{"status":"ok","message-type":"work","message-version":"1.0.0","message":{"indexed":{"date-parts":[[2026,9,24]],"date-time":"2026-09-24T06:00:35Z","timestamp":1790229635807,"version":"4.1.0"},"reference-count":126,"publisher":"Wiley","license":[{"start":{"date-parts":[[2026,9,23]],"date-time":"2026-09-23T00:00:00Z","timestamp":1790121600000},"content-version":"vor","delay-in-days":0,"URL":"http:\/\/creativecommons.org\/licenses\/by\/4.0\/"},{"start":{"date-parts":[[2026,9,23]],"date-time":"2026-09-23T00:00:00Z","timestamp":1790121600000},"content-version":"tdm","delay-in-days":0,"URL":"http:\/\/doi.wiley.com\/10.1002\/tdm_license_1.1"}],"content-domain":{"domain":["onlinelibrary.wiley.com"],"crossmark-restriction":true},"short-container-title":["Cladistics"],"abstract":"<jats:title>Abstract<\/jats:title>\n                  <jats:p>Larvae, a novel type of immature stage, are a crucial innovation of the Holometabola. Larval features therefore provide key evidence for reconstructing evolutionary transformations in this megadiverse group. Here we present the first formal reconstruction of larval character evolution, using maximum parsimony and Bayesian ancestral\u2010state reconstruction. Our analyses are based on an updated matrix of 88 larval characters and a phylogenetic topology assembled from published transcriptomic and phylogenomic studies. A series of key larval transformations is associated with the origin of Holometabola. That includes partial reductions of antennal and labial structures, simplification of the tarsus and developmental shifts in the appearance of compound eyes and wing buds. The reconstructed holometabolan groundplan larva was terrestrial and non\u2010predaceous, with an orthognathous head, well\u2010developed biting mouthparts, well\u2010developed thoracic legs with an undivided tarsus, no abdominal appendages and a relatively slender postcephalic body. Within Holometabola, larval character support for major clades was highly uneven. Some lineages, especially Neuroptera, Strepsiptera, Vespina, Glossata and Brachycera, showed dense clusters of inferred transformations and distinctive larval apomorphies. In contrast, several major clades, including Coleoptera, Lepidoptera, Mecopterida and Antliophora, were weakly supported or unsupported by larval synapomorphies, despite strong support from adult morphology in some cases. Major shifts in larval morphology were associated with ecological and functional transitions, including predation, parasitism, parasitoidism, phytophagy, aquatic habits, wood boring and development in moist substrates. Our results show a distinctive mosaic pattern in larval evolution in Holometabola, with major transformations largely restricted to some of the phylogenetic lineages and closely associated with key ecological transitions.<\/jats:p>","DOI":"10.1111\/cla.70067","type":"journal-article","created":{"date-parts":[[2026,9,24]],"date-time":"2026-09-24T05:07:09Z","timestamp":1790226429000},"update-policy":"https:\/\/doi.org\/10.1002\/crossmark_policy","source":"Crossref","is-referenced-by-count":0,"title":["Larval character evolution in Holometabola (Insecta)"],"prefix":"10.1111","author":[{"ORCID":"https:\/\/orcid.org\/0000-0002-0433-7626","authenticated-orcid":false,"given":"Rolf Georg","family":"Beutel","sequence":"first","affiliation":[{"name":"Entomology Group Institut f\u00fcr Zoologie und Evolutionsforschung, Friedrich\u2010Schiller\u2010Universit\u00e4t Jena  Jena Germany"},{"name":"Entomology II Section Senckenberg Forschungsinstitut und Naturmuseum Frankfurt  Frankfurt Germany"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0003-2538-6686","authenticated-orcid":false,"given":"Jakub","family":"Gocza\u0142","sequence":"additional","affiliation":[{"name":"Department of Forest Ecosystem Protection Faculty of Forestry University of Agriculture in Krakow  Krakow Poland"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0003-4443-0806","authenticated-orcid":false,"given":"Di","family":"Li","sequence":"additional","affiliation":[{"name":"Entomology II Section Senckenberg Forschungsinstitut und Naturmuseum Frankfurt  Frankfurt Germany"}],"role":[{"vocabulary":"crossref","role":"author"}]}],"member":"311","published-online":{"date-parts":[[2026,9,23]]},"reference":[{"key":"e_1_2_9_2_1","first-page":"109","article-title":"The larva of Apterobittacus apterus Maclachlan (Mecoptera: Panorpidae [sic])","volume":"4","author":"Applegarth A.G.","year":"1939","journal-title":"Microentomology"},{"key":"e_1_2_9_3_1","first-page":"451","article-title":"Verbliebene Vielfalt vergangener Bl\u00fcte. Zur Evolution, Phylogenie und Biodiversit\u00e4t der Neuropterida (Insecta: Endopterygota)","volume":"20","author":"Asp\u00f6ck U.","year":"2007","journal-title":"Denisia"},{"key":"e_1_2_9_4_1","doi-asserted-by":"crossref","first-page":"864","DOI":"10.1016\/B978-0-12-374144-8.00226-5","volume-title":"Encyclopedia of Insects","author":"Asp\u00f6ck U.","year":"2009"},{"key":"e_1_2_9_5_1","doi-asserted-by":"crossref","first-page":"830","DOI":"10.1007\/s13744-022-00993-4","article-title":"Distinguishing four Calliphoridae species (Diptera) from Jamaica using the cephalopharyngeal skeleton: application to forensic investigations","volume":"51","author":"Barrett T.A.","year":"2022","journal-title":"Neotrop. Entomol."},{"key":"e_1_2_9_6_1","volume-title":"Insect Metamorphosis: From Natural History to Regulation of Development and Evolution","author":"Bell\u00e9s X.","year":"2020"},{"key":"e_1_2_9_7_1","doi-asserted-by":"crossref","first-page":"60","DOI":"10.1016\/j.ibmb.2014.06.009","article-title":"The MEKRE93 (methoprene tolerant\u2010Kr\u00fcppel homolog 1\u2010E93) pathway in the regulation of insect metamorphosis, and the homology of the pupal stage","volume":"52","author":"Belles X.","year":"2014","journal-title":"Insect Biochem. Mol. Biol."},{"key":"e_1_2_9_8_1","doi-asserted-by":"crossref","first-page":"127","DOI":"10.1111\/j.1365-3113.1993.tb00658.x","article-title":"Phylogenetic analysis of Adephaga (Coleoptera) based on characters of the larval head","volume":"18","author":"Beutel R.G.","year":"1993","journal-title":"Syst. Entomol."},{"key":"e_1_2_9_9_1","doi-asserted-by":"crossref","first-page":"145","DOI":"10.1111\/j.1439-0469.1995.tb00222.x","article-title":"Phylogenetic analysis of Elateriformia (Coleoptera: Polyphaga) based on larval characters","volume":"33","author":"Beutel R.G.","year":"1995","journal-title":"J. Zoolog. Syst. Evol. Res."},{"key":"e_1_2_9_10_1","doi-asserted-by":"crossref","first-page":"9","DOI":"10.1163\/22119434-99900016","article-title":"Morphology and evolution of the larval head structures of Hydrophiloidea and Histeroidea (Coeloptera: Staphyliniformia)","volume":"142","author":"Beutel R.G.","year":"1999","journal-title":"Tijdschr. Entomol."},{"key":"e_1_2_9_11_1","doi-asserted-by":"crossref","first-page":"917","DOI":"10.14411\/eje.2008.119","article-title":"Comparative study of larval head structures of Megaloptera (Hexapoda)","volume":"105","author":"Beutel R.G.","year":"2008","journal-title":"Eur. J. Entomol."},{"key":"e_1_2_9_12_1","volume-title":"Handbook of Zoology","author":"Beutel R.G.","year":"2019"},{"key":"e_1_2_9_13_1","doi-asserted-by":"crossref","first-page":"89","DOI":"10.1016\/j.zool.2007.03.005","article-title":"The larval head of Raphidia (Raphidioptera, Insecta) and its phylogenetic significance","volume":"11","author":"Beutel R.G.","year":"2008","journal-title":"Zoology"},{"key":"e_1_2_9_14_1","first-page":"103","article-title":"Larval head morphology of Hydroscapha natans LeConte, 1874 (Coleoptera, Myxophaga, Hydroscaphidae) with special reference to miniaturization","volume":"118","author":"Beutel R.G.","year":"1998","journal-title":"Zoomorphology"},{"key":"e_1_2_9_15_1","doi-asserted-by":"crossref","first-page":"103","DOI":"10.1111\/j.1096-0031.2000.tb00350.x","article-title":"Phylogenetic relationships of the suborders of Coleoptera (Insecta)","volume":"16","author":"Beutel R.G.","year":"2000","journal-title":"Cladistics"},{"key":"e_1_2_9_16_1","doi-asserted-by":"crossref","first-page":"169","DOI":"10.1046\/j.1365-3113.2002.00172.x","article-title":"Larval morphology and phylogenetic position of Micromalthus debilis LeConte (Coleoptera: Micromalthidae)","volume":"27","author":"Beutel R.G.","year":"2002","journal-title":"Syst. Entomol."},{"key":"e_1_2_9_17_1","doi-asserted-by":"crossref","first-page":"53","DOI":"10.14411\/eje.2002.011","article-title":"Description of the larva of Rhipsideigma raffrayi (Coleoptera: Archostemata), with phylogenetic and functional implications","volume":"99","author":"Beutel R.G.","year":"2002","journal-title":"Eur. J. Entomol."},{"key":"e_1_2_9_18_1","doi-asserted-by":"crossref","first-page":"510","DOI":"10.1111\/j.1365-3113.2005.00293.x","article-title":"Phylogenetic analysis of Staphyliniformia (Coleoptera) based on characters of larvae and adults","volume":"30","author":"Beutel R.G.","year":"2005","journal-title":"Syst. Entomol."},{"key":"e_1_2_9_19_1","first-page":"37","article-title":"Comparative morphological study of larvae of Staphylinoidea (Coleoptera, Polyphaga) with phylogenetic implications","volume":"236","author":"Beutel R.G.","year":"1997","journal-title":"Zool. Anz."},{"key":"e_1_2_9_20_1","doi-asserted-by":"crossref","first-page":"825","DOI":"10.1071\/IT00030","article-title":"Larval head morphology of Phycosecis litoralis (Pascoe) (Coleoptera: Phycosecidae) with phylogenetic implications","volume":"14","author":"Beutel R.G.","year":"2000","journal-title":"Invertebr. Taxon."},{"key":"e_1_2_9_21_1","doi-asserted-by":"crossref","first-page":"171","DOI":"10.1046\/j.1365-3113.1999.00075.x","article-title":"Phylogenetic analysis of Myxophaga (Coleoptera) using larval characters","volume":"24","author":"Beutel R.G.","year":"1999","journal-title":"Syst. Entomol."},{"key":"e_1_2_9_22_1","doi-asserted-by":"crossref","first-page":"118","DOI":"10.1111\/j.1439-0469.2007.00446.x","article-title":"The larval head morphology of Xyela sp. (Xyelidae, Hymenoptera) and its phylogenetic implications","volume":"46","author":"Beutel R.G.","year":"2008","journal-title":"J. Zoolog. Syst. Evol. Res."},{"key":"e_1_2_9_23_1","doi-asserted-by":"crossref","first-page":"427","DOI":"10.1016\/j.asd.2009.05.002","article-title":"Resolving insect phylogeny: The significance of cephalic structures of the Nannomecoptera in understanding endopterygote relationships","volume":"38","author":"Beutel R.G.","year":"2009","journal-title":"Arthropod Struct. Dev."},{"key":"e_1_2_9_24_1","doi-asserted-by":"crossref","first-page":"533","DOI":"10.1111\/j.1096-3642.2009.00560.x","article-title":"The larval head of Nevrorthidae and the phylogeny of Neuroptera (Insecta)","volume":"158","author":"Beutel R.G.","year":"2010","journal-title":"Zool. J. Linn. Soc."},{"key":"e_1_2_9_25_1","doi-asserted-by":"crossref","first-page":"341","DOI":"10.1111\/j.1096-0031.2010.00338.x","article-title":"Morphological and molecular evidence converge upon a robust phylogeny of the megadiverse Holometabola","volume":"27","author":"Beutel R.G.","year":"2011","journal-title":"Cladistics"},{"key":"e_1_2_9_26_1","doi-asserted-by":"crossref","DOI":"10.1515\/9783110264043","volume-title":"Insect Morphology and Phylogeny: A Textbook for Students of Entomology","author":"Beutel R.G.","year":"2013"},{"key":"e_1_2_9_27_1","doi-asserted-by":"crossref","first-page":"75","DOI":"10.1111\/syen.12316","article-title":"The phylogeny of Coleopterida (Hexapoda) \u2013 morphological characters and molecular phylogenies","volume":"44","author":"Beutel R.G.","year":"2019","journal-title":"Syst. Entomol."},{"key":"e_1_2_9_28_1","doi-asserted-by":"crossref","first-page":"378","DOI":"10.1111\/syen.12403","article-title":"The morphological evolution of the Adephaga (Coleoptera)","volume":"45","author":"Beutel R.G.","year":"2020","journal-title":"Syst. Entomol."},{"key":"e_1_2_9_29_1","doi-asserted-by":"crossref","first-page":"227","DOI":"10.1111\/cla.12483","article-title":"Patterns of morphological simplification and innovation in the megadiverse Holometabola (Insecta)","volume":"38","author":"Beutel R.G.","year":"2022","journal-title":"Cladistics"},{"issue":"3","key":"e_1_2_9_30_1","doi-asserted-by":"crossref","first-page":"355","DOI":"10.1111\/syen.12623","article-title":"The evolutionary history of Coleoptera (Insecta) in the late Palaeozoic and the Mesozoic","volume":"49","author":"Beutel R.G.","year":"2024","journal-title":"Syst. Entomol."},{"key":"e_1_2_9_31_1","doi-asserted-by":"crossref","first-page":"149","DOI":"10.1146\/annurev-ento-121423-013358","article-title":"Evolutionary adaptations in larvae of Holometabola","volume":"71","author":"Beutel R.G.","year":"2026","journal-title":"Annu. Rev. Entomol."},{"key":"e_1_2_9_32_1","first-page":"49","article-title":"Der Larvenkopf von Panorpa communis L. und seine Verwandlung, mit besonderer Ber\u00fccksichtigung des Gehirns und der Augen","volume":"68","author":"Bierbrodt E.","year":"1942","journal-title":"Zool. Jb. Anat."},{"key":"e_1_2_9_33_1","doi-asserted-by":"crossref","DOI":"10.1038\/s41467-023-36868-4","article-title":"Key innovations and the diversification of Hymenoptera","volume":"14","author":"Blaimer B.B.","year":"2023","journal-title":"Nat. Commun."},{"key":"e_1_2_9_34_1","unstructured":"Blank S.M. 2002.Biosystematics of the extant Xyelidae with particular emphasis on the Old World taxa (Insecta: Hymenoptera). Unpublished PhD thesis Freie Universit\u00e4t Berlin."},{"issue":"1","key":"e_1_2_9_35_1","doi-asserted-by":"crossref","first-page":"69","DOI":"10.1111\/syen.12562","article-title":"Permian parallelisms: reanalysis of \u2020Tshekardocoleidae sheds light on the earliest evolution of the Coleoptera","volume":"48","author":"Boudinot B.E.","year":"2023","journal-title":"Syst. Entomol."},{"issue":"1","key":"e_1_2_9_36_1","doi-asserted-by":"crossref","first-page":"203","DOI":"10.1146\/annurev.en.28.010183.001223","article-title":"Biology of the Mecoptera","volume":"28","author":"Byers G.W.","year":"1983","journal-title":"Annu. Rev. Entomol."},{"key":"e_1_2_9_37_1","first-page":"36","article-title":"The morphology of the larval heads of certain Culicidae","volume":"9","author":"Cook E.F.","year":"1944","journal-title":"Microentomol. Palo Alto"},{"key":"e_1_2_9_38_1","first-page":"69","article-title":"On the morphology of the larval head of Chironomus (Diptera: Chironomidae)","volume":"9","author":"Cook E.F.","year":"1944","journal-title":"Microentomol. Palo Alto"},{"key":"e_1_2_9_39_1","first-page":"1","article-title":"The evolution of the head in the larvae of Diptera","volume":"14","author":"Cook E.F.","year":"1949","journal-title":"Microentomol. Palo Alto"},{"key":"e_1_2_9_40_1","volume-title":"The Biology of the Coleoptera","author":"Crowson R.A.","year":"1981"},{"key":"e_1_2_9_41_1","doi-asserted-by":"crossref","first-page":"957","DOI":"10.1002\/jmor.20033","article-title":"Kauri seeds and larval somersaults: The larval trunk of the seed mining basal moth Agathiphaga vitensis (Lepidoptera: Agathiphagidae)","volume":"273","author":"Dupont S.","year":"2012","journal-title":"J. Morphol."},{"key":"e_1_2_9_42_1","doi-asserted-by":"crossref","first-page":"1239","DOI":"10.1002\/jmor.20176","article-title":"Early leaf miners and the ground plan of the lepidopteran larval trunk: Caterpillar morphology of the basal moths Heterobathmia, Eriocrania, and Acanthopteroctetes","volume":"274","author":"Dupont S.","year":"2013","journal-title":"J. Morphol."},{"key":"e_1_2_9_43_1","doi-asserted-by":"crossref","first-page":"797","DOI":"10.1002\/jmor.20264","article-title":"Chambered cuticle, pellicles, strange sensilla, and extraordinary muscle arrangements: A study of the micropterigid larval trunk (Lepidoptera: Micropterigidae)","volume":"275","author":"Dupont S.T.","year":"2014","journal-title":"J. Morphol."},{"key":"e_1_2_9_44_1","doi-asserted-by":"crossref","first-page":"385","DOI":"10.3897\/asp.73.e31825","article-title":"The larval cephalic morphology of the enigmatic boreid Caurinus dectes (Mecoptera) and its phylogenetic significance","volume":"73","author":"Fabian B.","year":"2015","journal-title":"Arthropod Syst. Phylogeny"},{"key":"e_1_2_9_45_1","doi-asserted-by":"crossref","DOI":"10.1002\/jmor.70048","article-title":"Transformations of head structures during the larval development of the black soldier fly Hermetia illucens (Stratiomyidae, Diptera)","volume":"286","author":"Fabian B.","year":"2025","journal-title":"J. Morphol."},{"key":"e_1_2_9_46_1","doi-asserted-by":"crossref","first-page":"175","DOI":"10.1111\/syen.12386","article-title":"Reliable placement of beetle fossils via phylogenetic analyses \u2013 Triassic Leehermania as a case study: Staphylinidae or Myxophaga?","volume":"45","author":"Fik\u00e1\u010dek M.","year":"2020","journal-title":"Syst. Entomol."},{"key":"e_1_2_9_47_1","doi-asserted-by":"crossref","first-page":"1505","DOI":"10.1002\/jmor.20475","article-title":"The larval head anatomy of Rhyacophila (Rhyacophilidae) with discussion on mouthpart homology and the groundplan of Trichoptera","volume":"276","author":"Friedrich F.","year":"2015","journal-title":"J. Morphol."},{"key":"e_1_2_9_48_1","doi-asserted-by":"crossref","first-page":"323","DOI":"10.1146\/annurev.en.39.010194.001543","article-title":"Form and function of stemmata in larvae of holometabolous insects","volume":"39","author":"Gilbert C.","year":"1994","journal-title":"Annu. Rev. Entomol."},{"key":"e_1_2_9_49_1","doi-asserted-by":"crossref","DOI":"10.1093\/zoolinnean\/zlaf102","article-title":"Formation of beetle elytra and the most spectacular animal radiation in the history of Earth","volume":"204","author":"Gocza\u0142 J.","year":"2025","journal-title":"Zool. J. Linn. Soc."},{"key":"e_1_2_9_50_1","unstructured":"Grabarek V. 2008.Die Kopfmorphologie vonNeodiprion sertifer(Geoffroy 1785) (Diprionidae) und ihre phylogenetische Bedeutung. Unpubl. Diploma thesis FSU Jena."},{"key":"e_1_2_9_51_1","doi-asserted-by":"crossref","first-page":"141","DOI":"10.1016\/0040-8166(80)90057-9","article-title":"Cryptonephric malpighian tubule system in a dipteran larva, the New Zealand glow\u2010worm, Arachnocampa luminosa (Diptera: Mycetophilidae): a structural study","volume":"12","author":"Green L.F.","year":"1980","journal-title":"Tissue Cell"},{"key":"e_1_2_9_52_1","first-page":"133","volume-title":"Lepidoptera, Moths and Butterflies. Vol. 2: Morphology, Physiology, and Development. Handbook of Zoology. Vol. IV Arthropoda: Insecta. Part 36","author":"Hasenfuss I.","year":"2003"},{"key":"e_1_2_9_53_1","doi-asserted-by":"crossref","first-page":"23","DOI":"10.3897\/zookeys.798.28606","article-title":"A new species of Boloponera from Sekhukhuneland, South Africa (Hymenoptera, Formicidae, Ponerinae)","volume":"798","author":"Hawkes P.G.","year":"2018","journal-title":"ZooKeys"},{"key":"e_1_2_9_54_1","volume-title":"Die Stammesgeschichte der Insekten","author":"Hennig W.","year":"1970"},{"key":"e_1_2_9_55_1","volume-title":"Diptera (Zweifl\u00fcgler). Handbuch der Zoologie IV, Insecta. Lieferung 20","author":"Hennig W.","year":"1973"},{"key":"e_1_2_9_56_1","doi-asserted-by":"crossref","first-page":"111","DOI":"10.1079\/9781800623545.0006","volume-title":"Chalcidoidea of the World","author":"Heraty J.M.","year":"2025"},{"key":"e_1_2_9_57_1","doi-asserted-by":"crossref","first-page":"282","DOI":"10.1111\/j.1096-3642.1946.tb00124.x","article-title":"A new classification of insect pupae","volume":"116","author":"Hinton H.E.","year":"1946","journal-title":"Proc. Zool. Soc. London"},{"key":"e_1_2_9_58_1","doi-asserted-by":"crossref","first-page":"395","DOI":"10.1111\/j.1365-2311.1948.tb01227.x","article-title":"On the origin and function of the pupal stage","volume":"99","author":"Hinton H.E.","year":"1948","journal-title":"Trans. R. Entomol. Soc. London"},{"key":"e_1_2_9_59_1","doi-asserted-by":"crossref","first-page":"181","DOI":"10.1146\/annurev.en.03.010158.001145","article-title":"The phylogeny of the panorpoid Orders","volume":"3","author":"Hinton H.E.","year":"1958","journal-title":"Annu. Rev. Entomol."},{"key":"e_1_2_9_60_1","first-page":"29","volume-title":"Handbook of Zoology, Vol. IV Arthropoda: Insecta. Part 38. Coleoptera, Vol. 1: Morphology and Systematics (Archostemata, Adephaga, Myxophaga, Polyphaga (partim))","author":"H\u00f6rnschemeyer T.","year":"2005"},{"key":"e_1_2_9_61_1","doi-asserted-by":"crossref","first-page":"3026","DOI":"10.1098\/rspb.2011.0134","article-title":"Allopatric distribution and diversification without niche shift in a bryophyte\u2010feeding basal moth lineage (Lepidoptera: Micropterigidae)","volume":"278","author":"Imada Y.","year":"2011","journal-title":"Proc. R. Soc. B: Biol. Sci."},{"key":"e_1_2_9_62_1","first-page":"529","article-title":"Morphology of the primary larva of Mantispa aphavexelte Asp\u00f6ck and Asp\u00f6ck, 1994 (Neuroptera: Mantispidae) and phylogenetic implications to the order of Neuroptera","volume":"76","author":"Jandausch K.","year":"2018","journal-title":"Arthr. Syst. Phyl."},{"key":"e_1_2_9_63_1","doi-asserted-by":"crossref","first-page":"1742","DOI":"10.1002\/jmor.21060","article-title":"The larval morphology of the spongefly Sisyra nigra (Retzius, 1783) (Neuroptera: Sisyridae)","volume":"280","author":"Jandausch K.","year":"2019","journal-title":"J. Morphol."},{"key":"e_1_2_9_64_1","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 K.P.","year":"2018","journal-title":"Proc. Natl. Acad. Sci. U. S. A."},{"key":"e_1_2_9_65_1","volume-title":"Handbuch der Zoologie IV. Insecta. Inst. 25","author":"Kaltenbach A.","year":"1978"},{"key":"e_1_2_9_66_1","doi-asserted-by":"crossref","first-page":"107","DOI":"10.3897\/asp.74.e31842","article-title":"Serial block\u2010face scanning electron microscopy sheds new light on the head anatomy of an extremely miniaturized insect larva (Strepsiptera)","volume":"74","author":"Knauthe P.","year":"2016","journal-title":"Arthropod Syst. Phylogeny"},{"key":"e_1_2_9_67_1","doi-asserted-by":"crossref","first-page":"63","DOI":"10.1111\/j.1365-3113.1984.tb00502.x","article-title":"The larval head of Agathiphaga (Lepidoptera, Agathiphagidae) and the lepidopteran groundplan","volume":"9","author":"Kristensen N.P.","year":"1984","journal-title":"Syst. Entomol."},{"key":"e_1_2_9_68_1","first-page":"125","volume-title":"The Insects of Australia, vol. 1","author":"Kristensen N.P.","year":"1991"},{"key":"e_1_2_9_69_1","first-page":"237","article-title":"Phylogeny of endopterygote insects, the most successful lineage of living organisms","volume":"96","author":"Kristensen N.P.","year":"2013","journal-title":"Eur. J. Entomol."},{"key":"e_1_2_9_70_1","first-page":"7","volume-title":"Lepidoptera, Moths and Butterflies. Vol. 1: Evolution, Systematics and Biogeography. Handbook of Zoology, Vol. IV Arthropoda: Insecta. Part 35","author":"Kristensen N.P.","year":"1999"},{"key":"e_1_2_9_71_1","doi-asserted-by":"crossref","first-page":"699","DOI":"10.11646\/zootaxa.1668.1.30","article-title":"Lepidoptera phylogeny and systematics: the state of inventorying moth and butterfly diversity","volume":"1668","author":"Kristensen N.P.","year":"2007","journal-title":"Zootaxa"},{"key":"e_1_2_9_72_1","doi-asserted-by":"crossref","first-page":"181","DOI":"10.4039\/Ent125181-2","article-title":"Evolution of the hind wing in Coleoptera","volume":"125","author":"Kukalov\u00e1\u2010Peck J.","year":"1993","journal-title":"Can. Entomol."},{"key":"e_1_2_9_73_1","first-page":"9","article-title":"Evidence for an earliest Late Carboniferous divergence time and the early larval ecology and diversification of major Holometabola lineages","volume":"117","author":"Labandeira C.C.","year":"2011","journal-title":"Entomol. Am."},{"key":"e_1_2_9_74_1","doi-asserted-by":"crossref","first-page":"309","DOI":"10.1111\/syen.12657","article-title":"Lacewing\u2010specific Universal Single Copy Orthologs designed towards resolution of backbone phylogeny of Neuropterida","volume":"50","author":"Lai Y.","year":"2025","journal-title":"Syst. Entomol."},{"key":"e_1_2_9_75_1","doi-asserted-by":"crossref","first-page":"369","DOI":"10.1071\/IT99008","article-title":"The Australian Ommatidae (Coleoptera: Archostemata): new species, larva and discussion of relationships","volume":"13","author":"Lawrence J.F.","year":"1999","journal-title":"Invertebr. Taxon."},{"key":"e_1_2_9_76_1","doi-asserted-by":"crossref","first-page":"261","DOI":"10.1146\/annurev.es.13.110182.001401","article-title":"Evolution and classification of beetles","volume":"13","author":"Lawrence J.F.","year":"1982","journal-title":"Annu. Rev. Ecol. Syst."},{"key":"e_1_2_9_77_1","doi-asserted-by":"crossref","first-page":"1","DOI":"10.3161\/000345411X576725","article-title":"Phylogeny of the Coleoptera based on morphological characters of adults and larvae","volume":"61","author":"Lawrence J.F.","year":"2011","journal-title":"Ann. Zool."},{"key":"e_1_2_9_78_1","doi-asserted-by":"crossref","first-page":"913","DOI":"10.1080\/106351501753462876","article-title":"A likelihood approach to estimating phylogeny from discrete morphological character data","volume":"50","author":"Lewis P.O.","year":"2001","journal-title":"Syst. Biol."},{"key":"e_1_2_9_79_1","doi-asserted-by":"crossref","first-page":"1445","DOI":"10.1111\/1744-7917.13175","article-title":"Cephalic anatomy highlights morphological adaptation to underground habitats in a minute lacewing larva of Dilar (Dilaridae) and conflicting phylogenetic signal in Neuroptera","volume":"30","author":"Li D.","year":"2023","journal-title":"Insect. Sci."},{"key":"e_1_2_9_80_1","first-page":"161","article-title":"The larval morphology of the clonal raider ant, Ooceraea biroi (Forel, 1907) (Dorylinae), with broader implications for the Formicidae and Hymenoptera","volume":"35","author":"Li D.","year":"2025","journal-title":"Myrmecol. News"},{"key":"e_1_2_9_81_1","doi-asserted-by":"crossref","first-page":"180","DOI":"10.14411\/eje.2015.018","article-title":"Azalea sawfly Nematus lipovskyi (Hymenoptera: Tenthredinidae), a new invasive species in Europe","volume":"112","author":"Macek J.","year":"2015","journal-title":"Eur. J. Entomol."},{"key":"e_1_2_9_82_1","unstructured":"Maddison W.PandMaddison D.R.2019.Mesquite: a modular system for evolutionary analysis. 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Gen."},{"key":"e_1_2_9_88_1","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 B.","year":"2014","journal-title":"Science"},{"key":"e_1_2_9_89_1","doi-asserted-by":"crossref","first-page":"213","DOI":"10.1016\/j.jcz.2009.10.001","article-title":"The morphology of the larval head of Tipulidae (Diptera, Insecta) \u2013 the dipteran groundplan and evolutionary trends","volume":"248","author":"Neugart C.","year":"2009","journal-title":"Zool. Anz."},{"key":"e_1_2_9_90_1","doi-asserted-by":"crossref","first-page":"1309","DOI":"10.1016\/j.cub.2012.05.018","article-title":"Genomic and morphological evidence converge to resolve the enigma of Strepsiptera","volume":"22","author":"Niehuis O.","year":"2012","journal-title":"Curr. Biol."},{"key":"e_1_2_9_91_1","volume-title":"WinClada, version 1.00. 08","author":"Nixon K.C.","year":"2002"},{"key":"e_1_2_9_92_1","first-page":"93","article-title":"Life history traits, larval habits and larval morphology of a leafminer, Coptotriche japoniella (Tischeriidae), on an evergreen tree, Eurya japonica (Theaceae)","volume":"63","author":"Oishi M.","year":"2009","journal-title":"J. Lepid. Soc."},{"key":"e_1_2_9_93_1","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 E.","year":"2004","journal-title":"Bioinformatics"},{"key":"e_1_2_9_94_1","first-page":"157","volume-title":"Recent Research in Neuropterology. 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Entomol."},{"key":"e_1_2_9_98_1","doi-asserted-by":"crossref","first-page":"328","DOI":"10.1111\/j.1096-0031.2005.00074.x","article-title":"The phylogeny of Strepsiptera (Hexapoda)","volume":"21","author":"Pohl H.","year":"2005","journal-title":"Cladistics"},{"key":"e_1_2_9_99_1","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 L.J.","year":"2012","journal-title":"Methods Ecol. Evol."},{"key":"e_1_2_9_100_1","first-page":"61","article-title":"Morphologie des Kopfes und des Vorderdarmes der Larve und Imago von Sialis flavilatera","volume":"67","author":"R\u00f6ber H.","year":"1942","journal-title":"Zool. Jahrb., Abt. Anat."},{"issue":"3","key":"e_1_2_9_101_1","doi-asserted-by":"crossref","first-page":"539","DOI":"10.1093\/sysbio\/sys029","article-title":"MrBayes 3.2: efficient Bayesian phylogenetic inference and model choice across a large model space","volume":"61","author":"Ronquist F.","year":"2012","journal-title":"Syst. Biol."},{"key":"e_1_2_9_102_1","doi-asserted-by":"crossref","first-page":"531","DOI":"10.1111\/syen.12545","article-title":"The unresolved phylogenomic tree of butterflies and moths (Lepidoptera): Assessing the potential causes and consequences","volume":"47","author":"Rota J.","year":"2022","journal-title":"Syst. Entomol."},{"key":"e_1_2_9_103_1","doi-asserted-by":"crossref","first-page":"193","DOI":"10.1016\/B978-0-12-814545-6.00005-5","volume-title":"Mosquitoes, Communities, and Public Health in Texas","author":"Rueda L.M.","year":"2020"},{"key":"e_1_2_9_104_1","doi-asserted-by":"crossref","first-page":"293","DOI":"10.1016\/j.asd.2012.01.002","article-title":"The larvae of Nymphomyiidae (Diptera, Insecta) \u2013 ancestral and highly derived?","volume":"41","author":"Schneeberg K.","year":"2012","journal-title":"Arthropod Struct. Dev."},{"key":"e_1_2_9_105_1","doi-asserted-by":"crossref","first-page":"465","DOI":"10.1098\/rstb.1937.0007","article-title":"On the internal anatomy of the larva of the rat\u2010flea Nosopsyllus fasciatus (Bosc)","volume":"227","author":"Sharif M.","year":"1937","journal-title":"Philos. Trans. R. Soc. Lond. B Biol. Sci."},{"key":"e_1_2_9_106_1","doi-asserted-by":"crossref","first-page":"233","DOI":"10.1111\/j.1365-3113.1992.tb00335.x","article-title":"A phylogenetic interpretation of the Brachycera (Diptera) based on the larval mandible and associated mouthpart structures","volume":"17","author":"Sinclair B.J.","year":"1992","journal-title":"Syst. Entomol."},{"key":"e_1_2_9_107_1","doi-asserted-by":"crossref","first-page":"161","DOI":"10.1007\/s13127-021-00511-1","article-title":"Morphological and molecular inference of immature stages of Larinus hedenborgi (Col: Curculionidae), a trehala\u2010constructing weevil","volume":"22","author":"Skuhrovec J.","year":"2022","journal-title":"Organ. Divers. Evol."},{"key":"e_1_2_9_108_1","first-page":"618","volume-title":"Immature Insects","author":"Smith D.R.","year":"1987"},{"key":"e_1_2_9_109_1","volume-title":"Immature Insects Vol. 1","author":"Stehr F.","year":"1987"},{"key":"e_1_2_9_110_1","volume-title":"Immature Insects Vol. 2","author":"Stehr F.","year":"1991"},{"key":"e_1_2_9_111_1","doi-asserted-by":"crossref","first-page":"83","DOI":"10.1080\/20961790.2018.1428078","article-title":"Chrysomya chani Kurahashi (Diptera: Calliphoridae), a blow fly species of forensic importance: morphological characters of the third larval instar and a case report from Thailand","volume":"3","author":"Sukontason K.L.","year":"2018","journal-title":"Forensic Sci. Res."},{"key":"e_1_2_9_112_1","doi-asserted-by":"crossref","first-page":"447","DOI":"10.1038\/46737","article-title":"The origins of insect metamorphosis","volume":"401","author":"Truman J.W.","year":"1999","journal-title":"Nature"},{"key":"e_1_2_9_113_1","article-title":"Morphology and biological notes of Macrosiagon octomaculata (Gerstaecker) triungulinid and a summary of the knowledge on immature stages of the genus (Ripiphoridae)","volume":"60","author":"Ulyss\u00e9a M.A.","year":"2020","journal-title":"Pap. Avulsos Zool."},{"issue":"1","key":"e_1_2_9_114_1","doi-asserted-by":"crossref","first-page":"165","DOI":"10.14411\/eje.2004.021","article-title":"Larval morphology of Heterogynis (Lepidoptera: Heterogynidae)","volume":"101","author":"Vegliante F.","year":"2004","journal-title":"Eur. J. Entomol."},{"key":"e_1_2_9_115_1","doi-asserted-by":"crossref","first-page":"393","DOI":"10.1111\/j.1096-3642.2001.tb01320.x","article-title":"Phylogeny and classification of the extant basal lineages of the Hymenoptera (Insecta)","volume":"131","author":"Vilhelmsen L.","year":"2001","journal-title":"Zool. J. Linn. Soc."},{"key":"e_1_2_9_116_1","first-page":"346","article-title":"Larval anatomy of Orussidae (Hymenoptera)","volume":"12","author":"Vilhelmsen L.","year":"2003","journal-title":"J. Hymenopt. Res."},{"key":"e_1_2_9_117_1","article-title":"Xenos vesparum (Strepsiptera: Xenidae)\u2014A new insect model and its endoparasitic secondary larva","volume":"7","author":"Weingardt M.","year":"2023","journal-title":"Ins. Syst. Div."},{"key":"e_1_2_9_118_1","first-page":"497","article-title":"Funktionsanatomie des Larvenkopfes von Ctenocephalis felis felis B","volume":"92","author":"Widhalm\u2010Finke S.","year":"1974","journal-title":"Zool. Jb., Abt. Anat."},{"key":"e_1_2_9_119_1","doi-asserted-by":"crossref","DOI":"10.1186\/1741-7007-7-34","article-title":"Single\u2010copy nuclear genes resolve the phylogeny of the holometabolous insect orders","volume":"7","author":"Wiegmann B.M.","year":"2009","journal-title":"BMC Biol."},{"key":"e_1_2_9_120_1","first-page":"253","volume-title":"Immature Insects, Vol. I","author":"Wiggins G.B.","year":"1987"},{"key":"e_1_2_9_121_1","doi-asserted-by":"crossref","first-page":"112","DOI":"10.1002\/mmnd.19590060112","article-title":"Die Muskulatur der Larve von Limnophilus flavicornis Fabr","volume":"6","author":"Winkler D.","year":"1959","journal-title":"Dtsch. Entomol. Z. (N.F.)"},{"key":"e_1_2_9_122_1","doi-asserted-by":"crossref","first-page":"241","DOI":"10.1111\/j.1365-3113.2010.00556.x","article-title":"On the head morphology of Grylloblattodea (Insecta) and the systematic position of the order, with a new nomenclature for the head muscles of Dicondylia","volume":"36","author":"Wipfler B.","year":"2011","journal-title":"Syst. Entomol."},{"key":"e_1_2_9_123_1","doi-asserted-by":"crossref","first-page":"47","DOI":"10.1016\/j.asd.2012.09.005","article-title":"The skeletomuscular system of the larva of Drosophila melanogaster (Drosophilidae, Diptera) \u2013 A contribution to the morphology of a model organism","volume":"42","author":"Wipfler B.","year":"2013","journal-title":"Arthropod Struct. 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