{"status":"ok","message-type":"work","message-version":"1.0.0","message":{"indexed":{"date-parts":[[2026,9,11]],"date-time":"2026-09-11T05:06:45Z","timestamp":1789103205840,"version":"build-2803163510"},"reference-count":85,"publisher":"MDPI AG","issue":"9","license":[{"start":{"date-parts":[[2026,9,9]],"date-time":"2026-09-09T00:00:00Z","timestamp":1788912000000},"content-version":"vor","delay-in-days":0,"URL":"https:\/\/creativecommons.org\/licenses\/by\/4.0\/"}],"funder":[{"DOI":"10.13039\/100000001","name":"US National Science Foundation","doi-asserted-by":"crossref","award":["DEB-2244871"],"award-info":[{"award-number":["DEB-2244871"]}],"id":[{"id":"10.13039\/100000001","id-type":"DOI","asserted-by":"crossref"}]},{"award":["DEB-2244871"],"award-info":[{"award-number":["DEB-2244871"]}],"id":[{"id":"https:\/\/ror.org\/021nxhr62","id-type":"ROR","asserted-by":"publisher"}]}],"content-domain":{"domain":[],"crossmark-restriction":false},"short-container-title":["Insects"],"abstract":"<jats:p>Vector-borne plant pathogens exhibit a remarkable capacity to colonize distantly related hosts, reflecting long-term ecological and evolutionary interactions among pathogens, plants, and insect vectors. Phytoplasmas, which are obligate bacterial parasites transmitted by hemipteran insects, represent a model system for investigating how host\u2013pathogen associations diversify across ecological and evolutionary timescales. Despite extensive knowledge of phytoplasma diversity and vector relationships, the historical processes shaping these associations remain poorly understood. Here, we investigated the eco-evolutionary dynamics of global phytoplasma\u2013hemipteran associations using complementary cophylogenetic approaches that integrate phylogenetic relationships, divergence time estimates, and host association data. We tested the hypothesis that community assembly of phytoplasma\u2013vector networks over the last ~300 million years has been driven by multiple processes, including host switching, lineage duplication, and cospeciation, with additional influences from environmental change and historical biogeography. Our analyses revealed limited evidence for cospeciation across the evolution of the associations. Instead, duplication, host switching, and lineage losses were the predominant processes shaping current associations, with more than 90 inferred incongruent links between phytoplasmas and their insect hosts. A significant cophylogenetic signal was detected only between Psyllidae and phytoplasma group 16SrX, suggesting a history of relative ecological stability or specialization following their concurrent emergence in the Late Cretaceous. Reconstruction of the origins of phytoplasma associations with particular families of insect vectors indicates that the common ancestor of modern Cicadellidae may have already acquired an association with phytoplasmas, but that associations with other vector families were acquired more recently. These findings indicate that phytoplasma\u2013vector associations are largely dynamic and shaped by ecological opportunity rather than strict codiversification. Integrating evolutionary history with ecological interactions provides a framework to better understand the emergence, persistence, and potential future spread of vector-borne plant diseases.<\/jats:p>","DOI":"10.3390\/insects17090942","type":"journal-article","created":{"date-parts":[[2026,9,9]],"date-time":"2026-09-09T09:24:02Z","timestamp":1788945842000},"page":"942","update-policy":"https:\/\/doi.org\/10.3390\/mdpi_crossmark_policy","source":"Crossref","is-referenced-by-count":0,"title":["First Cophylogenetic Reconstruction of Hemipteran Insect\u2013Phytoplasma Associations Reveals Eco-Evolutionary Dynamics"],"prefix":"10.3390","volume":"17","author":[{"ORCID":"https:\/\/orcid.org\/0000-0003-1415-4097","authenticated-orcid":false,"given":"Valeria","family":"Trivellone","sequence":"first","affiliation":[{"name":"Illinois Natural History Survey, Prairie Research Institute, University of Illinois at Urbana-Champaign, Champaign, IL 61820, USA"}],"role":[{"vocabulary":"crossref","role":"author"},{"role":"conceptualization","vocabulary":"credit"},{"role":"methodology","vocabulary":"credit"},{"role":"formal-analysis","vocabulary":"credit"},{"role":"investigation","vocabulary":"credit"},{"role":"resources","vocabulary":"credit"},{"role":"data-curation","vocabulary":"credit"},{"role":"writing-review-editing","vocabulary":"credit"},{"role":"writing-original-draft","vocabulary":"credit"},{"role":"visualization","vocabulary":"credit"},{"role":"funding-acquisition","vocabulary":"credit"},{"role":"project-administration","vocabulary":"credit"}]},{"ORCID":"https:\/\/orcid.org\/0000-0003-4005-4305","authenticated-orcid":false,"given":"Christopher H.","family":"Dietrich","sequence":"additional","affiliation":[{"name":"Illinois Natural History Survey, Prairie Research Institute, University of Illinois at Urbana-Champaign, Champaign, IL 61820, USA"}],"role":[{"vocabulary":"crossref","role":"author"},{"role":"conceptualization","vocabulary":"credit"},{"role":"methodology","vocabulary":"credit"},{"role":"resources","vocabulary":"credit"},{"role":"writing-review-editing","vocabulary":"credit"},{"role":"supervision","vocabulary":"credit"},{"role":"funding-acquisition","vocabulary":"credit"}]}],"member":"1968","published-online":{"date-parts":[[2026,9,9]]},"reference":[{"key":"ref_1","doi-asserted-by":"crossref","first-page":"10","DOI":"10.1007\/s44297-025-00049-0","article-title":"Bacterial Vector-Borne Plant Diseases: Global Issues Caused by Three-Way Interactions","volume":"3","author":"Huang","year":"2025","journal-title":"Crop Health"},{"key":"ref_2","doi-asserted-by":"crossref","first-page":"1379","DOI":"10.1016\/j.molp.2020.08.010","article-title":"Bacterial Vector-Borne Plant Diseases: Unanswered Questions and Future Directions","volume":"13","author":"Huang","year":"2020","journal-title":"Mol. Plant"},{"key":"ref_3","doi-asserted-by":"crossref","unstructured":"Trivellone, V., Araujo, S., and Dietrich, C.H. (2025). Evolutionary Insights into Hemipteran Insect-Phytoplasma Associations: Phylogenetic Conservatism and Historical Ecology as Tools to Predict Future Associations. Phytoplasmas: Genomes, Plant Hosts and Vectors, CABI.","DOI":"10.1079\/9781800627031.0013"},{"key":"ref_4","doi-asserted-by":"crossref","first-page":"605","DOI":"10.1111\/nph.16092","article-title":"An Evolutionary Framework for Host Shifts\u2014Jumping Ships for Survival","volume":"224","author":"Thines","year":"2019","journal-title":"New Phytol."},{"key":"ref_5","doi-asserted-by":"crossref","unstructured":"Gardner, S.L., Brooks, D.R., Boeger, W.A., and Hoberg, E.P. (2023). An Evolutionary Pathway for Coping with Emerging Infectious Disease, Zea Books.","DOI":"10.32873\/unl.dc.zea.1504"},{"key":"ref_6","first-page":"185942","article-title":"Current View on Phytoplasma Genomes and Encoded Metabolism","volume":"1","author":"Kube","year":"2012","journal-title":"Sci. World J."},{"key":"ref_7","doi-asserted-by":"crossref","first-page":"27","DOI":"10.1038\/ng1277","article-title":"Reductive Evolution Suggested from the Complete Genome Sequence of a Plant-Pathogenic Phytoplasma","volume":"36","author":"Oshima","year":"2004","journal-title":"Nat. Genet."},{"key":"ref_8","doi-asserted-by":"crossref","unstructured":"Oshima, K., Maejima, K., and Namba, S. (2013). Genomic and Evolutionary Aspects of Phytoplasmas. Front. Microbiol., 4.","DOI":"10.3389\/fmicb.2013.00230"},{"key":"ref_9","doi-asserted-by":"crossref","unstructured":"Orlovskis, Z., and Hogenhout, S.A. (2016). A Bacterial Parasite Effector Mediates Insect Vector Attraction in Host Plants Independently of Developmental Changes. Front. Plant Sci., 7.","DOI":"10.3389\/fpls.2016.00885"},{"key":"ref_10","doi-asserted-by":"crossref","first-page":"106826","DOI":"10.1016\/j.ympev.2020.106826","article-title":"A Timetree for Phytoplasmas (Mollicutes) with New Insights on Patterns of Evolution and Diversification","volume":"149","author":"Cao","year":"2020","journal-title":"Mol. Phylogenet. Evol."},{"key":"ref_11","doi-asserted-by":"crossref","first-page":"555","DOI":"10.1111\/j.1574-6976.2011.00264.x","article-title":"Insects as Alternative Hosts for Phytopathogenic Bacteria","volume":"35","author":"Nadarasah","year":"2011","journal-title":"FEMS Microbiol. Rev."},{"key":"ref_12","doi-asserted-by":"crossref","first-page":"147","DOI":"10.1111\/j.1462-2920.2009.02056.x","article-title":"A Single Genetic Locus in the Phytopathogen Pantoea stewartii Enables Gut Colonization and Pathogenicity in an Insect Host","volume":"12","author":"Stavrinides","year":"2010","journal-title":"Environ. Microbiol."},{"key":"ref_13","doi-asserted-by":"crossref","first-page":"RP98992","DOI":"10.7554\/eLife.98992","article-title":"The Phytoplasma SAP54 Effector Acts as a Molecular Matchmaker for Leafhopper Vectors by Targeting Plant MADS-Box Factor SVP","volume":"13","author":"Orlovskis","year":"2025","journal-title":"eLife"},{"key":"ref_14","doi-asserted-by":"crossref","first-page":"144","DOI":"10.1016\/j.jip.2005.03.001","article-title":"Effect of Two Strains of Flavescence Dor\u00e9e Phytoplasma on the Survival and Fecundity of the Experimental Leafhopper Vector Euscelidius variegatus Kirschbaum","volume":"89","author":"Bressan","year":"2005","journal-title":"J. Invertebr. Pathol."},{"key":"ref_15","doi-asserted-by":"crossref","first-page":"e00042-18","DOI":"10.1128\/IAI.00042-18","article-title":"Two Phytoplasmas Elicit Different Responses in the Insect Vector Euscelidius variegatus Kirschbaum","volume":"86","author":"Galetto","year":"2018","journal-title":"Infect. Immun."},{"key":"ref_16","doi-asserted-by":"crossref","first-page":"3552","DOI":"10.1074\/jbc.M808334200","article-title":"Evf, a Virulence Factor Produced by the Drosophila Pathogen Erwinia carotovora, Is an S-Palmitoylated Protein with a New Fold That Binds to Lipid Vesicles","volume":"284","author":"Leulliot","year":"2009","journal-title":"J. Biol. Chem."},{"key":"ref_17","doi-asserted-by":"crossref","first-page":"299","DOI":"10.1093\/sysbio\/28.3.299","article-title":"Testing the Context and Extent of Host-Parasite Coevolution","volume":"28","author":"Brooks","year":"1979","journal-title":"Syst. Biol."},{"key":"ref_18","doi-asserted-by":"crossref","first-page":"347","DOI":"10.1111\/nph.12150","article-title":"Cospeciation vs Host-Shift Speciation: Methods for Testing, Evidence from Natural Associations and Relation to Coevolution","volume":"198","author":"Vienne","year":"2013","journal-title":"New Phytol."},{"key":"ref_19","doi-asserted-by":"crossref","first-page":"275","DOI":"10.1146\/annurev-ecolsys-102320-112823","article-title":"Cophylogenetic Methods to Untangle the Evolutionary History of Ecological Interactions","volume":"53","author":"Dismukes","year":"2022","journal-title":"Annu. Rev. Ecol. Evol. Syst."},{"key":"ref_20","doi-asserted-by":"crossref","first-page":"660","DOI":"10.2307\/2399219","article-title":"Historical Ecology: A New Approach to Studying the Evolution of Ecological Associations","volume":"72","author":"Brooks","year":"1985","journal-title":"Ann. Mo. Bot. Gard."},{"key":"ref_21","unstructured":"Page, R.D.M. (2003). Tangled Trees: Phylogeny, Cospeciation, and Coevolution, University of Chicago Press."},{"key":"ref_22","doi-asserted-by":"crossref","first-page":"137","DOI":"10.1093\/aesa\/saaa048","article-title":"Evolutionary Diversification in Insect Vector-Phytoplasma-Plant Associations","volume":"114","author":"Trivellone","year":"2021","journal-title":"Ann. Entomol. Soc. Am."},{"key":"ref_23","doi-asserted-by":"crossref","first-page":"e32910","DOI":"10.3897\/BDJ.7.e32910","article-title":"An Online Global Database of Hemiptera-Phytoplasma-Plant Biological Interactions","volume":"7","author":"Trivellone","year":"2019","journal-title":"Biodivers. Data J."},{"key":"ref_24","doi-asserted-by":"crossref","first-page":"355","DOI":"10.1111\/j.1471-8286.2007.01678.x","article-title":"Bold: The Barcode of Life Data System (http:\/\/www.barcodinglife.org)","volume":"7","author":"Ratnasingham","year":"2007","journal-title":"Mol. Ecol. Notes"},{"key":"ref_25","doi-asserted-by":"crossref","first-page":"D136","DOI":"10.1093\/nar\/gkr1178","article-title":"The NCBI Taxonomy Database","volume":"40","author":"Federhen","year":"2012","journal-title":"Nucleic Acids Res."},{"key":"ref_26","doi-asserted-by":"crossref","first-page":"2451","DOI":"10.1099\/ijs.0.63797-0","article-title":"\u201cCandidatus Phytoplasma Graminis\u201d and \u201cCandidatus Phytoplasma Caricae\u201d, Two Novel Phytoplasmas Associated with Diseases of Sugarcane, Weeds and Papaya in Cuba","volume":"55","author":"Arocha","year":"2005","journal-title":"Int. J. Syst. Evol. Microbiol."},{"key":"ref_27","doi-asserted-by":"crossref","first-page":"127","DOI":"10.1080\/07060661.2011.558854","article-title":"Identification and Molecular Characterization of the Phytoplasma Associated with Peach Rosette-like Disease at the Canadian Clonal Genebank Based on the 16S rRNA Gene Analysis","volume":"33","author":"Krukovets","year":"2011","journal-title":"Can. J. Plant Pathol."},{"key":"ref_28","doi-asserted-by":"crossref","first-page":"790","DOI":"10.1094\/PDIS.2001.85.7.790","article-title":"Revised Subgroup Classification of Group 16SrV Phytoplasmas and Placement of Flavescence Dor\u00e9e-Associated Phytoplasmas in Two Distinct Subgroups","volume":"85","author":"Davis","year":"2001","journal-title":"Plant Dis."},{"key":"ref_29","doi-asserted-by":"crossref","first-page":"3463","DOI":"10.1099\/ijsem.0.001218","article-title":"\u201cCandidatus Phytoplasma Hispanicum\u201d, a Novel Taxon Associated with Mexican Periwinkle Virescence Disease of Catharanthus Roseus","volume":"66","author":"Davis","year":"2016","journal-title":"Int. J. Syst. Evol. Microbiol."},{"key":"ref_30","doi-asserted-by":"crossref","first-page":"229","DOI":"10.1078\/0944-5013-00201","article-title":"Differential Amplification of Sequence Heterogeneous Ribosomal RNA Genes and Classification of the \u201cFragaria Multicipita\u201d Phytoplasma","volume":"158","author":"Davis","year":"2003","journal-title":"Microbiol. Res."},{"key":"ref_31","doi-asserted-by":"crossref","first-page":"1605","DOI":"10.1099\/00207713-49-4-1605","article-title":"The Phytoplasma Associated with Ash Yellows and Lilac Witches\u2019-Broom: \u201cCandidatus Phytoplasma Fraxini\u201d","volume":"49","author":"Griffiths","year":"1999","journal-title":"Int. J. Syst. Bacteriol."},{"key":"ref_32","doi-asserted-by":"crossref","first-page":"1349","DOI":"10.1099\/ijs.0.02842-0","article-title":"Clover Proliferation Phytoplasma: \u201cCandidatus Phytoplasma Trifolii.\u201d","volume":"54","author":"Hiruki","year":"2004","journal-title":"Int. J. Syst. Evol. Microbiol."},{"key":"ref_33","doi-asserted-by":"crossref","first-page":"1925","DOI":"10.1099\/ijs.0.02531-0","article-title":"\u201cCandidatus Phytoplasma Oryzae\u201d, a Novel Phytoplasma Taxon Associated with Rice Yellow Dwarf Disease","volume":"53","author":"Jung","year":"2003","journal-title":"Int. J. Syst. Evol. Microbiol."},{"key":"ref_34","doi-asserted-by":"crossref","first-page":"2887","DOI":"10.1099\/ijs.0.019695-0","article-title":"Phylogenetic Analysis and Delineation of Phytoplasmas Based on secY Gene Sequences","volume":"60","author":"Lee","year":"2010","journal-title":"Int. J. Syst. Evol. Microbiol."},{"key":"ref_35","doi-asserted-by":"crossref","first-page":"1037","DOI":"10.1099\/ijs.0.02843-0","article-title":"\u201cCandidatus Phytoplasma Asteris\u201d, a Novel Phytoplasma Taxon Associated with Aster Yellows and Related Diseases","volume":"54","author":"Lee","year":"2004","journal-title":"Int. J. Syst. Evol. Microbiol."},{"key":"ref_36","doi-asserted-by":"crossref","first-page":"337","DOI":"10.1099\/ijs.0.02697-0","article-title":"Classification of Phytoplasma Strains in the Elm Yellows Group (16SrV) and Proposal of \u201cCandidatus Phytoplasma Ulmi\u201d for the Phytoplasma Associated with Elm Yellows","volume":"54","author":"Lee","year":"2004","journal-title":"Int. J. Syst. Evol. Microbiol."},{"key":"ref_37","doi-asserted-by":"crossref","first-page":"2129","DOI":"10.1099\/ijs.0.025411-0","article-title":"Genetic Diversity of European Phytoplasmas of the 16SrV Taxonomic Group and Proposal of \u201cCandidatus Phytoplasma Rubi\u201d","volume":"61","author":"Salar","year":"2011","journal-title":"Int. J. Syst. Evol. Microbiol."},{"key":"ref_38","doi-asserted-by":"crossref","first-page":"1025","DOI":"10.1099\/ijs.0.02838-0","article-title":"\u201cCandidatus Phytoplasma Spartii\u201d, \u201cCandidatus Phytoplasma Rhamni\u201d and \u201cCandidatus Phytoplasma Allocasuarinae\u201d, Respectively Associated with Spartium Witches\u2019-Broom, Buckthorn Witches\u2019-Broom and Allocasuarina Yellows Diseases","volume":"54","author":"Marcone","year":"2004","journal-title":"Int. J. Syst. Evol. Microbiol."},{"key":"ref_39","first-page":"655","article-title":"Molecular Characterization and Transmission of Bermuda Grass White Leaf Phytoplasma in Iran","volume":"91","author":"Salehi","year":"2009","journal-title":"J. Plant Pathol."},{"key":"ref_40","doi-asserted-by":"crossref","first-page":"388","DOI":"10.1094\/PDIS-94-4-0388","article-title":"Genetic Diversity and Geographical Distribution of Phytoplasmas Associated with Potato Purple Top Disease in Mexico","volume":"94","year":"2010","journal-title":"Plant Dis."},{"key":"ref_41","first-page":"225","article-title":"Taxonomic Relatedness and Phylogenetic Positions of Phytoplasmas Associated with Diseases of Faba Bean, Sunnhemp, Sesame, Soybean, and Eggplant\/Taxonomische Verwandtschaft Und Phylogenetische Stellung von Phytoplasmen, Die Krankheiten Bei Pferdebohne, Crotalaria, Sesam, Sojabohne Und Aubergine Hervorrufen","volume":"102","author":"Schneider","year":"1995","journal-title":"J. Plant Dis. Prot."},{"key":"ref_42","doi-asserted-by":"crossref","first-page":"440","DOI":"10.1099\/00207713-44-3-440","article-title":"Phylogenetic Classification of Phytopathogenic Mollicutes by Sequence Analysis of 16S Ribosomal DNA","volume":"44","author":"Schneider","year":"1994","journal-title":"Int. J. Syst. Bacteriol."},{"key":"ref_43","doi-asserted-by":"crossref","first-page":"1217","DOI":"10.1099\/ijs.0.02823-0","article-title":"\u201cCandidatus Phytoplasma Mali\u201d, \u201cCandidatus Phytoplasma Pyri\u201d and \u201cCandidatus Phytoplasma Prunorum\u201d, the Causal Agents of Apple Proliferation, Pear Decline and European Stone Fruit Yellows, Respectively","volume":"54","author":"Schneider","year":"2004","journal-title":"Int. J. Syst. Evol. Microbiol."},{"key":"ref_44","first-page":"3","article-title":"Current Status of Molecular Classification of the Phytoplasmas","volume":"80","author":"Marcone","year":"1998","journal-title":"J. Plant Pathol."},{"key":"ref_45","doi-asserted-by":"crossref","first-page":"113","DOI":"10.1046\/j.1439-0329.2003.00312.x","article-title":"Molecular Characterization of a Phytoplasma from the Aster Yellows (16SrI) Group Naturally Infecting Populus nigra L. \u201cItalica\u201d Trees in Croatia","volume":"33","author":"Seruga","year":"2003","journal-title":"For. Pathol."},{"key":"ref_46","doi-asserted-by":"crossref","first-page":"127","DOI":"10.3989\/ajbm.2005.v62.i2.12","article-title":"Grapevine Yellows Diseases in Spain: Eight Year Survey of Disease Spread and Molecular Characterization of Phytoplasmas Involved","volume":"62","author":"Torres","year":"2005","journal-title":"An. Jard. Bot. Madr."},{"key":"ref_47","doi-asserted-by":"crossref","first-page":"437","DOI":"10.1111\/j.1744-7348.1998.tb05220.x","article-title":"Phylogenetic Relationships of Coconut Phytoplasmas and the Development of Specific Oligonucleotide PCR Primers","volume":"132","author":"Tymon","year":"1998","journal-title":"Ann. Appl. Biol."},{"key":"ref_48","doi-asserted-by":"crossref","first-page":"833","DOI":"10.1099\/ijs.0.02453-0","article-title":"\u201cCandidatus Phytoplasma Phoenicium\u201d sp. Nov., a Novel Phytoplasma Associated with an Emerging Lethal Disease of Almond Trees in Lebanon and Iran","volume":"53","author":"Verdin","year":"2003","journal-title":"Int. J. Syst. Evol. Microbiol."},{"key":"ref_49","doi-asserted-by":"crossref","first-page":"13","DOI":"10.1016\/j.cropro.2017.12.005","article-title":"Multilocus Genotyping Identifies a Highly Homogeneous Phytoplasma Lineage Associated with Sweet Cherry Virescence Disease in China and Its Carriage by an Erythroneurine Leafhopper","volume":"106","author":"Wang","year":"2018","journal-title":"Crop Prot."},{"key":"ref_50","doi-asserted-by":"crossref","first-page":"941","DOI":"10.1099\/00207713-48-3-941","article-title":"Phylogenetic Positions of Phytoplasmas Associated with Dieback, Yellow Crinkle and Mosaic Diseases of Papaya, and Their Proposed Inclusion in \u201cCandidatus Phytoplasma Australiense\u201d and a New Taxon, \u201cCandidatus Phytoplasma Australasia\u201d","volume":"48","author":"White","year":"1998","journal-title":"Int. J. Syst. Bacteriol."},{"key":"ref_51","doi-asserted-by":"crossref","first-page":"449","DOI":"10.1099\/00207713-45-3-449","article-title":"Characterization of the Mycoplasmalike Organism Associated with Witches\u2019-Broom Disease of Lime and Proposition of a Candidatus Taxon for the Organism, \u201cCandidatus Phytoplasma Aurantifolia\u201d","volume":"45","author":"Zreik","year":"1995","journal-title":"Int. J. Syst. Bacteriol."},{"key":"ref_52","doi-asserted-by":"crossref","first-page":"1792","DOI":"10.1093\/nar\/gkh340","article-title":"MUSCLE: Multiple Sequence Alignment with High Accuracy and High Throughput","volume":"32","author":"Edgar","year":"2004","journal-title":"Nucleic Acids Res."},{"key":"ref_53","doi-asserted-by":"crossref","first-page":"1547","DOI":"10.1093\/molbev\/msy096","article-title":"MEGA X: Molecular Evolutionary Genetics Analysis across Computing Platforms","volume":"35","author":"Kumar","year":"2018","journal-title":"Mol. Biol. Evol."},{"key":"ref_54","doi-asserted-by":"crossref","first-page":"1695","DOI":"10.1093\/molbev\/mss020","article-title":"Partitionfinder: Combined Selection of Partitioning Schemes and Substitution Models for Phylogenetic Analyses","volume":"29","author":"Lanfear","year":"2012","journal-title":"Mol. Biol. Evol."},{"key":"ref_55","first-page":"772","article-title":"PartitionFinder 2: New Methods for Selecting Partitioned Models of Evolution for Molecular and Morphological Phylogenetic Analyses","volume":"34","author":"Lanfear","year":"2016","journal-title":"Mol. Biol. Evol."},{"key":"ref_56","doi-asserted-by":"crossref","first-page":"1312","DOI":"10.1093\/bioinformatics\/btu033","article-title":"RAxML Version 8: A Tool for Phylogenetic Analysis and Post-Analysis of Large Phylogenies","volume":"30","author":"Stamatakis","year":"2014","journal-title":"Bioinformatics"},{"key":"ref_57","doi-asserted-by":"crossref","first-page":"373","DOI":"10.1111\/2041-210X.13512","article-title":"raxmlGUI 2.0: A Graphical Interface and Toolkit for Phylogenetic Analyses Using RAxML","volume":"12","author":"Edler","year":"2021","journal-title":"Methods Ecol. Evol."},{"key":"ref_58","doi-asserted-by":"crossref","unstructured":"Balbuena, J.A., M\u00edguez-Lozano, R., and Blasco-Costa, I. (2013). PACo: A Novel Procrustes Application to Cophylogenetic Analysis. PLoS ONE, 8.","DOI":"10.1371\/journal.pone.0061048"},{"key":"ref_59","doi-asserted-by":"crossref","first-page":"1212","DOI":"10.1093\/sysbio\/syaa033","article-title":"Random Tanglegram Partitions (Random TaPas): An Alexandrian Approach to the Cophylogenetic Gordian Knot","volume":"69","author":"Balbuena","year":"2020","journal-title":"Syst. Biol."},{"key":"ref_60","doi-asserted-by":"crossref","first-page":"932","DOI":"10.1111\/2041-210X.12736","article-title":"Paco: Implementing Procrustean Approach to Cophylogeny in R","volume":"8","author":"Hutchinson","year":"2017","journal-title":"Methods Ecol. Evol."},{"key":"ref_61","unstructured":"Oksanen, J., Blanchet, F.G., Kindt, R., Legendre, P., Minchin, P.R., O\u2019Hara, R.B., Simpson, G.L., Solymos, P., Stevens, M.H.H., and Wagner, H. (2026, July 01). Vegan: Community Ecology Package. R Package Version 2.2-7. Available online: http:\/\/CRAN.Rproject.org\/package=vegan."},{"key":"ref_62","doi-asserted-by":"crossref","first-page":"217","DOI":"10.1080\/10635150252899734","article-title":"A Statistical Test for Host-Parasite Coevolution","volume":"51","author":"Legendre","year":"2002","journal-title":"Syst. Biol."},{"key":"ref_63","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":"ref_64","doi-asserted-by":"crossref","first-page":"1428","DOI":"10.1111\/j.1420-9101.2007.01340.x","article-title":"When Can Host Shifts Produce Congruent Host and Parasite Phylogenies? A Simulation Approach","volume":"20","author":"Giraud","year":"2007","journal-title":"J. Evol. Biol."},{"key":"ref_65","unstructured":"R Core Team (2024). R: A Language and Environment for Statistical Computing, R Core Team."},{"key":"ref_66","doi-asserted-by":"crossref","first-page":"16","DOI":"10.1186\/1748-7188-5-16","article-title":"Jane: A New Tool for the Cophylogeny Reconstruction Problem","volume":"5","author":"Conow","year":"2010","journal-title":"Algorithms Mol. Biol."},{"key":"ref_67","unstructured":"Ronquist, F. (TreeFitter, 2002). TreeFitter, version 1.2."},{"key":"ref_68","unstructured":"Charleston, M.A., and Page, R.D.M. (2002). TreeMap 2. A Macintosh Program for Cophylogeny Mapping, University of Glasgow."},{"key":"ref_69","doi-asserted-by":"crossref","first-page":"725","DOI":"10.1093\/sysbio\/syt034","article-title":"Identifying Hidden Rate Changes in the Evolution of a Binary Morphological Character: The Evolution of Plant Habit in Campanulid Angiosperms","volume":"62","author":"Beaulieu","year":"2013","journal-title":"Syst. Biol."},{"key":"ref_70","unstructured":"Trivellone, V. (2020, March 15). Hemiptera-Phytoplasma-Plant Database Links. Available online: http:\/\/trivellone.speciesfile.org\/."},{"key":"ref_71","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":"ref_72","unstructured":"Orme, D., Freckleton, R., Thomas, G., Petzoldt, T., Fritz, S., Isaac, N., and Pearse, W. (2025). Caper: Comparative Analyses of Phylogenetics and Evolution in R, CRAN."},{"key":"ref_73","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":"ref_74","doi-asserted-by":"crossref","unstructured":"Wei, W., and Zhao, Y. (2022). Phytoplasma Taxonomy: Nomenclature, Classification, and Identification. Biology, 11.","DOI":"10.3390\/biology11081119"},{"key":"ref_75","doi-asserted-by":"crossref","unstructured":"Merkle, D., Middendorf, M., and Wieseke, N. (2010). A Parameter-Adaptive Dynamic Programming Approach for Inferring Cophylogenies. BMC Bioinform., 11.","DOI":"10.1186\/1471-2105-11-S1-S60"},{"key":"ref_76","doi-asserted-by":"crossref","first-page":"946","DOI":"10.1093\/sysbio\/syad016","article-title":"Rtapas: An R Package to Assess Cophylogenetic Signal Between Two Evolutionary Histories","volume":"72","author":"Krasnov","year":"2023","journal-title":"Syst. Biol."},{"key":"ref_77","doi-asserted-by":"crossref","first-page":"556","DOI":"10.1111\/j.2006.0030-1299.15025.x","article-title":"On Ecological Fitting, Plant-Insect Associations, Herbivore Host Shifts, and Host Plant Selection","volume":"114","author":"Agosta","year":"2006","journal-title":"Oikos"},{"key":"ref_78","doi-asserted-by":"crossref","first-page":"e02356-21","DOI":"10.1128\/aem.02356-21","article-title":"Phylogenetics of Historical Host Switches in a Bacterial Plant Pathogen","volume":"88","author":"Kahn","year":"2022","journal-title":"Appl. Environ. Microbiol."},{"key":"ref_79","doi-asserted-by":"crossref","first-page":"1309","DOI":"10.1111\/mec.14523","article-title":"Shifts in Diversification Rates and Host Jump Frequencies Shaped the Diversity of Host Range Among Sclerotiniaceae Fungal Plant Pathogens","volume":"27","author":"Navaud","year":"2018","journal-title":"Mol. Ecol."},{"key":"ref_80","doi-asserted-by":"crossref","first-page":"ieag059","DOI":"10.1093\/jisesa\/ieag059","article-title":"Mitochondrial Genomes of Psylloidea (Hemiptera): Candidate Broad-Range Primers for Long-Polymerase Chain Reaction, Genewise Variability, and Calibration-Sensitive Time-Calibrated Phylogeny","volume":"26","author":"Wu","year":"2026","journal-title":"J. Insect Sci."},{"key":"ref_81","doi-asserted-by":"crossref","first-page":"45","DOI":"10.1038\/nature14324","article-title":"Global Effects of Land Use on Local Terrestrial Biodiversity","volume":"520","author":"Newbold","year":"2015","journal-title":"Nature"},{"key":"ref_82","doi-asserted-by":"crossref","first-page":"6493","DOI":"10.1002\/ece3.7502","article-title":"Screening Potential Insect Vectors in a Museum Biorepository Reveals Undiscovered Diversity of Plant Pathogens in Natural Areas","volume":"11","author":"Trivellone","year":"2021","journal-title":"Ecol. Evol."},{"key":"ref_83","doi-asserted-by":"crossref","unstructured":"Wei, W., Trivellone, V., Dietrich, C.H., Zhao, Y., Bottner-Parker, K.D., and Ivanauskas, A. (2021). Identification of Phytoplasmas Representing Multiple New Genetic Lineages from Phloem-Feeding Leafhoppers Highlights the Diversity of Phytoplasmas and Their Potential Vectors. Pathogens, 10.","DOI":"10.3390\/pathogens10030352"},{"key":"ref_84","doi-asserted-by":"crossref","unstructured":"Trivellone, V., Mackay, A.J., Stone, C.M., and Dietrich, C.H. (2026). Leveraging Existing Biodiversity and Zoonosis Monitoring Infrastructure for Integrative Plant Pathogen Surveillance in Natural Ecosystems. Insects, 17.","DOI":"10.20944\/preprints202602.0241.v1"},{"key":"ref_85","doi-asserted-by":"crossref","first-page":"e70035","DOI":"10.1111\/syen.70035","article-title":"Analyses of Expanded Anchored-Hybrid Phylogenomic Datasets Support Revisions to the Higher Classification of Leafhoppers (Hemiptera: Cicadellidae)","volume":"51","author":"Cao","year":"2026","journal-title":"Syst. Entomol."}],"container-title":["Insects"],"original-title":[],"language":"en","link":[{"URL":"https:\/\/www.mdpi.com\/2075-4450\/17\/9\/942\/pdf","content-type":"unspecified","content-version":"vor","intended-application":"similarity-checking"}],"deposited":{"date-parts":[[2026,9,11]],"date-time":"2026-09-11T04:23:12Z","timestamp":1789100592000},"score":1,"resource":{"primary":{"URL":"https:\/\/www.mdpi.com\/2075-4450\/17\/9\/942"}},"subtitle":[],"short-title":[],"issued":{"date-parts":[[2026,9,9]]},"references-count":85,"journal-issue":{"issue":"9","published-online":{"date-parts":[[2026,9]]}},"alternative-id":["insects17090942"],"URL":"https:\/\/doi.org\/10.3390\/insects17090942","relation":{},"ISSN":["2075-4450"],"issn-type":[{"value":"2075-4450","type":"electronic"}],"subject":[],"published":{"date-parts":[[2026,9,9]]}}}