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.

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.

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 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.

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.

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.

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.

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.

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.

Morphological reassessment of the movable calcar of delphacid planthoppers (Hemiptera: Fulgoromorpha: Delphacidae).

This study presents the morphology of calcar in adult Delphacidae based on representatives of the genera Ugyops Guérin-Meneville, 1834, Notuchus Fennah, 1969 (Ugyopini), Asiraca Latreille, 1798 (Asiracini), Kelisia Fieber, 1866, (Kelisini), Stenocranus Fieber, 1866 (Stenocranini), Chloriona Fieber, 1866, Megadelphax Wagner, 1963, Muellerianella Wagner, 1963, Javesella Fennah, 1963, Conomelus Fieber, 1866, Euconomelus Haupt, 1929, Hyledelphax Vilbaste, 1968, Stiroma Fieber, 1866, Struebingianella Wagner, 1963 and Xanthodelphax Wagner, 1963 (Delphacini). We used SEM electron microscopy, to define seven types of calcar structure (Types 1, 2, 5, 6, 7, 8, and 9) based on combinations of characters including shape, number of teeth and differentiation of sensory structures in species from fifteen genera. Additionally, two other types (Types 3 and 4) were determined based on the calcar descriptions from previous studies. Similarities and differences in calcar structure and function were discussed and emerging relationships between planthopper species and their particular habitats were indicated.

Phylogeny and biogeography of the enigmatic ghost lineage Cylindrotomidae (Diptera, Nematocera).

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.

The mid-Miocene Zhangpu biota reveals an outstandingly rich rainforest biome in East Asia.

During the Mid-Miocene Climatic Optimum [MMCO, ~14 to 17 million years (Ma) ago], global temperatures were similar to predicted temperatures for the coming century. Limited megathermal paleoclimatic and fossil data are known from this period, despite its potential as an analog for future climate conditions. Here, we report a rich middle Miocene rainforest biome, the Zhangpu biota (~14.7 Ma ago), based on material preserved in amber and associated sedimentary rocks from southeastern China. The record shows that the mid-Miocene rainforest reached at least 24.2°N and was more widespread than previously estimated. Our results not only highlight the role of tropical rainforests acting as evolutionary museums for biodiversity at the generic level but also suggest that the MMCO probably strongly shaped the East Asian biota via the northern expansion of the megathermal rainforest biome. The Zhangpu biota provides an ideal snapshot for biodiversity redistribution during global warming.

Nomenclatorial note on Bernaeinae (Hemiptera, Sternorrhyncha, Aleyrodidae).

The Bernaeinae, currently regarded as a subfamily of Aleyrodidae (Szwedo Drohojowska 2016, Drohojowska et al. 2019), is the only extinct subfamily of whiteflies with a fossil record from the Callovian (late Middle Jurassic) to Cenomanian (early Upper Cretaceous). Currently, it comprises seven species in six genera (Schlee 1970, Shcherbakov 2000, Drohojowska et al. 2019, Chen et al. 2020)-Bernaea neocomica Schlee, 1970; Burmoselis evelynae Shcherbakov, 2000; Heidea cretacica Schlee, 1970 (see comment below); Juleyrodes Shcherbakov, 2000 (J. gilli Shcherbakov, 2000, J. visnyai Shcherbakov, 2000), Paraburmoselis kachinensis Chen, Zhang, Wang et Zheng, 2020 and Sinicoselis weberi Drohojowska, Wegierek, Evans et Huang, 2019. Heidea by mistake was figured in Drohojowska Szwedo (2011a, p. 192, Fig. 23) in Aleyrodinae, but no taxonomic decisions were taken regarding this fossil. Later, in the checklist of fossil Aleyrodidae, Szwedo Drohojowska (2016: supplement p. 6), listed Heidea in Bernaeinae. This statement was not noted by Chen et al. (2020) listing again Heidea in Aleyrodinae referring to erroneous placement on the figure in Drohojowska Szwedo (2011a).

Wing Interference Patterns in patterned wings of Culicoides Latreille, 1809 (Diptera: Ceratopogonidae)-exploring potential identification tool.

Biting midges, nematoceran flies family Ceratopogonidae, is represented by more than 6200 valid species. This group is still poorly known and not easy in identification. Structural coloration of insect wings (WIP, or Wing Interference Pattern) is an example of an optical phenomenon called iridescence, which is based on the formation of pigment-free, rainbow colours caused by the interference of white light reflected from transparent or semi-transparent structures. Preliminary results indicate that the structural colouring of the wings of these insects may be helpful in the delimitation of closely related and cryptic species.

New fossil from mid-Cretaceous Burmese amber confirms monophyly of Liadopsyllidae (Hemiptera: Psylloidea).

Amecephala pusilla gen. et sp. nov. is described and illustrated on the basis of a well-preserved female psyllid (Liadopsyllidae) in a piece of Cretaceous Myanmar amber. The new genus differs from other members of Liadopsyllidae in details of the antennae and forewings. For the first time, the presence of a circumanal ring is documented for Mesozoic psyllids. Based on differences in the length of female terminalia, it is suggested that Liadopsyllidae may have displayed a diversified oviposition biology. As far as known, Liadopsyllidae lack a pulvillus, a putative autapomorphy supporting the monophyly of Liadopsyllidae. An identification key to genera and an annotated checklist of known Liadopsyllidae species are provided. New synonyms and combinations are proposed and the status of the subfamily Miralinae is discussed.

Fossils reshape the Sternorrhyncha evolutionary tree (Insecta, Hemiptera).

The Sternorrhyncha, which comprise about 18,700 described recent species, is a suborder of the Hemiptera, one of big five most diverse insect orders. In the modern fauna, these tiny phytophages comprise insects of great ecological and economic importance, like aphids (Aphidomorpha), scale insects (Coccidomorpha), whiteflies (Aleyrodomorpha) and psyllids (Psylloidea). Their evolutionary history can be traced back to the Late Carboniferous, but the early stages of their evolution and diversification is poorly understood, with two known extinct groups-Pincombeomorpha and Naibiomorpha variously placed in classifications and relationships hypotheses. Most of the recent Sternorrhyncha groups radiated rapidly during the Cretaceous. Here we report the new finding of very specialised sternorrhynchans found as inclusions in mid-Cretaceous amber from Kachin state (northern Myanmar), which represent another extinct lineage within this hemipteran suborder. These fossils, proposed to be placed in a new infraorder, are revealed to be related to whiteflies and psyllids. We present, also for the first time, the results of phylogenetic analyses covering extinct and extant lineages of the Sternorrhyncha.