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Abstract Applying BEAST v1.10.4, we constructed a Bayesian Inference tree comprising 322 taxa, primarily representing Paleoptera (Odonata and Ephemeroptera; Pterygota), Zygentoma and Archaeognatha (Apterygota; paraphyly), and Neoptera (Plecoptera; Pterygota), based on a 2685 bp sequence dataset. Our analyses revealed that robust dating required the incorporation of both Quaternary and pre‐Quaternary dates. To achieve this, our dating incorporated a 1.55 Ma (Quaternary) geological event (the formation of the Ryukyu Islands) and a set of chronologically well‐founded fossil dates, spanning from up to 400 Ma (Devonian) for the stem Archaeognatha, 320 Ma (Carboniferous) for the crown of Paleoptera, 300 Ma (Carboniferous) for the crown Ephemeroptera, and 280 Ma (Permian) for the crown Odonata, down to 1.76 Ma (Quaternary) for Calopteryx japonica, encompassing a total of 22 calibration points (events: 6, fossils: 16; Quaternary: 7, pre‐Quaternary: 15). The resulting dated tree aligns with previous research, albeit with some dates being overestimated. This overestimation was mainly due to the lack of Quaternary calibration and the exclusive dependence on pre‐Quaternary calibration, though the application of maximum age constraints also played a role. Our minimum age dating demonstrates that the molecular clock did not uniformly progress, rendering rate dating an inapplicable approach. We observed that the base substitution rate is time‐dependent, with an exponential increase evident from around 20 Ma (Miocene) to the present time, exceeding an order of magnitude. The extensive radiation and speciation of Insecta and Paleoptera, potentially resulting from the severe climatic changes associated with the Quaternary, including the commencement of glacial and interglacial cycles, may have significantly contributed to this increase in base substitution rates. Additionally, we identified a potential peak in base substitution rates during the Carboniferous period, around 320 million years ago, possibly corresponding to the Late Paleozoic Ice Age.

The order Psocodea, including barklice, booklice, and parasitic lice, is diverse and widely distributed since the Cretaceous. That is particularly the case for the speciose extinct family Empheriidae (Psocodea, Trogiomorpha, Atropetae), recently fused with the ‘Archaeatropidae’. Understanding the evolution of barklice is dependent in part on studying this family, as its representatives have been found from the Early Cretaceous to the Eocene, surviving the K/Pg extinction event. The phylogenetic relationships of Empheriidae in relation to other families, such as Lepidopsocidae or Psoquillidae, have been extensively debated. However, distinguishing diagnostic characters for the Empheriidae has proven challenging. In this study, we describe the new empheriid Santonipsocus mimeticus gen. et sp. nov. from Cretaceous Charentese amber (France). It is the third empheriid species found in this locality. The new genus is compared with the other genera in the family, and Proprionoglaris guyoti and Proprionoglaris axioperierga are revised based on the type material and new specimens. We explore the phylogeny of Empheriidae, both the relationships with other families and the inner relationships between the genera, through maximum parsimony analysis and Bayesian inference analysis. Our results suggest that Empheriidae may represent a paraphyletic evolutionary grade to the rest of Atropetae. The phylogenetic relationships between genera align with the biogeography of the family and support previous hypotheses. In addition, we discuss the possible biology of the members of the family, shedding light on the evolutionary history of Empheriidae.

Studying insect fossils, particularly those preserved as compressions in sedimentary matrices, can be difficult due to the taphonomic processes that often result to poor preservation and contrast of structures compared to the embedding matrix. To address this, we propose a user-friendly and simple methodology based on UV-light to study insect fossils and select specimens of interest for more advanced imagery exploration. While UV-light imaging has been previously applied to compressions of arthropod fossils, it typically involved laser light sources. Our approach allows the investigation of fossils using an affordable, compact, and portable UV-light source, along with a simple and replicable low-cost protocol. • The methodology is based on UV-light induced natural fluorescence of sediment and fossil remains. • UV-light is effective on compression fossils to gain natural contrast and enhance observation of body structures like veins or setae on wings. • UV-light is effective to reveal palaeoecological information such as pollen grains preserved on specimens, especially near or on putative pollinator or pollen-eating taxa.

The first fossil limoniid fly from the Miocene Fossil-Lagerstätte of Foulden Maar in New Zealand is described on the basis of an isolated well-preserved wing. The specimen is tentatively attributed to a new species Gynoplistia fouldensensis sp. nov. in the large extant genus Gynoplistia, which is well diversified in the country. It is the second fossil record of this genus, the first one being an isolated wing from the Cretaceous Weald Clay Formation in the United Kingdom.

Fossil insects are valuable indicators of the evolutionary history of the clades to which they belong. According to their state of preservation, fossil insects are often partially described for key morphological characters, such as forewing venation in crickets (Orthoptera, Grylloidea). In parallel, the use of 3D microtomography is increasingly becoming common for studying some fossils, which allowed here the precise reconstruction and interpretation of the venation pattern in the smallest known cricket with a stridulatory apparatus, †Picogryllus carentonensis, found in opaque amber. The 3D reconstructions have revealed the general structure of the venation of the forewing and have enabled the identification of all its veins and cells, validating its similarity with that of extant crickets. Putative homologies are established according to previous studies, and some particularities are observed, such as the presence of two crossveins in the mirror, a rare feature in extant crickets that is discussed in the frame of cricket venation evolution. These findings highlight the importance of 3D microtomography as a powerful tool for examining fossil insects and also provide crucial information for taxonomic identification and evolutionary studies, offering a validated morphological basis for future phylogenetic analyses incorporating fossils.

The erasipterid Sinoerasipteron xiaheyanensis Nel & Huang gen. et sp. nov. from the Moscovian Tupo Formation in Xiaheyan locality (China), is described and illustrated. It is the sixth species of the odonatopteran griffenflies from this locality. This new discovery confirms the high diversity of these flying predators in the insect assemblage.

Abstract While not generally used for relatively flat structures, X‐ray microtomography can provide decisive insights for comparative anatomy and evolutionary studies of flat structures, thereby solving long‐standing ecological and evolutionary issues. This is particularly true for the study of insect wings, organs that have played a central role in the evolution and ecological success of insects, the first organisms to develop powered flight. We present the challenges inherent in studying insect wings by microtomography, as well as solutions to overcome them. We highlight the impact of insect preparation (i.e. dry or ethanol‐preserved specimens, complete specimens or only part of the wings, wings stained with iodine or with a 0.3% solution of phosphotungtic acid—PTA) and of the acquisition mode (phase contrast or not), on scan quality and ease of interpretation. We compare our results with those obtained by more ‘traditional’ techniques, based on examination of the wings with a binocular or SEM microscope. We show that X‐ray microtomography is effective in establishing vein homologies—a crucial step in many evolutionary studies in insects—by accessing the internal structures of the wing base itself, where veins originate. It can also be used for difficult tasks such as distinguishing between crossveins and main veins, establishing vein polarity (i.e. concave vs. convex veins) or vein fusions, situations that complicate the establishment of vein homology hypotheses and the completion of subsequent evolutionary studies. Unstained, ethanol‐preserved specimens give the least conclusive results, while dry specimens and ethanol‐preserved wings stained with PTA give the best results. We show that 3D tools provide crucial insights for the study of insect wings, illustrating how CT‐scan tools may contribute to a change in dimension, introducing quantification and functional approaches and hopefully stimulating new studies for other seemingly 2D organs in several other fields, including plant studies. We highlight, nonetheless, that this tool must be used in coordination with more traditional observational techniques, at least for insect wings, to provide a more complete picture of these fundamental organs.

The new “libelluloid” family Cordulibellulidae is described to accommodate a new genus and species Cordulibellula inopinata from the Eocene Green River Formation, Utah, USA. Even if its affinities remain somewhat uncertain because of the lack of information on structures others than those of the hind wing, ts putative closest relatives are known from the Late Jurassic and Early Cretaceous. Therefore, this new fossil is interpreted as a “relict” taxon surviving until the Paleogene. This new addition to the odonatan paleoentomofauna of the Green River Formation confirms the high diversity of this clade during the Paleogene and in this formation.

Palaeosphryon menatensis gen. et sp. nov., first unambiguous representative of the longhorn beetle subfamily Prioninae from the Paleocene of Menat (France), is described and illustrated. The new fossil is placed into the tribe Prionini, showing some similarities with some species of the extant genera Osphryon (Papua New Guinea) and Titanus (Brazil, Colombia, Guianas, Ecuador, Peru), viz. in general body shape, antennomere 3 as long as first and second together but shorter than the length of fourth plus fifth, elongate elytra, and small spines on the lateral margin of the pronotum disposed in a relatively similar way as in Osphryon. Nevertheless, the exact affinities of the new fossil within the Prionini remain uncertain because of the lack of a recent phylogenetic analysis in which it could be integrated. This fossil beetle is exceptional for its very large size, with a body 70 mm long. Some other large longhorn beetles have been found in the same outcrop, and are awaiting description. The positions of the previously described Cerambycidae from Menat are also discussed. This exceptional fauna of Cerambycidae is in accordance with the current palaeoenvironmental reconstruction for the Menat Konservat-Lagerstätte, as a small maar lake surrounded by a warm and humid, probably evergreen forest.

Luberotenthredo cerestensis gen. et sp. nov. is the first record of the sawfly family Tenthredinidae from the Oligocene of Céreste (Southern France). This taxon is described and illustrated based on a well-preserved specimen. This genus resembles the extant genus Perineura (subfamily Tenthredininae, tribe Perineurini) with which it shares forewing venation similarities and numerous morphological characters. This new taxon is the first fossil representative of the tribe Perineurini and can be used as a calibration point for future investigation of the diversification of the family Tenthredinidae.

The impact of three extinction events during the Permo–Triassic interval on terrestrial invertebrates is unclear. Here, the authors find that key abiotic and biotic factors, including changes in floral assemblages, were correlated with changes in insect diversity through this interval.

A new middle Permian fossil locality featuring a large abundance of exceptionally preserved plants and insects is identified in the Karoo Basin of South Africa.

The current knowledge on the Paleocene Odonata is rather limited despite the fact that it is a crucial period for the history of this order. An overview of the fossil odonatans from the Paleocene of Menat (France) is provided. We describe the anisopteran Macrogomphus menatensis sp. nov., first fossil representative of the family Epigomphidae, together with two zygopteran, viz. the dysagrionid Menatagrion hervetae gen. et sp. nov., and the new family Menatlestidae fam. nov., with its type species Menatlestes palaeocenicus gen. et sp. nov. The genus Menatagrion gen. nov. is the first Paleocene record of the Dysagrionidae, otherwise known by a putative Cretaceous genus and several Eocene to Miocene genera. Menatlestes gen. nov., putatively attributed to the stem-group of the Lestinoidea (Megalestidae and Lestidae), would correspond to the oldest record of this clade. With these three new taxa, and the previously described Thanetophilosina menatensis, Valerea multicellulata, “Lestes” zalesskyi, and an Aeshna species indet., the total number of Odonata from Menat goes up to seven species in total; two Anisoptera and five Zygoptera. Furthermore, we propose new evidences showing that the head characters defining the putative suborder Cephalozygoptera are due to deformations, very frequent among the fossil Odonatoptera. We treat the Cephalozygoptera as a junior synonym of Zygoptera.

Oligoptilomera luberonensis gen. et sp. nov., first fossil representative of the gerrid subfamily Ptilomerinae, is described and figured from the Oligocene of Murs (Vaucluse, Southern France). Extant Ptilomerinae live in streams in warm climates, of the Indo-Malaysian, eastern Palaearctic, and Papouan regions. The discovery of this Oligocene French Ptilomerinae is in accordance with the putative age of the subfamily, at least older than the Eocene, and with the Indo-Malaysian affinities previously recorded for some other insects from the Oligocene of France. The two insect assemblages of Murs and Céreste are compared and the differences discussed. Although of similar ages, that from Murs was possibly corresponding to a more shallow water paleolake than that of Céreste.

The new archaeorthopteran family Archaeogeraridae, based on Archaeogerarus schubneli gen. et sp. nov., is described from the Moscovian of Avion. It has several unique wing venation characters allowing to separate it from the other groups of this superorder, viz. a vein RP separating from RA very close to wing base and closely parallel to it; M shortly fused with RP; CuA+CuPa simple and straight; and CuPb branched. This last character is convergently present in a few Palaeozoic Panorthoptera, a feature of great interest for a future phylogenetic analysis of the whole superorder. Avionxixia gui gen. et sp. nov., second Cnemidolestidae from the Moscovian of Avion, is described and illustrated. It shares numerous characters with the Chinese Namurian genus Xixia and the European (Germany and France) Moscovian genus Piesbergopterum, suggesting possible phylogenetic affinities between these three genera. It is mainly separated from these two genera by the costal area much narrower than the subcostal one and the presence of only one posterior branch of the anterior branch of MP+CuA+CuPa. This new taxon confirms the high diversity of the Cnemidolestidae during the late Carboniferous.

Lodevoisadia coheni gen. et sp. nov. is described as the ninth species of ‘Grylloblattodea’ from the middle Permian of the Salagou Formation, near Lodève town (France). It is currently not reasonable to place this species into a specific family, even though it seems to share most characters with the small family Tunguskapteridae. The lack of phylogenetic analysis and the current poor delineation of the majority of the grylloblattodean families (lacking synapomorphies) render any attribution of new taxa to a particular family often uncertain.

Abstract Full body impressions and resting traces of Hexapoda can be of extreme importance because they bring crucial information on behavior and locomotion of the trace makers, and help to better define trophic relationships with other organisms (predators or preys). However, these ichnofossils are much rarer than trackways, especially for winged insects. Here we describe a new full-body impression of a winged insect from the Middle Permian of Gonfaron (Var, France) whose preservation is exceptional. The elongate body with short prothorax and legs and long wings overlapping the body might suggests a plant mimicry as for some extant stick insects. These innovations are probably in relation with an increasing predation pressure by terrestrial vertebrates, whose trackways are abundant in the same layers. This discovery would possibly support the recent age estimates for the appearance of phasmatodean-like stick insects, nearly 30 million years older than the previous putative records. The new exquisite specimen is fossilized on a slab with weak ripple-marks, suggesting the action of microbial mats favoring the preservation of its delicate structures. Further prospections in sites with this type of preservation could enrich our understanding of early evolutionary history of insects.

Abstract New Caledonia was, until recently, considered an old continental island harbouring a rich biota with outstanding Gondwanan relicts. However, deep marine sedimentation and tectonic evidence suggest complete submergence of the island during the latest Cretaceous to the Paleocene. Molecular phylogenies provide evidence for some deeply-diverging clades that may predate the Eocene and abundant post-Oligocene colonisation events. Extinction and colonization biases, as well as survival of some groups in refuges on neighbouring paleo-islands, may have obscured biogeographic trends over long time scales. Fossil data are therefore crucial for understanding the history of the New Caledonian biota, but occurrences are sparse and have received only limited attention. Here we describe five exceptional fossil assemblages that provide important new insights into New Caledonia’s terrestrial paleobiota from three key time intervals: prior to the submersion of the island, following re-emergence, and prior to Pleistocene climatic shifts. These will be of major importance for elucidating changes in New Caledonia’s floristic composition over time.

Schubnel, Nel and colleagues present the first Carboniferous representative of Titanoptera, Theiatitan azari. Specially modified wing zones suggest that Titanoptera were capable of wing-based communication using light or sound, as seen in modern species.

Abstract Ghost lineages have always challenged the understanding of organism evolution. They participate in misinterpretations in phylogenetic, clade dating, biogeographic, and paleoecologic studies. They directly result from fossilization biases and organism biology. The Cylindrotomidae are a perfect example of an unexplained ghost lineage during the Mesozoic, as its sister family Tipulidae is already well diversified during the Cretaceous, while the oldest Cylindrotomidae are Paleogene representatives of the extant genus Cylindrotoma and of the enigmatic fossil genus Cyttaromyia. Here we clarify the phylogenetic position of Cyttaromyia in the stem group of the whole family, suggesting that the crown group of the Cylindrotomidae began to diversify during the Cenozoic, unlike their sister group Tipulidae. We make a comparative analysis of all species in Cyttaromyia, together with the descriptions of the two new species, C. gelhausi sp. nov. and C. freiwaldi sp. nov., and the revision of C. obdurescens. The cylindrotomid biogeography seems to be incongruent with the phylogenetic analysis, the apparently most derived subfamily Stibadocerinae having apparently a ‘Gondwanan’ distribution, with some genera only known from Australia or Chile, while the most inclusive Cylindrotominae are Holarctic.

In several insect eusocial lineages, e.g., some aphids, thrips, ants, some stingless bees, and termites, task specialization is brought to its climax with a sterile soldier caste solely devoted to colony defense. In Isoptera, while the reproductives are defenseless, the soldiers have unique morpho-physiological specializations whose origin and evolution remain unresolved. Here we report on two instances of Cretaceous fossil termite reproductives belonging to different families († Valkyritermes inopinatus gen. et sp. nov. and an unpublished specimen from the Crato Formation), with intriguing phragmotic soldier-like heads and functional wings. These individuals, herein called Valkyries, are the first termite reproductives known with defensive features and suggest that phragmosis arose at least in the Early Cretaceous. Valkyries resemble modern neotenic soldiers except for their complete wings. Their discovery supports the hypothesis that the division between reproductive (indicated by the winged condition of Valkyries) and defensive tasks (indicated by the phragmotic head) has not always been complete in termite history. We explore two alternative scenarios regarding the origin of Valkyries (i.e., relatively recent and convergent origins vs. plesiomorphic condition) and discuss how they might relate to the development of soldiers. We argue that, in both cases, Valkyries likely evolved to face external threats, a selective pressure that could also have favored the origin of soldiers from helpers. Valkyries highlight the developmental flexibility of termites and illustrate the tortuous paths that evolution may follow.

Glaphyrophlebia victoiriensis sp. nov. (Paoliida: Blattinopsidae) is the third Gzhelian representative of the genus and is described based on a beautiful forewing from the Var department in Southern France. Together with the description of another forewing fragment of a Glaphyrophlebia sp. from the Province of León in NW Spain, they improve our knowledge of fossil insects from French and Spanish upper Carboniferous deposits. The specimen of Glaphyrophlebia sp. is the first mention of the family in the Carboniferous of Spain and extends the geographical distribution of the genus. These descriptions suggest that the genus Glaphyrophlebia was speciose during the Upper Pennsylvanian, while otherwise very diverse in the lower and middle Permian strata of the Russian Federation. We proposed the first hypothesis to explain the diversification of the family and of its most speciose genera and to argue that their diversity dynamics were likely linked with the major environmental changes that followed the collapse of the Carboniferous rainforest, notably the extension of arid biomes during the Permian period. The exquisite preservation and the fineness of the sediment from Tante Victoire, in which the new species was found, suggests that the locality is suitable for preserving other fossil insects and will require additional investigations.

The systematic status of the genus Athalia and related genera is a perennial controversy in sawfly taxonomy. Several authors have hypothesized that the placement of Athalia within the Tenthredinidae is artificial, but no studies have focused on this topic. If the hypothesis that Athalia does not belong to Tenthredinidae can be supported, the taxonomic framework of Tenthredinoidea needs revision. We present a comprehensive phylogenomic study of Tenthredinoidae, focusing on the positions of Athalia and related genera by sampling 80 representatives mainly of the Tenthredinoidea, including Heptamelinae and Blasticotomidae. Our phylogenetic reconstructions based on nuclear genes and mitochondrial (mt) sequences support Athalia and related genera as a distinct clade sister to Tenthredinidae + (Cimbicidae + Diprionidae). A comparison of symphytan mitochondrial genomes reveals an innovative gene rearrangement pattern in Athaliidae, in which Dentathalia demonstrates a more ancestral pattern than Athalia and Hypsathalia. The lineage specificity of mt rRNA secondary structures also provides sufficient support to consider Athaliidae as a separate family. In summary, the phylogeny and genomic structural changes unanimously support the taxonomic treatment of Athaliidae as a family and the re-establishment of Dentathalia as a valid genus.

Sysciophlebia ‘sp. form Villablino’, the first Iberian representative of the Palaeozoic–Early Mesozoic family Spiloblattinidae, is described and illustrated. Its forewing colour pattern is strongly similar to those of the Gzhelian–early-middle Asselian species Sysciophlebia euglyptica, Sysciophlebia ilfeldensis, Sysciophlebia rubida, and ‘Sysciophlebia sp. form KBQ’, supporting the currently proposed Gzhelian age for its type locality. It supports the use of the representatives of the Spiloblattinidae for stratigraphic purposes. The diagnoses and limits of the families Subioblattidae, Phyloblattidae, Compsoblattidae, Spiloblattinidae, and of the spiloblattinid genera are discussed.

Two new genera and species of Coniopterygidae (Neuroptera) are described and illustrated from mid Cretaceous (Cenomanian) amber of Myanmar. Mulleroconis hyalina gen. n. et sp. n., attributed to the Coniopteryginae, bears a unique combination of venation characters and an abdomen without plicatures. The second new genus, attributed to the Aleuropteryginae, i.e. Palaeoconis azari gen. n. et sp. n., displays a unique pattern of crossveins 1m-cua and 2mp2-cua, with the latter crossing the pigmented spot. A check-list of all fossil genera and species of Coniopterygidae is provided.

Triassopsyllidiida pectinata gen. et sp. nov. is described from the latest Middle Triassic Tongchuan biota of China and tentatively attributed to the superfamily Protopsyllidioidea. Its forewing venation is unique among this superfamily in the anteriorly pectinate vein ScP + RA and the presence of a veinlet between R and M + CuA. Its exact position in this group remains uncertain, mainly because of the weak diagnostic value of the wing venation characters in these insects. The phylogenetic relationships of the Protopsyllidioidea are discussed.

The taxa Cubicostissus palaeocaeni gen. et sp. nov. are described from the Paleocene of Menat (Central France) as the oldest representative of the family Issidae for which all currently known fossils are reviewed. With a unique combination of characters, the specimen represents the first fossil described for the tribe Hysteropterini. Its occurrence in the European Paleocene shows that its lineage, and therefore the Issidae, were already present at least some 60 Ma. It allows to discard the hypothesis of an Eocene origin for the family, which is in accordance with a recent molecular calibration of the family, reporting the tribe as old as the Upper Cretaceous.

Hydrothermal palaeoenvironments are very uncommon in Upper Jurassic and Lower Cretaceous deposits worldwide. We present new plant and insect remains from travertines formed during the Jurassic-Cretaceous boundary in northern Spain (Aguilar Fm., Palencia province). A total of 136 plant specimens and three insect wings were collected and studied. This entomofauna consists of dragonfly (Odonata) wings including Cymatophlebiidae and an undetermined new genus and species of Aktassiidae, representing the first report of these families for the Iberian Peninsula. The fossil flora shows different morphotypes of plants, which have been tentatively assigned to three different genera. The taphocoenosis of the flora was dominated by Bennettitales (98.5%) including cf. Pterophyllum sp., Ptilophyllum cf. acutifolium, Ptilophyllum cf. pecten, Ptilophyllum cf. pectiniformis and cf. Ptilophyllum sp., and the occasional presence of ferns (1.5%) represented by the taxon Cladophlebis cf. denticulata. The presence of the Anisoptera Cymatophlebia cf. longialata suggests a higher affinity for a Tithonian age of the studied site, and the anatomy and palaeogeographical distribution of this species suggest capacity to migrate for rather long distances. The floristic composition of the site differs remarkably from other Tithonian-Berriasian floras of the Iberian Peninsula. The presence of Odonata and the distinctive flora in (semi)arid conditions could be due to the hot-spring providing an environmental niche with constant conditions of warmth and humidity forming an ‘ecological oasis’.

Emphytusmiocenicus sp. nov., first fossil representative of this genus, is described from the upper Miocene of the diatom paleolake of Montagne d’Andance (Ardèche, France). Its placement is ensured by an in-depth comparison with all the extant and fossil genera of the subfamily Allantinae. The representatives of Emphytus are distributed in the Palearctic, Nearctic, and Oriental regions. Empria sammuti sp. nov., second representative of the latter genus, is described from the latest Miocene of the diatom paleolake of Sainte-Reine (Cantal, France). The placement of this new species is based on a detailed comparison with the extant genera of the tribe Empriini. The larvae of the extant Emphytus and Empria spp. are known to be phytophagous on angiosperm leaves of several families, all present as fossils in the taphocenoses of la Montagne d’Andance and Sainte-Reine. Emphytus miocenicus sp. nov. represents the oldest record of this genus and of its crown group, corroborating the estimate of a middle Eocene–middle Oligocene age for its stem group. Throughout our study, it appears that the first described fossil of the genus Empria, E. oligocaenica, from the Oligocene of Germany, needs to be revised and redescribed. It should preferably be treated until the revision as incertae sedis in Allantinae sensu lato.

The Oligocene Oligopipiza quadriguttata Nidergras, Hadrava, and Nel gen et sp. nov. is the first fossil Pipizinae found in the lacustrine outcrop of Céreste (South-East of France). It differs from the other Pipizinae in the male genitalia, with a surstylus without tooth and shorter than epandrium, and a long epandrium with a very deep and narrow median theca. It is compared to other extant and fossil Pipizinae. Its position in this clade is supported by its inclusion in previous morphological phylogenetic analysis of the Syrphidae. Palaeoecological inferences for the paleobiota of Céreste are made based on this taxon and point to the presence of a mixed forest. The taphonomy of these flies is discussed. They were probably embedded in surface microbial mats. The pollinator role of Oligopipiza quadriguttata is also discussed on the basis of the presence of pollen surrounding the fossil flies.

Abstract The majority of the analyses of the evolutionary history of the megadiverse class Insecta are based on the documented taxonomic palaeobiodiversity. A different approach, poorly investigated, is to focus on morphological disparity, linked to changes in the organisms’ functioning. Here we establish a hierarchy of the great geological epochs based on a new method using Wagner parsimony and a ‘presence/absence of a morphological type of mouthpart of Hexapoda’ dataset. We showed the absence of major rupture in the evolution of the mouthparts, but six epochs during which numerous innovations and few extinctions happened, i.e., Late Carboniferous, Middle and Late Triassic, ‘Callovian-Oxfordian’, ‘Early’ Cretaceous, and ‘Albian-Cenomanian’. The three crises Permian-Triassic, Triassic-Jurassic, and Cretaceous-Cenozoic had no strong, visible impact on mouthparts types. We particularly emphasize the origination of mouthparts linked to nectarivory during the Cretaceous Terrestrial Revolution. We also underline the origination of mouthparts linked to phytophagy during the Middle and the Late Triassic, correlated to the diversification of the gymnosperms, especially in relation to the complex ‘flowers’ producing nectar of the Bennettitales and Gnetales.

Many histerid beetles are necrophilous on carrion during both active and advanced stages of decomposition. In this study, 13 species of Saprinus were recorded on carrion from Lebanon, containing eight that are new for the Lebanese fauna. The following Saprinus species are newly recorded from Lebanon: 1) Saprinus (S.) caerulescens caerulescens (Hoffmann, 1803); 2) S. (S.) calatravensis Fuente, 1899; 3) S. (S.) chalcites (Illiger, 1807); 4) S. (S.) godet (Brullé, 1832); 5) S. (S.) maculatus (P. Rossi, 1792); 6) S. (S.) strigil Marseul, 1855; 7) S. (S.) submarginatus J. Sahlberg, 1913; and 8) S. (S.) tenuistrius sparsutus Solsky, 1876. The peak activity was recorded, key for the species is provided, and habitus images and male genitalia are illustrated in order to facilitate their taxonomic identifications. Saprinus species are diverse and common on animal carcass; they were likewise collected from a human cadaver in Lebanon. Preliminary comments on biology and distribution of the studied species are given. Our paper represents the first faunistic study on Histeridae from Lebanon. A rigorous research program regarding the biology of Saprinus in Lebanon and the neighbouring countries would greatly improve the knowledge of the diversity, activity, and possible forensic value of Saprinus.

Abstract Background The systematic positions of the extinct insect orders Hypoperlida, Miomoptera and Permopsocida were enigmatic and unstable for nearly a century. The recent studies based on new material, especially from the Cenomanian Burmese amber, shed light on evolutionary history of Acercaria resolving Permopsocida as the stem group of Condylognatha. However, the knowledge of the remaining two orders differs significantly. Results In this study, we describe new specimens and evaluate morphology of various structures with emphasis on the mouthparts and wing venation. Our results are primary based on revisions of the type specimens with a proper delimitation of taxa Hypoperlida and Miomoptera followed by their significance for the evolutionary history of Acercaria. Three new genera as Belmomantis gen. nov., Elmomantis gen. nov., and Mazonopsocus gen. nov. are designated as members of Palaeomanteidae. The Pennsylvanian Mazonopsocus provides a minimum age for calibration, in accordance to the presence of crown acercarians during the late Carboniferous. Conclusions This contribution demonstrates that Hypoperlida and Miomoptera are stem groups of Acercaria. The putative clade (Hypoperlida + Miomoptera) is appearing as potential sister group of (Psocodea + (Permopsocida + (Thripida + Hemiptera))).

Abstract When the same complex trait is exhibited by closely related species, a single evolutionary origin is frequently invoked. The complex stridulatory apparatus present in the forewings of extant crickets, mole crickets, katydids, and prophalangopsids, is currently interpreted as sharing a single common origin due to their similarity and unique function. An alternative hypothesis of convergent evolution in these ensiferan groups has challenged this common view, but remained controversial because of competing interpretations of wing venation. Here we propose another hypothesis for the widely and long debated homology of ensiferan stridulatory apparatus, performing the first 3D reconstruction of hidden structures at the wing bases. This approach allowed defining the homology of each vein from its very origin rather than after its more distal characteristics, which may be subjected to environmental pressure of selection. The stridulatory apparatus involves different veins in these four singing clades. In light of the most recent phylogenetic evidence, this apparatus developed four times in Ensifera, illustrating extraordinary convergent evolutions between closely related clades, by far exceeding the number of evolutionary steps ever proposed for calling ability in this group.

Abstract For a long time, New Caledonia was considered a continental island, a fragment of Gondwana harbouring old clades that originated by vicariance and so were thought to be locally ancient. Recent molecular phylogenetic studies dating diversification and geological data indicating important events of submergence during the Paleocene and Eocene (until 37 Ma) brought evidence to dismiss this old hypothesis. In spite of this, some authors still insist on the idea of a local permanence of a Gondwanan biota, justifying this assumption through a complex scenario of survival by hopping to and from nearby and now-vanished islands. Based on a comprehensive review of the literature, we found 40 studies dating regional clades of diverse organisms and we used them to test the hypothesis that New Caledonian and inclusive Pacific island clades are older than 37 Ma. The results of this meta-analysis provide strong evidence for refuting the hypothesis of a Gondwanan refuge with a biota that originated by vicariance. Only a few inclusive Pacific clades (6 out of 40) were older than the oldest existing island. We suggest that these clades could have extinct members either on vanished islands or nearby continents, emphasizing the role of dispersal and extinction in shaping the present-day biota.

Abstract The clade Triadophlebiomorpha represents a morphological ‘link’ between the Paleozoic griffenflies (Meganisoptera) and the modern taxa. Nevertheless they are relatively poorly known in the body structures and paleobiogeography. The Triassic dragonfly is extremely rare in China with only one previously recorded. A new family, Sinotriadophlebiidae Zheng, Nel et Zhang fam. nov., for the genus and species Sinotriadophlebia lini Zheng, Nel et Zhang gen. et sp. nov., is described from the Upper Triassic Baijiantan Formation of Xinjiang, northwestern China. It is the second Chinese Triassic odonatopteran and the second largest Mesozoic representative of this superorder in China. The discovery provides new information for the clade Triadophlebiomorpha during the Late Triassic and expands its distribution and diversity in Asia. The find reflects a close relationship between the two Triassic entomofaunas from Kyrgyzstan and the Junggar Basin, and provides a Carnian age constraint on the lowermost part of the Baijiantan Formation.

Many insects mimic plants in order to avoid detection by predators. Here, Garrouste and colleagues describe a katydid fossil that extends the record of leaf mimicry to the Middle Permian, more than 100 million years earlier than previously known fossil specimens of plant mimicry.

The Late Palaeozoic insect superorder Palaeodictyopterida exhibits a remarkable disparity of larval ecomorphotypes, enabling these animals to occupy diverse ecological niches. The widely accepted hypothesis presumed that their immature stages only occupied terrestrial habitats, although authors more than a century ago hypothesized they had specializations for amphibious or even aquatic life histories. Here, we show that different species had a disparity of semiaquatic or aquatic specializations in larvae and even the supposed retention of abdominal tracheal gills by some adults. While a majority of mature larvae in Palaeodictyoptera lack unambiguous lateral tracheal gills, some recently discovered early instars had terminal appendages with prominent lateral lamellae like in living damselflies, allowing support in locomotion along with respiratory function. These results demonstrate that some species of Palaeodictyopterida had aquatic or semiaquatic larvae during at least a brief period of their post-embryonic development. The retention of functional gills or gill sockets by adults indicates their amphibious lifestyle and habitats tightly connected with a water environment as is analogously known for some modern Ephemeroptera or Plecoptera. Our study refutes an entirely terrestrial lifestyle for all representatives of the early diverging pterygote group of Palaeodictyopterida, a greatly varied and diverse lineage which probably encompassed many different biologies and life histories.

The position of the Palaeozoic Anthracoptilidae has been a major problem of insect systematics for over a century. The previous hypotheses suggested affinities of Anthracoptilidae with the Palaeodictyopteroida, Protorthoptera, Hypoperlida, stem-Mantodea, Paraneoptera, Holometabola, or Eoblattida. Herein we put forward a new hypothesis based mainly on a comprehensive re-evaluation of the wing venation characters and re-examination of the type material of Anthracoptilus perrieri and Mesoptilus dolloi. The Anthracoptilidae are considered as belonging to the paoliid lineage, sister group of the Dictyoptera. In particular this result refutes the presence of Mantodea in the Paleozoic fossil record. The two families Strephocladidae and Strephoneuridae, are considered as junior synonyms of the Anthracoptilidae, while the previous synonymy of the Anthracoptilidae with the Ischnoneuridae is rejected. We consider the Permarrhaphidae, previously considered as synonym of the Anthracoptilidae, (and Permarrhaphus) as Insecta incertae sedis. The following new taxa are proposed: Mesoptilus carpenteri sp. nov. from the early Permian of Wellington Formation in Oklahoma (USA) which extends the range and distribution of the genus; Pseudomesoptilus gen. nov. is designated to include Mesoptilus sellardsi; Strephocladus permianus sp. nov. from the middle Permian of South of France; Westphaloptilus gallicus gen. et sp. nov. from the Bashkirian of the North of France.

We describe herein a new aphid genus and species Daoaphis magnalata gen. et sp. n., belonging to the extinct family Oviparosiphidae from the Middle Jurassic (ca. 165 Ma) Daohugou beds, Ningcheng County, Inner Mongolia, Northeast China. The new genus is considered to be the earliest certain representative of this family and the body remains of one of the oldest aphids. The evolutionary position of this new genus and distribution of the family are discussed.

Pondweed bugs (Hemiptera: Mesoveliidae), considered a sister group to all other Gerromorpha, are exceedingly rare as fossils. Therefore, each new discovery of a fossil mesoveliid is of high interest, giving new insight into their early evolutionary history and diversity and enabling the testing of their proposed relationships. Here, we report the discovery of new mesoveliid material from Spanish Lower Cretaceous (Albian) amber, which is the first such find in Spanish amber. To date, fossil records of this family only include one species from French Kimmeridgian as compression fossils, two species in French amber (Albian-Cenomanian boundary), and one in Dominican amber (Miocene). The discovery of two males and one female described and figured as Glaesivelia pulcherrima Sánchez-García & Solórzano Kraemer gen. et sp. n., and a single female described and figured as Iberovelia quisquilia Sánchez-García & Nel, gen. et sp. n., reveals novel combinations of traits related to some genera currently in the subfamily Mesoveliinae. Brief comments about challenges facing the study of fossil mesoveliids are provided, showing the necessity for a revision of the existing phylogenetic hypotheses. Some of the specimens were studied using infrared microscopy, a promising alternative to the systematic study of organisms preserved in amber that cannot be clearly visualised. The new taxa significantly expand the fossil record of the family and shed new light on its palaeoecology. The fossils indicate that Mesoveliidae were certainly diverse by the Cretaceous and that numerous tiny cryptic species living in humid terrestrial to marginal aquatic habitats remain to be discovered. Furthermore, the finding of several specimens as syninclusions suggests aggregative behaviour, thereby representing the earliest documented evidence of such ethology.

Sinoeuthemis daohugouensis gen. et sp. n. is the first record of the isophlebiopteran family Euthemistidae from Middle Jurassic of northeast China, while previously this family was restricted to the early Late Jurassic Kazakhstan. This new finding allows us to emend the family diagnosis with hindwing characters. This new species shows a mixture of characters alternatively present in different genera of the two families Euthemistidae and Sphenophlebiidae.

Paleoropronia salamonei gen. et sp. nov., the first Cenozoic Roproniidae, is described from the Paleocene of Menat (Massif Central, France) on the basis of its fore wing venation. The Roproniidae range between the Mesozoic and the present time. P. salamonei gen. et sp. nov. was perhaps a parasitoid on tenthredinid sawfly larvae, as these insects were present in the wasp fauna from Menat outcrop.

New material of the wasp family Maimetshidae (Apocrita) is presented from four Cretaceous amber deposits – the Neocomian of Lebanon, the Early Albian of Spain, the latest Albian/earliest Cenomanian of France, and the Campanian of Canada. The new record from Canadian Cretaceous amber extends the temporal and paleogeographical range of the family. New material from France is assignable to Guyotemaimetsha enigmatica Perrichot et al. including the first females for the species, while a series of males and females from Spain are described and figured as Iberomaimetsha Ortega-Blanco, Perrichot, and Engel gen. n., with the two new species Iberomaimetsha rasnitsyni Ortega-Blanco, Perrichot, and Engel sp. n. and I. nihtmara Ortega-Blanco, Delclòs, and Engel sp. n.; a single female from Lebanon is described and figured as Ahiromaimetsha najlae Perrichot, Azar, Nel, and Engel gen. et sp. n., and a single male from Canada is described and figured as Ahstemiam cellula McKellar and Engel gen. et sp. n. The taxa are compared with other maimetshids, a key to genera and species is given, and brief comments made on the family.

Three new palaeopteran insects are described from the Middle Permian (Guadalupian) of Salagou Formation in the Lodève Basin (South of France), viz. the diaphanopterodean Alexrasnitsyniidae fam. n., based on Alexrasnitsynia permiana gen. et sp. n., the Parelmoidae Permelmoa magnifica gen. et sp. n., and Lodevohymen lapeyriei gen. et sp. n. (in Megasecoptera or Diaphanopterodea, family undetermined). In addition the first record of mayflies attributed to family Syntonopteridae (Ephemeroptera) is reported. These new fossils clearly demonstrate that the present knowledge of the Permian insects remains very incomplete. They also confirm that the Lodève entomofauna was highly diverse providing links to other Permian localities and also rather unique, with several families still not recorded in other contemporaneous outcrops.

Libanopsyllipsocus alexanderasnitsyni gen. et sp. n., of Psyllipsocidae is described and figured from the Lower Cretaceous amber of Lebanon. The position of the new taxon is discussed and the fossil is compared to other psyllipsocids. The species represents the earliest record of the family Psyllipsocidae.

A commented list of fossil Odonata from the Oligocene outcrop of Rott is given, together with descriptions of new traces of oviposition in plant tissues, very similar to ichnotaxa already known from the early Eocene Laguna del Hunco floras of Patagonia. The joint presences of odonatan larvae and traces of oviposition demonstrate the autochthony of these insects in the palaeolake of Rott, confirming the existence of a diverse and abundant aquatic entomofauna, a situation strikingly different to that in the contemporaneous Oligocene palaeolake of Céreste (France).

Protomyrmeleon daohugouensis sp. n. and Protomyrmeleon lini sp. n., two new species of Protomyrmeleontidae from the Middle Jurassic of Jiulongshan Formation are the first Chinese representatives of this Mesozoic odonatopteran family. The type specimen of P. lini is exceptionally well preserved, showing several particular wing and body structures that were unknown, viz. unique shape of tarsal claws, extreme thoracic skewness, presence of three pairs of long spurs on all femora and tibiae. The type "A" (sensu Nel et al., 2005) of wing venation (i.e. with a very long bridge between IR2 and RP3/4) corresponds to that of the protomyrmeleontid fore wing. The nearly complete absence of the meso-metathoracic interpleural suture, newly discovered in the Protomyrmeleontidae, can be considered as a synapomorphy of the clade Protozygoptera + Odonata.

A new genus and a new species of Lygistorrhinidae, Parisognoriste eocenica is described from the Eocene Oise amber of the Paris Basin. Palaeognoriste sciariforme Meunier, 1904 and Palaeognoriste affine Meunier, 1912 are re-described. Lectotypes are designated for both species of Palaeognoriste. The phylogenetic positions of the new genus and Palaeognoriste Meunier are discussed.

Zygokaratawia reni, a new campterophlebiid genus and species is described from the Middle Jurassic of China. This fossil has a wing shape unique for this clade, i.e. a fore- and hind wing of the same width and very shortly petiolated, and hind wing cubito-anal area nearly as narrow as that of the forewing. This wing shape is convergently similar to that of recent Zygoptera: Calopterygidae, as well as to several other Cenozoic zygopteran clades, suggesting similar styles of flight and habits, i.e. predation on small insects and flight along trees of river banks.