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A stem-group amphiesmenopteran, lepidopterans, and a trichopteran. (A), A stem-group amphiesmenopteran, Necrotaulius tener , modified after Grimaldi and Engel [5]. (B-F), Lepidopteran forewings. (B), Archaeolepis mane , from Grimaldi and Engel [5]. (C), Forewing of an Early Jurassic Germany lepidopteran, from Grimaldi and Engel [5]. (D), Undopterix sukatshevae , modified after Skalski [10]; the red arrowhead showing Rs 3 + 4 furcation beyond the M 1 + 2 furcation. (E), Netoxena nana , modified after Martins-Neto [24]; red line indicates a single-Y configuration. (F), Mesokristensenia sinica , modified after Huang et al . [25]. (G), A trichopteran, Juraphilopotamus lubricus , modified after Gao et al . [43]. doi:10.1371/journal.pone.0079500.g001 

A stem-group amphiesmenopteran, lepidopterans, and a trichopteran. (A), A stem-group amphiesmenopteran, Necrotaulius tener , modified after Grimaldi and Engel [5]. (B-F), Lepidopteran forewings. (B), Archaeolepis mane , from Grimaldi and Engel [5]. (C), Forewing of an Early Jurassic Germany lepidopteran, from Grimaldi and Engel [5]. (D), Undopterix sukatshevae , modified after Skalski [10]; the red arrowhead showing Rs 3 + 4 furcation beyond the M 1 + 2 furcation. (E), Netoxena nana , modified after Martins-Neto [24]; red line indicates a single-Y configuration. (F), Mesokristensenia sinica , modified after Huang et al . [25]. (G), A trichopteran, Juraphilopotamus lubricus , modified after Gao et al . [43]. doi:10.1371/journal.pone.0079500.g001 

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The early history of the Lepidoptera is poorly known, a feature attributable to an inadequate preservational potential and an exceptionally low occurrence of moth fossils in relevant mid-Mesozoic deposits. In this study, we examine a particularly rich assemblage of morphologically basal moths that contribute significantly toward the understanding o...

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... 195 Ma) of England (Whalley 1985(Whalley , 1986Sohn et al. 2015). While these body-fossil discoveries clearly indicate a Jurassic lepidopteran diversification, the diversity and abundance of lepidopteran specimens remain low throughout the Mesozoic Era (Zhang et al. 2013). A survey of lepidopteran fossil specimens from the Jurassic and Cretaceous periods found a total of 177 specimens (100 body fossils, 77 ichnofossils), which accounts for approximately 3% of all identified lepidopteran fossil specimens . ...
Article
A new fossil leaf mine ichnogenus and species, Leucopteropsa spiralae gen. et sp. nov. (Lepidoptera: Lyonetiidae), from the Upper Cretaceous Kaiparowits Formation (Campanian age, c. 76.6 to 74.5 Ma) in Utah, USA, is the earliest record (75.6 ± 0.18 Ma) and only fossil evidence of a lyonetiid-like leaf-mining moth, as well as one of the oldest known fossils within the Yponomeutoidea-Gracillarioidea clade. The blotch-style mine consists of a central oviposition, or egg laying, site with a spiral trail packed with faecal pellets that concentrically spirals outwards. The trail increases in width from oviposition site to terminus, the trajectory of the trail does not cross itself, and there is no apparent pupation chamber present. The morphology of the fossil mine is reliably associated with Cemiostominae and is most similar to mines produced by extant members of the genus Leucoptera, such as the mountain ash bent-wing moth, Leucoptera malifoliella, and the laburnum leaf miner L. laburnella. The leaf-mining moth responsible for L. spiralae may be an early member of the genus Leucoptera or other Cemiostominae genera, and the new ichnogenus Leucopteropsa is erected because the morphology, and therefore the taxonomy, of the moth specifically responsible for L. spiralae mines was not preserved in association with the leaf mine. Despite the phylogenetic analysis of 15 lyonetiids undertaken in this study and the need for further phylogenetic work for the Lepidoptera, particularly the placement of the family Lyonetiidae and the subfamily Cemiostominae within the larger Yponomeutoidea-Gracillarioidea phylogeny, this fossil provides an important Late Cretaceous (∼76 Ma) calibration point for lepidopteran phylogeny and serves as an indicator for the antiquity of the most diverse lepidopteran group, Ditrysia. Because lepidopteran body fossils are extremely rare, owing to their small and lightly sclerotized bodies, this discovery also underscores the importance of ichnofossils in the lepidopteran fossil record. http://zoobank.org/urn:lsid:zoobank.org:pub:52B20B15-5C1A-4916-9096-A490DFED7D1C
... and in the absence of an epiphysis (Huang et al. 2010). The latter differs from Agathiphagidae and Mesokristenseniidae by lacking medial spurs on the metatibiae (Zhang et al. 2013). ...
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We report some surprising recent distributional range extensions of one extant genus and two more families of primitive moths discovered in amber fossils from Southeast Asia which were previously only known
... Among extant insects, the first four of these are still among the most species-diverse orders. In contrast, only 18 species of Lepidoptera have been reported from the Mesozoic of Northeastern China (29,99); Lepidoptera diversified significantly 18 Five orders, Odonata (dragonflies and damselflies), Blattaria (cockroaches), Orthoptera (grasshoppers and katydids), Plecoptera (stoneflies), and Mecoptera (scorpionflies), have moderately high numbers of described species (Figure 1). A member of an extinct group of Odonata, Hsiufua chaoi, from the Yanliao Entomofauna was considered to be the largest known fossil dragonfly in China; it has a forewing length of 107.6 mm (98). ...
Article
During the past 20 years, more than 1,600 species of well-preserved fossil insects, including members of over 270 families within 24 orders, have been described from the Middle Jurassic Yanliao Entomofauna and Early Cretaceous Jehol Entomofauna in Northeastern China. Diversified fossil insects not only document the origin, systematics, and early evolution of many lineages, but also reveal these lineages’ behaviors and interactions with coexisting plants, vertebrates, and other insects in their ecosystems. For example, fossil evidence has been documented, for example, regarding insects’ feeding and pollination mutualism with gymnosperms; ectoparasitic feeding on blood of vertebrates; camouflage, mimicry of gymnosperm plants, and eyespot warning; sound stridulation for attracting potential mates; and sexual display, mating, egg-laying, and parental care. In this article, we review the diverse taxonomy of mid-Mesozoic insects of Northeastern China and elucidate their behaviors and interactions within their ecosystems, which have impacted their early evolution and development into extant insects. Expected final online publication date for the Annual Review of Entomology, Volume 66 is January 11, 2020. Please see http://www.annualreviews.org/page/journal/pubdates for revised estimates.
... Eolepidopterigidae, Mesokristenseniidae and Ascololepidopterigidae, and an extant family, vis. Micropterigidae (Huang et al., 2010;Zhang et al., 2013Zhang et al., , 2015Zhang et al., , 2017. Micropterigidae, as a small family within Zeugloptera, are the most primitive type of surviving Lepidoptera (Gibbs, 2014). ...
Article
A new genus and species, Archmosaicus comitatus gen. et sp. nov., assigned to the family Micropterigidae, is described based on three well-preserved specimens from the mid-Cretaceous Burmese amber. It is established mainly based on antennal scape and pedicel greatly swollen, scape with a strong indentation on mesal surface, forewing with veins Sc and R deeply forked, and hind wing without free R. The minor differences on these three specimens could be attributable to either individual variation or sexual dimorphism.
... On the basis of our estimates, the last common ancestor of extant Lepidoptera occurred approximately 271 Ma (95% CI 208-331 Ma). This is slightly earlier than the first glossatan fossilized wing scales, recently discovered in an Upper Triassic deposit [73], but well before the first fossils reliably identified as members of the Lepidoptera dated to the Early Jurassic [74][75][76][77]. Therefore, suctorial mouthparts for feeding on gymnosperm pollination drops might have evolved earlier than generally thought. ...
Article
Insects are a highly diverse group of organisms and constitute more than half of all known animal species. They have evolved an extraordinary range of traits, from flight and complete metamorphosis to complex polyphenisms and advanced eusociality. Although the rich insect fossil record has helped to chart the appearance of many phenotypic innovations, data are scarce for a number of key periods. One such period is that following the End-Permian Extinction, recognized as the most catastrophic of all extinction events. We recently discovered several 240-million-year-old insect fossils in the Mount San Giorgio Lagerstätte (Switzerland-Italy) that are remarkable for their state of preservation (including internal organs and soft tissues), and because they extend the records of their respective taxa by up to 200 million years. By using these fossils as calibrations in a phylogenomic dating analysis, we present a revised time scale for insect evolution. Our date estimates for several major lineages, including the hyperdiverse crown groups of Lepidoptera, Hemiptera: Heteroptera and Diptera, are substantially older than their currently accepted post-Permian origins. We found that major evolutionary innovations, including flight and metamorphosis, appeared considerably earlier than previously thought. These results have numerous implications for understanding the evolution of insects and their resilience in the face of extreme events such as the End-Permian Extinction.
... The Jiulongshan Formation belongs to the well-recognized Yanliao Biota, which is known for yielding feathered dinosaurs, mammals, conifers, and numerous insect taxa such as Coleoptera (Chang et al., 2009), Trichoptera (Liu et al., 2014), Lepidoptera (Zhang et al., 2013), Grylloblattodea (Cui & Ren, 2013), Diptera (Shi et al., 2015) and Hymenoptera (Li et al., 2014b(Li et al., , 2015a, among others. Based on Ar-Ar and SHRIMP U-Pb dating, the Jiulongshan Formation corresponds to uppermost strata of the Middle Jurassic. ...
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The phylogeny of the superfamily Evanioidea is presented using morphology and DNA sequence data of selected extant and fossil genera by employing two phylogenetic methods, maximum parsimony and Bayesian inference. Based on our new results, the monophyly of Evanioidea is corroborated. Evanioidea, Anomopterellidae, Othniodellithidae, Andreneliidae and Evaniidae are monophyletic families, while Praeaulacidae, Aulacidae, Baissidae and Gasteruptiidae are paraphyletic families. In addition, four new genera (Sinuevania gen.n., Curtevania gen.n., Exilaulacus gen.n., Heterobaissa gen.n.) with five new species (Sinuevania mira sp.n., Curtevania enervia sp.n., Exilaulacus loculatus sp.n., Exilaulacus latus sp.n., Heterobaissa apetiola sp.n.), and five additionally new species (Newjersevania longa sp.n., Newjersevania brevis sp.n., Cretevania tenuis sp.n., Cretevania venae sp.n., Praeaulacus rectus sp.n.) and one new combination [Cretevania mitis (Li, Shih & Ren, 2014a) comb.n.] are described based on well‐preserved fossils from the Middle Jurassic Jiulongshan Formation in Inner Mongolia, China, the Early Cretaceous Yixian Formation in Liaoning, and mid‐Cretaceous amber from Myanmar. This study documents the diversification of one major lineage of the mid‐Mesozoic parasitoid revolution that dramatically changed food‐web relationships in terrestrial ecosystems. This published work has been registered in ZooBank, http://zoobank.org/urn:lsid:zoobank.org:pub:CBC04ADA‐0176‐402D‐9B43‐E1B3CDA080E1.
... In addition, we investigated 16 well-preserved lepidopterans from the Middle Jurassic of Daohugou, China. Although fossil insects from Daohugou are famous for their excellent macroscopic preservation (39,40), the Daohugou lepidopterans (Mesokristenseniidae) preserve only faint traces of wing scales ( fig. S1, F and G). ...
... In addition, we investigated 16 well-preserved lepidopterans from the Middle Jurassic of Daohugou, China. Although fossil insects from Daohugou are famous for their excellent macroscopic preservation (39,40), the Daohugou lepidopterans (Mesokristenseniidae) preserve only faint traces of wing scales ( fig. S1, F and G). ...
Article
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Lepidopteran scales exhibit remarkably complex ultrastructures, many of which produce structural colors that are the basis for diverse communication strategies. Little is known, however, about the early evolution of lepidopteran scales and their photonic structures. We report scale architectures from Jurassic Lepidoptera from the United Kingdom, Germany, Kazakhstan, and China and from Tarachoptera (a stem group of Amphiesmenoptera) from mid-Cretaceous Burmese amber. The Jurassic lepidopterans exhibit a type 1 bilayer scale vestiture: an upper layer of large fused cover scales and a lower layer of small fused ground scales. This scale arrangement, plus preserved herringbone ornamentation on the cover scale surface, is almost identical to those of some extant Micropterigidae. Critically, the fossil scale ultrastructures have periodicities measuring from 140 to 2000 nm and are therefore capable of scattering visible light, providing the earliest evidence of structural colors in the insect fossil record. Optical modeling confirms that diffraction-related scattering mechanisms dominate the photonic properties of the fossil cover scales, which would have displayed broadband metallic hues as in numerous extant Micropterigidae. The fossil tarachopteran scales exhibit a unique suite of characteristics, including small size, elongate-spatulate shape, ridged ornamentation, and irregular arrangement, providing novel insight into the early evolution of lepidopteran scales. Combined, our results provide the earliest evidence for structural coloration in fossil lepidopterans and support the hypothesis that fused wing scales and the type 1 bilayer covering are groundplan features of the group. Wing scales likely had deep origins in earlier amphiesmenopteran lineages before the appearance of the Lepidoptera.
... These three primitive families represent relict lineages of small moths with mandibulate, chewing mouthparts (14). Morphologically related Jurassic fossils have been included in four extinct mandibulate families: Archaeolepidae, Eolepidopterygidae, Mesokristenseniidae, and Ascololepidopterygidae (15). Only the Archaeolepidae, with the singlewing species A. mane, are characterized by the presence of relatively well-preserved, probably solid, wing scales, but no herringbone pattern has been observed (16). ...
Article
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On the basis of an assemblage of fossilized wing scales recovered from latest Triassic and earliest Jurassic sediments from northern Germany, we provide the earliest evidence for Lepidoptera (moths and butterflies). The diverse scales confirm a (Late) Triassic radiation of lepidopteran lineages, including the divergence of the Glossata, the clade that comprises the vast multitude of extant moths and butterflies that have a sucking proboscis. The microfossils extend the minimum calibrated age of glossatan moths by ca. 70 million years, refuting ancestral association of the group with flowering plants. Development of the proboscis may be regarded as an adaptive innovation to sucking free liquids for maintaining the insect’s water balance under arid conditions. Pollination drops secreted by a variety of Mesozoic gymnosperms may have been non-mutualistically exploited as a high-energy liquid source. The early evolution of the Lepidoptera was probably not severely interrupted by the end-Triassic biotic crisis.
... What remains to be evaluated is whether the family can be associated with one of the fossil families described in Lepidoptera or Trichoptera. The genera and families of Mesozoic Lepidoptera are summarised in Zhang et al. (2013). In addition to the Micropterigidae, there are four families that have been assigned to Lepidoptera: Archaeolepidae; Eolepidopterigidae; Mesokristenseniidae; and Ascololepidopterigidae. Concerning the Eolepidopterigidae, the suborder Eolepidopterigina was established (Rasnitsyn 1983). ...
... This character, however, is absent in females of Micropterigidae and extant Heterobathmiidae and is, therefore, of some taxonomic value. The other families are treated as suborder incertae sedis (Huang et al. 2010;Zhang et al. 2013). ...
Article
A small fossil insect with scales on the wings and body was identified as a representative of Aphiesmenoptera from Burmese amber. The species is introduced here as Tarachocelis microlepidopterella (†). The insect is described in detail, and photos and line drawings are provided for wing venation, head, mouthparts, scales, legs and abdomen. All characters shared with primitive Lepidoptera and Trichoptera are symplesiomorphies or groundplan traits of Amphiesmenoptera. In addition, the Burmese fossil has a number of remarkable autapomorphies, giving it an appearance that deviates clearly from known families of Lepidoptera and Trichoptera. The species, representing a family of its own, Tarachocelidae, is considered a separate and unique taxon in the stem-group of Amphiesmenoptera and is provisionally placed as Amphiesmenoptera incertae sedis .* * NB : See Note Added in Proof (Section 4).