The earliest known holometabolous insects.

The Eumetabola (Endopterygota (also known as Holometabola) plus Paraneoptera) have the highest number of species of any clade, and greatly contribute to animal species biodiversity. The palaeoecological circumstances that favoured their emergence and success remain an intriguing question. Recent molecular phylogenetic analyses have suggested a wide range of dates for the initial appearance of the Holometabola, from the Middle Devonian epoch (391 million years (Myr) ago) to the Late Pennsylvanian epoch (311 Myr ago), and Hemiptera (310 Myr ago). Palaeoenvironments greatly changed over these periods, with global cooling and increasing complexity of green forests. The Pennsylvanian-period crown-eumetabolan fossil record remains notably incomplete, particularly as several fossils have been erroneously considered to be stem Holometabola (Supplementary Information); the earliest definitive beetles are from the start of the Permian period. The emergence of the hymenopterids, sister group to other Holometabola, is dated between 350 and 309 Myr ago, incongruent with their current earliest record (Middle Triassic epoch). Here we describe five fossils--a Gzhelian-age stem coleopterid, a holometabolous larva of uncertain ordinal affinity, a stem hymenopterid, and early Hemiptera and Psocodea, all from the Moscovian age--and reveal a notable penecontemporaneous breadth of early eumetabolan insects. These discoveries are more congruent with current hypotheses of clade divergence. Eumetabola experienced episodes of diversification during the Bashkirian-Moscovian and the Kasimovian-Gzhelian ages. This cladogenetic activity is perhaps related to notable episodes of drying resulting from glaciations, leading to the eventual demise in Euramerica of coal-swamp ecosystems, evidenced by floral turnover during this interval. These ancient species were of very small size, living in the shadow of Palaeozoic-era 'giant' insects. Although these discoveries reveal unexpected Pennsylvanian eumetabolan diversity, the lineage radiated more successfully only after the mass extinctions at the end of the Permian period, giving rise to the familiar crown groups of their respective clades.

A swarm of whiteflies--the first record of gregarious behavior from Eocene Baltic amber.

A new whitefly Snotra christelae gen. et sp. n. is characterized, illustrated, and described from the Baltic amber. It represents the first record of gregarious behavior of Aleyrodinae (Aleyrodidae) whiteflies in fossil state. Implications of this finding on interpretation of whiteflies and their host-plant relationships and evolutionary traits of the group are discussed.

Tip of the clade on the top of the World--the first fossil Lophopidae (Hemiptera: Fulgoromorpha) from the Palaeocene of Tibet.

Lophopidae is a family of planthoppers (Hemiptera: Fulgoromorpha) present today in tropical and subtropical zones of the Old World. The most recent taxonomic studies and phylogeny of these insects do not include the extinct representatives. Therefore, each new discovery of a fossil lophopid is of high interest, giving new insights to their evolutionary history and enabling to test the proposed relationships. The recent findings of extinct Lophopidae in Europe, in various Palaeogene deposits, put in doubts their proposed evolutionary and biogeographic scenario. The new fossil from the Palaeocene of Northern Tibet is related to one of the Lophopidae clades, Apia(+) group, believed to be the most advanced one, and recently distributed in the recent Sundaland-New Guinea-Queensland area. A new genus and species Gesaris gnapo gen. et sp. n. provide information on early lophopids diversity and relationships and demonstrates the necessity for a revision of the existing hypotheses for the initial diversification and distributional pattern of the Lophopidae.

The discovery of a fossil whitefly from Lower Lusatia (Germany) presents a challenge to current ideas about Baltic amber.

The whiteflies (Hemiptera: Aleyrodidae) are small sternorrhynchan insects, which have the potential to cause significant economic damage to agricultural crops. There is a paucity of knowledge regarding the diversity, disparity, and evolutionary history of these insects, with classification based on the immatures, called puparia. The fossil record of whiteflies is sparse and incomplete, with the majority of fossils representing imaginal forms preserved as inclusions in fossilized resins. In this study, we present the first inclusion in succinite associated with the layers of Lower/Middle Miocene 2nd Lusatian Lignite Seam of Wanninchen in Brandenburg, Germany. The objective of the present study is to elaborate this fossil, and as a consequence, a new fossil genus and species, Pudrica christianottoi  gen. et sp. nov., is described. This fossil is a representative of the subfamily Aleyrodinae, and it is the third fossil genus of this whitefly subfamily to be described. The discovery of the fossil inclusion in the succinite from the lignite deposits of Lower Lusatia challenges the current understanding of the character and conditions of formation and deposition of central and east European Paleogene fossil resins. Succinite is a fossil resin that occurs in the Eocene deposits of the Gulf of Gdansk, belonging to the Prussian Formation, containing a glauconite-rich horizon known as the 'Blue Earth'. Similarly, glauconite-rich deposits are present in the Lublin area of Poland, where they are associated with the occurrence of succinite. Additionally, succinite has been found in deposits in the Rovno-Zhitomir area of Ukraine, which are alluvial deposits containing glauconite and lignite layers. Succinite was also identified in Eocene strata of Spitsbergen and in Axel Heiberg Island in the Canadian Arctic. Succinite has also been discovered in early Miocene deposits in Bitterfeld, Germany, where it occurs alongside lignite deposits (the deposit actually encompasses different fossil resins, so potentially originating from different source plants). Furthermore, it has been identified in younger (Pleistocene) deposits across Europe. The autochthonous (parautochthonous) character of the lignite deposits in Lower Lusatia raises questions regarding the time range of the succinite-producing gymnosperm trees and the autochthonous or allochthonous character of the lignite layers associated fossilized resins.

Chemical and palaeoentomological evidence of a relationship between early Eocene Belgian and Oise (France) ambers.

Of the early Eocene amber deposits known across the world, Belgian amber has been mostly absent from the relevant literature. We reinvestigated amber held in the palaeobotanical collection of the Royal Belgian Institute of Natural Sciences, Brussels, which derived from three localities in Belgium that originated from two geographical areas (Leval-Trahegnies and Orp-le-Grand). Using Fourier transform infrared (FTIR) spectroscopy we show the close chemical relationship of Belgian amber to the early Eocene Oise amber from the Paris Basin, and highlight the potential effect of weathering on the amber chemistry. The amber derives from a very similar botanical source as the Oise amber (Combretaceae or Leguminosae-Caesalpinioideae), but from different coeval basins. The two Leval-Trahegnies localities provided amber that exhibit different stages of weathering (heavily fissured and crazed, darkened) and lacking any inclusions. The Orp-le-Grand locality provided the least weathered amber, with one amber piece containing two inclusions: a mite and a new genus and species of hemipteran (Cativolcus uebruum gen. et sp. nov.), and a second one that preserved the impression of insect wings pressed into the surface.

Mesozoic evolution of cicadas and their origins of vocalization and root feeding.

Extant cicada (Hemiptera: Cicadoidea) includes widely distributed Cicadidae and relictual Tettigarctidae, with fossils ascribed to these two groups based on several distinct, minimally varying morphological differences that define their extant counterparts. However, directly assigning Mesozoic fossils to modern taxa may overlook the role of unique and transitional features provided by fossils in tracking their early evolutionary paths. Here, based on adult and nymphal fossils from mid-Cretaceous Kachin amber of Myanmar, we explore the phylogenetic relationships and morphological disparities of fossil and extant cicadoids. Our results suggest that Cicadidae and Tettigarctidae might have diverged at or by the Middle Jurassic, with morphological evolution possibly shaped by host plant changes. The discovery of tymbal structures and anatomical analysis of adult fossils indicate that mid-Cretaceous cicadas were silent as modern Tettigarctidae or could have produced faint tymbal-related sounds. The discovery of final-instar nymphal and exuviae cicadoid fossils with fossorial forelegs and piercing-sucking mouthparts indicates that they had most likely adopted a subterranean lifestyle by the mid-Cretaceous, occupying the ecological niche of underground feeding on root. Our study traces the morphological, behavioral, and ecological evolution of Cicadoidea from the Mesozoic, emphasizing their adaptive traits and interactions with their living environments.

An extraordinary tribe of Tropiduchidae from the Eocene Baltic amber (Hemiptera: Fulgoromorpha: Fulgoroidea).

The new tribe Patollini trib. n. of the Tropiduchidae with the extinct genus Patollo gen. n., comprising two species Patollo natangorum sp. n. and P. aestiorum sp. n. from Eocene Baltic amber is described. Taxonomic placement of some fossil taxa ascribed to Tropiduchidae is discussed. The classification of Tropiduchidae is discussed, as well as phylogenetic position and fossil record of Tropiduchidae and related taxa.

The first Aleyrodidae from the Lowermost Eocene Oise amber (Hemiptera: Sternorrhyncha).

The first records are provided of the family Aleyrodidae in the Lowermost Eocene amber of Oise, France. The following new taxa in the subfamily Aleurodicinae are described, figured and discussed, together with an identification key: Oisedicus maginus gen. et sp. n., Clodionusfizoli gen. et sp. n., Lukotekia menae gen. et sp. n. and Isaraselis cladiva gen. et sp. n. Unplaced species of Lukotekia are briefly described, and the diversity of the whiteflies from Oise amber is discussed. The importance of fossils for palaeoecological and palaeoclimatological reconstruction is briefly considered.

Toward a new classification of planthoppers Hemiptera Fulgoromorpha: 2. Higher taxa, their names and their composition.

A synthesis of planthopper suprafamilial and familial classification is provided and discussed, solving some nomenclatural issues for the group, and adopting an enriched presentation with a clear distinction between the taxon name and its concept.

New Surijokocixiidae (Insecta: Hemiptera: Fulgoromorpha) from the Middle Triassic of China.

A new Middle Triassic (Ladinian) planthopper of the family Surijokocixiidae (Surijokocixioidea, Fulgoromorpha) from the Tongchuan Formation in Shaanxi, NW China is established as Sinosurijikocixius tongchuanensis gen. et sp. nov. All the known taxa assigned to Surijokocixiidae are reviewed and compared. The palaeogeographic distributional pattern of Surijokocixiidae from the middle Permian to Late Triassic is summarized.

A new species of Eocene whiteflySnotra herczeki sp. nov. (Hemiptera, Sternorrhyncha, Aleyrodidae) from Baltic amber.

The second species of the Aleyrodinae whitefly genus Snotra Szwedo et Drohojowska, 2016 is described based on an inclusion in the Eocene Baltic amber. It is second species of the genus, adding new data to knowledge on disparity and taxonomic diversity of whiteflies in the Eocene Baltic amber. The diversity of the group and its fossil record in the Eocene fossil resins is briefly discussed.

A second Aleurodicinae from the Eocene Baltic amberMedocellodes blackmani gen. et sp. nov. (Hemiptera, Sternorrhyncha, Aleyrodidae).

A new genus and species of Aleurodicinae whiteflies from the Eocene Baltic amber is described. Medocellodes blackmani Drohojowska et Szwedo gen. et sp. nov. is the second representative of Aleurodicinae from Baltic amber, presenting mixture of plesiomorphic characters, as retention of median ocellus, with apomorphic states, shared also with other contemporaneous Aleurodicinae from Rovno amber. Morphological characters of the newly described fossil are briefly discussed.

Evolutionary implications of new Postopsyllidiidae from mid-Cretaceous amber from Myanmar and sternorrhynchan nymphal conservatism.

Nymphs of extinct sternorrhynchan hemipterans are extremely rare, although very important for understanding of evolutionary traits of these insects. A protopsyllidioid nymph, in mid-Cretaceous amber from Kachin, Myanmar, placed in the family Postopsyllidiidae, is the first nymph of this family to be found in the fossil. Postopsyllidiidae previously comprised the sole genus Postopsyllidium with a few species: P. rebeccae, P. grimaldii and P. burmaticum from Kachin amber (Cenomanian) and P. emilyae from Turonian amber of New Jersey. Here, we report a new genus and species of postopsyllid Megalophthallidion burmapteron gen. et sp. nov. (imago) and the first known nymph of the family ascribed to the same genus. An overview of the fossil record of sternorrhynchan nymphs, and the importance of this finding, unlocking a new and complementary window to understanding the evolutionary traits of Protopsyllidioidea and other Sternorrhyncha hemipterans are presented.

Spotted beautyGedanochila museisucini gen. et sp. nov.a new Achilidae from Eocene Baltic amber (Hemiptera: Fulgoromorpha) and its relation to Achilini.

Gedanochila museisucini gen. et sp. nov. is described, based on inclusions in the Eocene Baltic amber. A morphological phylogenetic analysis supports the placement of Gedanochila gen. nov. into the tribe Achilini. Definition, content and subdivisions of the tribe as well as position of extinct taxa placed within are briefly discussed.

New finding of Toxorhina (Ceratocheilus) limoniid fly in Eocene Baltic amber and the biogeographical context of the genus.

Based on new fossil materials, a new species Toxorhina (Ceratocheilus) christelius sp. nov. has been described herein with complete documentation of drawings and photographs. Features such as wide spine on the gonocoxite differentiating the new species of Toxorhina were discussed. Finding new interesting fossil materials also allowed for providing an emended diagnosis and additional description of known Eocene species-Toxorhina (Ceratocheilus) eridanus. Comparison of chosen morphological features of fossil and recent representatives of the genus were given and key for fossil species of subgenus Ceratocheilus was introduced. Distribution of recent Toxorhina and evolutionary history of the genus were discussed. The results of research on fossil materials prove that the stratigraphic range of the subgenus Ceratocheilus and the genus Toxorhina goes back to the Eocene, there is no evidence of their existence on Earth before. Moreover, these insects were probably associated with a warm climate, they were found for example in Baltic amber, the deposits of which were formed mainly in a subtropical climate. The reach diversity of the genus Toxorhina of recent fauna is strictly observed in tropical zones around the world.

A new giant dung midge from Miocene Ethiopian amber (Diptera: Scatopsidae).

Fossil new genus and species Afroscatopse haennii gen. et sp. nov. from Miocene Ethiopian amber are described and illustrated and placed in the tribe Colobostematini. The gigantic adults of this scavenger fly, largest within the family, with dull body, long antennae and broad wings evidently were adapted to live on bark of trees. It is suggested that it was a myrmecophilous species with larvae living in the nests of tree ants.

Genomic-Phenomic Reciprocal Illumination: Desyopone hereon gen. et sp. nov., an Exceptional Aneuretine-like Fossil Ant from Ethiopian Amber (Hymenoptera: Formicidae: Ponerinae).

Fossils are critical for understanding the evolutionary diversification, turnover, and morphological disparification of extant lineages. While fossils cannot be sequenced, phenome-scale data may be generated using micro-computed tomography (µ-CT), thus revealing hidden structures and internal anatomy, when preserved. Here, we adduce the male caste of a new fossil ant species from Miocene Ethiopian amber that resembles members of the Aneuretinae, matching the operational definition of the subfamily. Through the use of synchrotron radiation for µ-CT, we critically test the aneuretine-identity hypothesis. Our results indicate that the new fossils do not belong to the Aneuretinae, but rather the Ponerini (Ponerinae). Informed by recent phylogenomic studies, we were able to place the fossils close to the extant genus Cryptopone based on logical character analysis, with the two uniquely sharing absence of the subpetiolar process among all ponerine genera. Consequently, we: (1) revise the male-based key to the global ant subfamilies; (2) revise the definitions of Aneuretinae, Ponerinae, Platythyreini, and Ponerini; (3) discuss the evolution of ant mandibles; and (4) describe the fossils as †Desyopone hereon gen. et sp. nov. Our study highlights the value of males for ant systematics and the tremendous potential of phenomic imaging technologies for the study of ant evolution.

A New Genus of Spittlebugs (Hemiptera, Cercopidae) from the Eocene of Central Tibetan Plateau.

The superfamily Cercopoidea is commonly named as "spittlebugs", as its nymphs produce a spittle mass to protect themselves. Cosmoscartini (Cercopoidea: Cercopidae) is a large and brightly colored Old World tropical tribe, including 11 genera. A new genus Nangamostethos gen. nov. (type species: Nangamostethostibetense sp. nov.) of Cosmoscartini is described from Niubao Formation, the late Eocene of central Tibetan Plateau (TP), China. Its placement is ensured by comparison with all the extant genera of the tribe Cosmoscartini. The new fossil represents one of few fossil Cercopidae species described from Asia. It is likely that Nangamostethos was extinct from the TP due to the regional aridification and an overturn of plant taxa in the late Paleogene.

Widespread mineralization of soft-bodied insects in Cretaceous amber.

Fossilized tree resin, or amber, commonly contains fossils of animals, plants and microorganisms. These inclusions have generally been interpreted as hollow moulds or mummified remains coated or filled with carbonaceous material. Here, we provide the first report of calcified and silicified insects in amber from the mid-Cretaceous Kachin (Burmese) amber. Data from light microscopy, scanning electron microscopy (SEM), energy-dispersive and wavelength-dispersive X-ray spectroscopy (EDX and WDX), X-ray micro-computed tomography (Micro-CT) and Raman spectroscopy show that these Kachin fossils owe their preservation to multiple diagenetic mineralization processes. The labile tissues (e.g. eyes, wings and trachea) mainly consist of calcite, chalcedony and quartz with minor amounts of carbonaceous material, pyrite, iron oxide and phyllosilicate minerals. Calcite, quartz and chalcedony also occur in cracks as void-filling cements, indicating that the minerals formed from chemical species that entered the fossil inclusions through cracks in the resin. The results demonstrate that resin and amber are not always closed systems. Fluids (e.g. sediment pore water, diagenetic fluid and ground water) at different burial stages have chances to interact with amber throughout its geological history and affect the preservational quality and morphological fidelity of its fossil inclusions.

Bitara gen. nov. of Tropiduchidae (Hemiptera: Fulgoromorpha) east of Wallace line.

A new genus and species of the tropiduchid planthopper of the tribe Tropiduchini is described as Bitara augusta gen. et sp. nov. from Madang Province, Papua New Guinea. It is the 18th species of Tropiduchini and the 9th genus known east of the Wallace line. It is another taxon with characteristic asymmetric genitalia within this tribe.