title: New fossil insect order Permopsocida elucidates major radiation and evolution of suction feeding in hemimetabolous insects (Hexapoda: Acercaria).
abstract: With nearly 100,000 species, the Acercaria (lice, plant lices, thrips, bugs) including number of economically important species is one of the most successful insect lineages. However, its phylogeny and evolution of mouthparts among other issues remain debatable. Here new methods of preparation permitted the comprehensive anatomical description of insect inclusions from mid-Cretaceous Burmese amber in astonishing detail. These "missing links" fossils, attributed to a new order Permopsocida, provide crucial evidence for reconstructing the phylogenetic relationships in the Acercaria, supporting its monophyly, and questioning the position of Psocodea as sister group of holometabolans in the most recent phylogenomic study. Permopsocida resolves as sister group of Thripida + Hemiptera and represents an evolutionary link documenting the transition from chewing to piercing mouthparts in relation to suction feeding. Identification of gut contents as angiosperm pollen documents an ecological role of Permopsocida as early pollen feeders with relatively unspecialized mouthparts. This group existed for 185 million years, but has never been diverse and was superseded by new pollenivorous pollinators during the Cretaceous co-evolution of insects and flowers. The key innovation of suction feeding with piercing mouthparts is identified as main event that triggered the huge post-Carboniferous radiation of hemipterans, and facilitated the spreading of pathogenic vectors.
authors.affiliation: State Key Laboratory of Palaeobiology and Stratigraphy, Nanjing Institute of Geology and Palaeontology, Chinese Academy of Sciences, Nanjing, People's Republic of China.
authors.forename: Di-Ying
authors.initials: DY
authors.lastname: Huang
authors.affiliation: Staatliches Museum für Naturkunde Stuttgart, Stuttgart, Germany.
authors.forename: Günter
authors.initials: G
authors.lastname: Bechly
authors.affiliation: Institut de Systématique, Évolution, Biodiversité, Muséum national d'Histoire naturelle, Paris, France.|AgroParisTech, Paris, France.
authors.forename: Patricia
authors.initials: P
authors.lastname: Nel
authors.affiliation: Division of Entomology, Natural History Museum, and Department of Ecology &Evolutionary Biology, University of Kansas, Lawrence, Kansas, United States of America.|Division of Invertebrate Zoology, American Museum of Natural History, New York, New York, United States of America.
authors.forename: Michael S
authors.initials: MS
authors.lastname: Engel
authors.affiliation: Charles University, Faculty of Science, Department of Zoology, Prague, Czech Republic.
authors.forename: Jakub
authors.initials: J
authors.lastname: Prokop
authors.affiliation: Lebanese University, Faculty of Sciences II, Department of Biology, Beirut, Lebanon.
authors.forename: Dany
authors.initials: D
authors.lastname: Azar
authors.affiliation: State Key Laboratory of Palaeobiology and Stratigraphy, Nanjing Institute of Geology and Palaeontology, Chinese Academy of Sciences, Nanjing, People's Republic of China.
authors.forename: Chen-Yang
authors.initials: CY
authors.lastname: Cai
authors.affiliation: ANKA/ Institute for Photon Science and Synchrotron Radiation, Karlsruhe Institute of Technology (KIT), Eggenstein-Leopoldshafen, Germany.|Laboratory for Applications of Synchrotron Radiation, Karlsruhe Institute of Technology (KIT), Karlsruhe, Germany.
authors.forename: Thomas
authors.initials: T
authors.lastname: van de Kamp
authors.affiliation: Staatliches Museum für Naturkunde Stuttgart, Stuttgart, Germany.
authors.forename: Arnold H
authors.initials: AH
authors.lastname: Staniczek
authors.affiliation: Institut de Systématique, Évolution, Biodiversité, Muséum national d'Histoire naturelle, Paris, France.
authors.forename: Romain
authors.initials: R
authors.lastname: Garrouste
authors.affiliation: Staatliches Museum für Naturkunde Stuttgart, Stuttgart, Germany.
authors.forename: Lars
authors.initials: L
authors.lastname: Krogmann
authors.affiliation: ANKA/ Institute for Photon Science and Synchrotron Radiation, Karlsruhe Institute of Technology (KIT), Eggenstein-Leopoldshafen, Germany.
authors.forename: Tomy
authors.initials: T
authors.lastname: Dos Santos Rolo
authors.affiliation: ANKA/ Institute for Photon Science and Synchrotron Radiation, Karlsruhe Institute of Technology (KIT), Eggenstein-Leopoldshafen, Germany.|Laboratory for Applications of Synchrotron Radiation, Karlsruhe Institute of Technology (KIT), Karlsruhe, Germany.
authors.forename: Tilo
authors.initials: T
authors.lastname: Baumbach
authors.affiliation: Im Königsfeld 22A, Saarbrücken, Germany.
authors.forename: Rainer
authors.initials: R
authors.lastname: Ohlhoff
authors.affiliation: Arthropoda Laboratory, Palaeontological Institute, Russian Academy of Sciences, Moscow, Russia.
authors.forename: Alexey S
authors.initials: AS
authors.lastname: Shmakov
authors.affiliation: Institut de Systématique, Évolution, Biodiversité, Muséum national d'Histoire naturelle, Paris, France.
authors.forename: Thierry
authors.initials: T
authors.lastname: Bourgoin
authors.affiliation: Institut de Systématique, Évolution, Biodiversité, Muséum national d'Histoire naturelle, Paris, France.
authors.forename: André
authors.initials: A
authors.lastname: Nel
mesh_terms: D018647:Amber; D000818:Animals; D005075:Biological Evolution; D004463:Ecology; D005580:Fossils / Q000033:anatomy & histology*; D007313:Insecta / Q000033:anatomy & histology* / Q000528:radiation effects; D019684:Magnoliopsida / Q000737:chemistry; D010802:Phylogeny; D011058:Pollen / Q000737:chemistry; D011827:Radiation
chemical_list: D018647:Amber