi-asserted-by":"crossref","first-page":"103739","DOI":"10.1016\/j.ibmb.2022.103739","article-title":"Effect of Triatoma infestans saliva on mouse immune system cells: the role of the pore-forming salivary protein trialysin","volume":"143","author":"Rocha","year":"2022","journal-title":"Insect Biochem. Mol. Biol."},{"key":"2026090711121380500_bib77","doi-asserted-by":"crossref","first-page":"3287","DOI":"10.1038\/s41467-023-38924-5","article-title":"Coordinated adaptations define the ontogenetic shift from worm- to fish-hunting in a venomous cone snail","volume":"14","author":"Rogalski","year":"2023","journal-title":"Nat. Commun."},{"key":"2026090711121380500_bib78","doi-asserted-by":"crossref","first-page":"107","DOI":"10.1146\/annurev.en.07.010162.000543","article-title":"Chemical defenses of arthropods","volume":"7","author":"Roth","year":"1962","journal-title":"Annu. Rev. Entomol."},{"key":"2026090711121380500_bib79","doi-asserted-by":"crossref","first-page":"676","DOI":"10.1038\/nmeth.2019","article-title":"Fiji: an open-source platform for biological-image analysis","volume":"9","author":"Schindelin","year":"2012","journal-title":"Nat. Methods"},{"key":"2026090711121380500_bib80","volume-title":"The true bugs of the world (Hemiptera: Heteroptera): classification and natural history","author":"Schuh","year":"2020"},{"key":"2026090711121380500_bib81","first-page":"15","article-title":"Mouthparts and feeding","volume-title":"The insects: structure and function","author":"Simpson","year":"2013"},{"key":"2026090711121380500_bib82","doi-asserted-by":"crossref","first-page":"249","DOI":"10.1038\/s41467-023-35794-9","article-title":"Micro and macroevolution of sea anemone venom phenotype","volume":"14","author":"Smith","year":"2023","journal-title":"Nat. Commun."},{"key":"2026090711121380500_bib83","first-page":"77","article-title":"The morphology ofthe salivary glands of terrestrial Heteroptera (Geocorisae) and its bearing on classification","volume":"98","author":"Southwood","year":"1955","journal-title":"Tijdschr. Entomol."},{"key":"2026090711121380500_bib84","doi-asserted-by":"crossref","first-page":"13","DOI":"10.1007\/s00265-010-1037-6","article-title":"A brief guide to model selection, multimodel inference and model averaging in behavioural ecology using Akaike\u2019s information criterion","volume":"65","author":"Symonds","year":"2011","journal-title":"Behav. Ecol. Sociobiol."},{"key":"2026090711121380500_bib85","doi-asserted-by":"crossref","first-page":"100931","DOI":"10.1016\/j.asd.2020.100931","article-title":"Evolutionary trends of digestion and absorption in the major insect orders","volume":"56","author":"Terra","year":"2020","journal-title":"Arthropod Struct. Dev."},{"key":"2026090711121380500_bib86","doi-asserted-by":"crossref","first-page":"20201393","DOI":"10.1098\/rspb.2020.1393","article-title":"Eye size and investment in frogs and toads correlate with adult habitat, activity pattern and breeding ecology","volume":"287","author":"Thomas","year":"2020","journal-title":"Proc. R. Soc. B: Biol. Sci."},{"key":"2026090711121380500_bib87","doi-asserted-by":"crossref","first-page":"W232","DOI":"10.1093\/nar\/gkw256","article-title":"W-IQ-TREE: a fast online phylogenetic tool for maximum likelihood analysis","volume":"44","author":"Trifinopoulos","year":"2016","journal-title":"Nucleic. Acids. Res."},{"key":"2026090711121380500_bib88","doi-asserted-by":"crossref","first-page":"2301","DOI":"10.1038\/nprot.2016.136","article-title":"The MaxQuant computational platform for mass spectrometry-based shotgun proteomics","volume":"11","author":"Tyanova","year":"2016","journal-title":"Nat. Protoc."},{"key":"2026090711121380500_bib89","doi-asserted-by":"crossref","first-page":"3215","DOI":"10.1007\/s00018-018-2768-1","article-title":"Giant fish-killing water bug reveals ancient and dynamic venom evolution in Heteroptera","volume":"75","author":"Walker","year":"2018","journal-title":"Cell. Mol. Life Sci."},{"key":"2026090711121380500_bib90","doi-asserted-by":"crossref","first-page":"552","DOI":"10.1074\/mcp.M116.063321","article-title":"Melt with this kiss: paralyzing and liquefying venom of the assassin bug Pristhesancus plagipennis (Hemiptera: reduviidae)","volume":"16","author":"Walker","year":"2017","journal-title":"Mol. Cell. Proteomics"},{"key":"2026090711121380500_bib91","doi-asserted-by":"crossref","first-page":"755","DOI":"10.1038\/s41467-018-03091-5","article-title":"The assassin bug Pristhesancus plagipennis produces two distinct venoms in separate gland lumens","volume":"9","author":"Walker","year":"2018","journal-title":"Nat. Commun."},{"key":"2026090711121380500_bib92","doi-asserted-by":"crossref","first-page":"43","DOI":"10.3390\/toxins8020043","article-title":"Venoms of heteropteran insects: a treasure trove of diverse pharmacological toolkits","volume":"8","author":"Walker","year":"2016","journal-title":"Toxins"},{"key":"2026090711121380500_bib93","doi-asserted-by":"crossref","first-page":"897","DOI":"10.1016\/B978-0-12-374144-8.00237-X","article-title":"Salivary glands","volume-title":"Encyclopedia of insects","author":"Walker","year":"2009","edition":"2nd ed"},{"key":"2026090711121380500_bib94","doi-asserted-by":"crossref","first-page":"339","DOI":"10.1111\/syen.12465","article-title":"Diversification of true water bugs revealed by transcriptome-based phylogenomics","volume":"46","author":"Wang","year":"2021","journal-title":"Syst. Entomol."},{"key":"2026090711121380500_bib95","doi-asserted-by":"crossref","first-page":"msad221","DOI":"10.1093\/molbev\/msad221","article-title":"Horizontally transferred salivary protein promotes insect feeding by suppressing ferredoxin-mediated plant defenses","volume":"40","author":"Wang","year":"2023","journal-title":"Mol. Biol. Evol."},{"key":"2026090711121380500_bib96","doi-asserted-by":"crossref","first-page":"42","DOI":"10.1111\/cla.12232","article-title":"When did the ancestor of true bugs become stinky? Disentangling the phylogenomics of Hemiptera\u2013Heteroptera","volume":"35","author":"Wang","year":"2019","journal-title":"Cladistics"},{"key":"2026090711121380500_bib97","doi-asserted-by":"crossref","first-page":"287","DOI":"10.1016\/j.ympev.2009.05.039","article-title":"Molecular phylogeny of the assassin bugs (Hemiptera: reduviidae), based on mitochondrial and nuclear ribosomal genes","volume":"53","author":"Weirauch","year":"2009","journal-title":"Mol. Phylogenet. Evol."},{"key":"2026090711121380500_bib98","doi-asserted-by":"crossref","first-page":"487","DOI":"10.1146\/annurev-ento-120709-144833","article-title":"Systematics and evolution of heteroptera: 25 years of progress","volume":"56","author":"Weirauch","year":"2011","journal-title":"Annu. Rev. Entomol."},{"key":"2026090711121380500_bib99","doi-asserted-by":"crossref","first-page":"67","DOI":"10.1111\/cla.12233","article-title":"Revisiting habitat and lifestyle transitions in Heteroptera (Insecta: hemiptera): insights from a combined morphological and molecular phylogeny","volume":"35","author":"Weirauch","year":"2019","journal-title":"Cladistics"},{"key":"2026090711121380500_bib100","doi-asserted-by":"crossref","first-page":"8370","DOI":"10.1038\/ncomms9370","article-title":"Herbivory increases diversification across insect clades","volume":"6","author":"Wiens","year":"2015","journal-title":"Nat. Commun."},{"key":"2026090711121380500_bib101","doi-asserted-by":"crossref","first-page":"W587","DOI":"10.1093\/nar\/gkad359","article-title":"Tree visualization by one table (tvBOT): a web application for visualizing, modifying and annotating phylogenetic trees","volume":"51","author":"Xie","year":"2023","journal-title":"Nucleic. Acids. Res."},{"key":"2026090711121380500_bib102","doi-asserted-by":"crossref","first-page":"403","DOI":"10.1111\/cla.12501","article-title":"Diversification of the phytophagous lineages of true bugs (Insecta: hemiptera: heteroptera) shortly after that of the flowering plants","volume":"38","author":"Ye","year":"2022","journal-title":"Cladistics"},{"key":"2026090711121380500_bib103","doi-asserted-by":"crossref","first-page":"411","DOI":"10.1111\/1744-7917.12955","article-title":"Comparative analyses of the venom components in the salivary gland transcriptomes and saliva proteomes of some heteropteran insects","volume":"29","author":"Yoon","year":"2022","journal-title":"Insect Sci."},{"key":"2026090711121380500_bib104","doi-asserted-by":"crossref","first-page":"2777","DOI":"10.1093\/molbev\/msaa133","article-title":"Venom systems as models for studying the origin and regulation of evolutionary novelties","volume":"37","author":"Zancolli","year":"2020","journal-title":"Mol. Biol. Evol."}],"container-title":["Systematic Biology"],"original-title":[],"language":"en","link":[{"URL":"https:\/\/academic.oup.com\/sysbio\/advance-article-pdf\/doi\/10.1093\/sysbio\/syag067\/70685015\/syag067.pdf","content-type":"application\/pdf","content-version":"am","intended-application":"syndication"},{"URL":"https:\/\/academic.oup.com\/sysbio\/advance-article-pdf\/doi\/10.1093\/sysbio\/syag067\/70685015\/syag067.pdf","content-type":"unspecified","content-version":"vor","intended-application":"similarity-checking"}],"deposited":{"date-parts":[[2026,9,7]],"date-time":"2026-09-07T15:12:29Z","timestamp":1788793949000},"score":1,"resource":{"primary":{"URL":"https:\/\/academic.oup.com\/sysbio\/advance-article\/doi\/10.1093\/sysbio\/syag067\/8763810"}},"subtitle":[],"editor":[{"given":"Fabien","family":"Condamine","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"editor"}]}],"short-title":[],"issued":{"date-parts":[[2026,8,18]]},"references-count":104,"URL":"https:\/\/doi.org\/10.1093\/sysbio\/syag067","relation":{},"ISSN":["1063-5157","1076-836X"],"issn-type":[{"value":"1063-5157","type":"print"},{"value":"1076-836X","type":"electronic"}],"subject":[],"published":{"date-parts":[[2026,8,18]]},"article-number":"syag067"}}