The structure of wing in the earliest Permopsocida.

Permopsocids are small acercarian insects with mouthparts specialized for sucking. They are closely related to Hemiptera and Thysanoptera. The earliest known representatives are from the Early Permian. Here evidence is presented that the Permopsocida occurred even earlier in Pennsylvanian (Moscovian) deposits in the Piesberg quarry near Osnabrück (Lower Saxony, Germany). This material is assigned to the Permian family Psocidiidae; Carbonopsocus mercuryi gen. et sp. nov., based on the wing venation diagnosed by the unique branching pattern of the main veins, the shape of the areola postica being longer than wide, the angular shape of the pterostigma, the ir crossvein directed proximally mid of pterostigma (apomorphy) and the vannus formed by the three veins of PCu, A1 and A2. The shape of the veins, with a Y-vein formed by the distal fusion of PCu with A1, could be a putative symplesiomorphy of the Psocodea with Permopsocida and Hemiptera. C. mercuryi gen. et sp. nov. is the first appearance date for Permopsocida and roots the Acercaria tree. In addition, another specimen of Dichentomum cf. arroyo (Psocidiidae) from Carrizo Arroyo is presented and figured, confirming the presence of the genus Dichentomum near the Carboniferous-Permian boundary and linking it to the Artinskian species from Elmo in Kansas, USA.

A new mayfly subfamily sheds light on the early evolution and Pangean origin of Baetiscidae (Insecta: Ephemeroptera).

The family Baetiscidae Edmunds & Traver, 1954 is recognisable among mayflies due to its bizarre larvae, equipped with a robust and spiked thoracic notal shield covering part of the abdomen up to sixth segment. Originally being described as extant species from the USA and Canada, Baetiscidae were later found in the fossil record as well, specifically in Cretaceous of Brazil and Eocene Baltic amber. An enigmatic piece of fossil evidence are two larvae from the Early Cretaceous Koonwarra Fossil Bed in Australia, which have been presumed as attributable to Baetiscidae and briefly discussed in previous studies. In the present contribution, we reinvestigate these fossils and confirm their attribution to the family Baetiscidae. These larvae are depicted and described as Koonwarrabaetisca jelli gen. et sp. nov. and Koonwarrabaetisca duncani sp. nov. For both Cretaceous genera Protobaetisca Staniczek, 2007 and Koonwarrabaetisca gen. nov. we establish a new subfamily Protobaetiscinae subfam. nov. within the family Baetiscidae, based on the presence of markedly shortened thoracic sterna. The phylogenetic position of newly described subfamily is clarified using a cladistic analysis; Protobaetiscinae subfam. nov. forms a monophyletic clade, sister to Baetiscinae. The confirmation of the distribution of Baetiscidae in the Cretaceous of Australia suggests almost worldwide distribution of this family in the deep time. Given their limited dispersal abilities, this distributional pattern can be best explained by the Pangean origin for this family, moving the time of their origin at least to the Early Jurassic. The larvae of Koonwarrabaetisca gen. nov. exhibit the same ecomorphological specialization as the rest of Baetiscidae, that supporting with a high probability their lifestyle similar to extant Baetisca Walsh, 1862. The larvae probably lived in the flowing water with stony substrate densely covered by filamentous algae, and in the places of accumulation of dead plant and algae matter during the last instars. Thus, Koonwarrabaetisca gen. nov. could be the allochthonous component in mayfly fauna of the Koonwarra paleolake.

Thoracic and abdominal outgrowths in early pterygotes: a clue to the common ancestor of winged insects?

One of the fundamental questions in insect evolution is the origin of their wings and primary function of ancestral wing precursors. Recent phylogenomic and comparative morphological studies broadly support a terrestrial ancestor of pterygotes, but an aquatic or semiaquatic ancestor cannot be ruled out. Here new features of the branchial system of palaeodictyopteran larvae of several different instars of Katosaxoniapteron brauneri gen. et sp. nov. (Eugereonoidea) from the late Carboniferous collected at Piesberg (Germany) are described, which consist of delicate dorsolateral and lamellate caudal abdominal gills that support an aquatic or at least semiaquatic lifestyle for these insects. Moreover, the similar form and surface microstructures on the lateral abdominal outgrowths and thoracic wing pads indicate that paired serial outgrowths on segments of both tagmata presumably functioned as ancestral type of gills resembling a protopterygote model. This is consistent with the hypothesis that the wing sheaths of later stage damselfly larvae in hypoxic conditions have a respiratory role similar to abdominal tracheal gills. Hence, the primary function and driving force for the evolution of the precursors of wing pads and their abdominal homologues could be respiration.

Fluctuation in the diversity of mayflies (Insecta, Ephemerida) as documented in the fossil record.

Due to their aquatic larvae, the evolution of mayflies is intricately tied to environmental changes affecting lakes and rivers. Despite a rich fossil record, little is known about the factors shaping the pattern of diversification of mayflies in deep time. We assemble an unprecedented dataset encompassing all fossil occurrences of mayflies and perform a Bayesian analysis to identify periods of increased origination or extinction. We provide strong evidence for a major extinction of mayflies in the mid-Cretaceous. This extinction and subsequent faunal turnover were probably connected with the rise of angiosperms. Their dominance caused increased nutrient input and changed the chemistry of the freshwater environments, a trend detrimental mainly to lacustrine insects. Mayflies underwent a habitat shift from hypotrophic lakes to running waters, where most of their diversity has been concentrated from the Late Cretaceous to the present.

Muddying the unexplored post-industrial waters: Biodiversity and conservation potential of freshwater habitats in fly ash sedimentation lagoons.

Deposits of fly ash and other coal combustion wastes are common remnants of the energy industry. Despite their environmental risks from heavy metals and trace elements, they have been revealed as refuges for threatened terrestrial biodiversity. Surprisingly, freshwater biodiversity of fly ash sedimentation lagoons remains unknown despite such lack of knowledge strongly limits the efficient restoration of fly ash deposits. We bring the first comprehensive survey of freshwater biodiversity, including nekton, benthos, zooplankton, phytoplankton, and macrophytes, in fly ash lagoons across industrial regions of the Czech Republic. To assess their conservation potential, we compared their biodiversity with abandoned post-mining ponds, the known strongholds of endangered aquatic species in the region with a shortage of natural ponds. Of 28 recorded threatened species, 15 occurred in the studied fly ash lagoons, some of which were less abundant or even absent in the post-mining ponds. These are often species of nutrient-poor, fishless waters with rich vegetation, although some are specialised extremophiles. Species richness and conservation value of most groups in the fly ash lagoons did not significantly differ from the post-mining ponds, except for species richness of benthos, zooplankton, and macrophytes, which were slightly lower in the fly ash lagoons. Although the concentrations of some heavy metals (mainly Se, V, and As) were significantly higher in the fly ash lagoons, they did not significantly affect species richness or conservation value of the local communities. The differences in species composition therefore does not seem to be caused by water chemistry. Altogether, we have shown that fly ash lagoons are refuges for threatened aquatic species, and we thus suggest maintaining water bodies during site restoration after the cessation of fly ash deposition. Based on our analyses of environmental variables, we discuss suitable restoration practices that efficiently combine biodiversity protection and environmental risk reduction.

Big diversity in a small hotspot: two new species of Leptophlebiidae (Insecta, Ephemeroptera) from New Caledonia.

Two new species from Grande Terre Island, New Caledonia, namely Fasciamiruspetersorumsp. nov. and Simulacalararasp. nov. are described based on larval morphology and molecular data (COI sequences). Fasciamiruspetersorumsp. nov. is distributed in the southern part of the island and is characterised by a reduced third segment of the labial palps and all abdominal gills divided from the base. The species inhabits slow-flowing aquatic habitats with fine-grained substrate in forest brooks. Simulacalararasp. nov. is known from a single locality in the northern part of the island and is characterised by narrow and distinctly elongated abdominal gills 1-7. It was collected from fine substrates behind stones in riffles with slightly turbulent flow. Both species were recorded only in areas with ultramafic bedrock.

Mayfly Ephemeraglaucops (Ephemeroptera, Ephemeridae) recorded in the Czech Republic after almost a century.

Background: The mayfly Ephemera (Sinephemera) glaucops Pictet, 1843 has been considered regionally extinct in the Czech Republic, with the last occurrence dating from 1933. Its extinction was connected with the anthropogenic changes of lowland rivers, forming the original habitat of E.glaucops within the area of the Czech Republic. However, the species has been reported as spreading in man-made, often post-industrial waterbodies in Germany, The Netherlands and Austria since the 1970s. New information: We report E.glaucops from the Czech Republic, based on 27 larvae collected in the North Bohemia lignite basin in 2018. All individuals were found at one locality - a small kaolin pit in the shallow part near the shore, mostly without macrophytes. We provide details about the locality and morphological diagnostic characters of E.glaucops. This study highlights the importance of post-industrial sites for aquatic biodiversity as surrogate biotopes for degraded original habitats.

Two new species of Alainites (Ephemeroptera, Baetidae) from the Mediterranean biodiversity hotspot.

The mayfly genus Alainites Waltz & McCafferty, 1994 encompassed 20 species and was represented across the West Palaearctic region by six species. Based on morphological (nymphal characters) and molecular (mitochondrial COI sequences) evidence, two new species are described: A.bengunn sp. nov. from Sardinia and A.gasithi sp. nov. from Israel. Both species are confined to narrow distribution ranges, in line with most of their congeners from the region. The key nymphal traits are discussed and identified to distinguish species in the group.

Contribution to the knowledge of the genus Takobia Novikova & Kluge, 1987 (Ephemeroptera, Baetidae) in Central Asia.

Based on the original type material, the nymphal stage of the mayfly Takobiamaxillare is redescribed; in parallel, a lectotype is designated. Takobiamaxillare is the type species of the genus Takobia, and an accurate and complete knowledge of its morphology is crucial to the delimitation of this problematic genus and clarification of its phylogenetic affinities. Ambiguous characters, previously reported for this species in the literature are clarified. Furthermore, two new species in the same genus are described, namely Takobiasinusopalpata sp. nov. and Takobiashughnonica sp. nov. based on the morphology of nymphs from Central Asia, supplemented with COI sequences. Implications for the systematics of Takobia and related taxa are discussed and the need for an extensive phylogenetic study of this group is stressed.

A new species of Epeorus (Caucasiron) (Ephemeroptera, Heptageniidae) from Azerbaijan and Iran.

A new species, Epeorus (Caucasiron) hyrcanicussp. nov., is described based on larval morphology and molecular data (COI) containing sequences from all Caucasian Caucasiron species described to date. The species is distributed in the Hyrcanian forest of southeastern Azerbaijan and northwestern Iran. Based on our wide-range sampling, the new species is likely endemic to this area. The most pronounced larval morphological diagnostic characters are the coloration pattern of abdominal sterna (a pair of oblique stripes and stripe-like medio-lateral maculae) and terga (triangular medial maculae), poorly developed projection of the costal margin of gill plates III, presence of hair-like setae on the surface of abdominal terga, and relatively wide shape of gill plates VII (in natural position from ventral view). The diagnostic characters are compared to related species, and primary information to habitat is provided.

Life history, systematics and flight ability of the Early Permian stem-mayflies in the genus Misthodotes Sellards, 1909 (Insecta, Ephemerida, Permoplectoptera).

BACKGROUND: The stem-group of Ephemeroptera is phylogenetically important for understanding key steps in evolutionary history of early pterygote insects. However, these taxa have been mostly studied from the taxonomy point of view focused on the pattern of wing venation and often using only classical optical microscopy devices. In-depth studies on detailed morphology of the different body structures are scarcely performed, although the results are critical for elucidation of life history traits and their evolutionary pattern among the basal pterygotes. RESULTS: New information is presented on the morphology of two species of Misthodotes, which are stem-mayflies from the Early Permian. Based on new results obtained from a re-examination of the type specimens and supplementary material, we infer the life history traits of both the adult and larval stages of these Palaeozoic insects and reconsider previous interpretations. For the first time, we report the structure of the thoracic pleura and the articulation at the base of the wing in a stem-group of Ephemeroptera and compare them with those of extant mayflies. We also provide additional support for the systematic placement of investigated taxa and an amended diagnosis of the genus Misthodotes. CONCLUSIONS: Adult Misthodotes sharovi and Misthodotes zalesskyi had chewing mouthparts, which enabled them to scavenge or feed on plants. The wing apparatus was adapted for slow powered flapping flight and gliding, using long caudal filaments for steering. The wing base does not have rows of articulary sclerites as previously hypothesized for some Palaeozoic taxa but inflexible axilla similar to that found in modern mayflies. The structure of the thoracic pleura is also similar to that in the crown group of Ephemeroptera, while differences in the course of sutures may be explained by an evolutionary trend towards more powerful dorsoventral flying musculature and forewing-based flight (anteromotorism) in modern taxa. There is no evidence for swarming behaviour and mating in the air as occurs in modern mayflies as they had none of the associated morphological adaptations. Putative larvae of Misthodotes can not be unambiguously associated with the adults. They also exhibit some morphological specializations of Protereismatidae like 9 pairs of abdominal tracheal gills supporting their benthic lifestyle with legs adapted to burrowing.

Updated check-list of the mayflies (Insecta: Ephemeroptera) of Iraq.

Based on a recent field survey in Iraqi Kurdistan and a critical evaluation of previously published data, 37 mayfly species are listed as occurring in Iraq. We collected and identified nine species as new for the country and corrected some previously published records. For several species scarcely treated in the literature, we provide information allowing their identification in the larval stage to promote the acquisition of reliable faunistic data from Iraq in the future.

Identification guide to larvae of Caucasian Epeorus (Caucasiron) (Ephemeroptera, Heptageniidae).

The Caucasus and adjacent areas are inhabited by fifteen species of mayflies of the genus Epeorus, subgenus Caucasiron Kluge, 1997 (Heptageniidae). This identification guide aims to facilitate an accurate species identification of their larvae and sum up all available information on their taxonomy and distribution. An identification key is provided, and the important diagnostic characters of all species are described and illustrated. The larva of E. (C.) insularis (Braasch, 1983) is described for the first time. This study enables the routine identification of Caucasiron larvae necessary for biomonitoring and hydrobiological research in the Caucasus region.

Diversity and distribution of Epeorus (Caucasiron) (Ephemeroptera, Heptageniidae) in Iran, with descriptions of three new species.

Combining morphological and molecular data in an integrative approach, three new mayfly species of Epeorus (Caucasiron) are described. These include Epeorus (Caucasiron) alborzicus Hrivniak & Sroka, sp. nov. and Epeorus (Caucasiron) shargi Hrivniak & Sroka, sp. nov. from northern Iran, and Epeorus (Caucasiron) zagrosicus Hrivniak & Sroka, sp. nov. from central Iran. They are unambiguously delimited using both distance-based and likelihood-based approaches in the analyses of barcode COI sequences. Each new species is compared with other species of the subgenus and morphological diagnostic characters are provided. Based on extensive sampling of streams throughout the country, the distribution and habitat preferences of all Caucasiron species in Iran are assessed. Altogether, there are now six species recorded, among them also E. (C.) nigripilosus Sinitshenkova, 1976 is reported for the first time in Iran. Five species are distributed in the Alborz Mts. in northern Iran, one species was found in the Zagros Mts. in central Iran.

The impact of Miocene orogeny for the diversification of Caucasian Epeorus (Caucasiron) mayflies (Ephemeroptera: Heptageniidae).

A common hypothesis for the high biodiversity of mountains is the diversification driven by orogeny creating conditions for rapid in situ speciation of resident lineages. The Caucasus is a young mountain system considered as a biodiversity hotspot; however, the origin and evolution of its diversity remain poorly understood. This study focuses on mayflies of the subgenus Caucasiron, one of the most diversified stenotopic mayflies inhabiting various types of streams throughout the Caucasus. Using the time-calibrated phylogeny based on two mitochondrial (COI, 16S) and three nuclear (EF-1α, wg, 28S) gene fragments, we tested the role of Caucasian orogeny in biogeography, diversification patterns, and altitudinal diversification of Caucasiron mayflies. We found that orogeny promoted the lineage diversification of Caucasiron in the Miocene. The highest diversification rate corresponding with the uplift of mountains was followed by a significant slowdown towards the present suggesting minor influence of Pleistocene climatic oscillations on the speciation. The Caucasiron lineages cluster into three principal clades originating in the Upper Miocene. We found a strong support that one of the three clades diversified via allopatric speciation in the Greater Caucasus isolated in the Parathetys Sea. The other two clades originating most likely outside the Greater Caucasus diversified towards high and low altitude, respectively, indicating possible role of climatic factors and/or passive uplift on their differentiation. Current high Caucasiron diversity in the Greater Caucasus is a result of in situ speciation and later immigration from adjacent mountain ranges after the Parathetys Sea retreat. Our phylogeny supported the monophyly of Rhithrogeninae, Epeorus s.l., Caucasiron, and Iron. Epeorus subgenus Ironopsis was found paraphyletic, with its European representatives more closely related to Epeorus s.str. than to Iron. Therefore, we re-arranged taxa treated within Ironopsis to comply with the phylogeny recovered herein.

New Oligoneuriidae (Insecta, Ephemeroptera) from Iran.

Two new species of the mayfly family Oligoneuriidae are described based on larval specimens recently collected in Iran. The first new species, Oligoneuriella tuberculata Godunko & Staniczek, sp. nov., can be distinguished from all its congeners by the presence of pronounced protuberances posteromedially on abdominal terga, highly reduced paracercus, large lamella of gill I, and setation on hind margin of middle and hind femora confined to their basal halves. The second species, Oligoneuriopsis villosus Bojková, Godunko, & Staniczek, sp. nov., remarkably belongs to a mostly Afrotropical genus. The new species clearly differs from all its congeners in the shape of setae on the surface of gills and terga, pattern of body colouration, and the shape of posterolateral projections of abdominal segments. Except for the species description, the generic diagnosis of Oligoneuriopsis Crass, 1947 is briefly discussed. COI barcode sequences of both new species are provided and molecular species delimitation is tested using distance-based and likelihood-based approaches, with both new species unambiguously recognised as separate lineages. The analysis of COI also corroborates the respective affinities of both new species, estimated based on morphology. The two new species of Oligoneuriidae described herein highlight the importance of the Middle East as a centre of diversity of this mayfly family within the Palaearctic.

A new Epeorus (Caucasiron) (Ephemeroptera: Heptageniidae) species from Turkey based on molecular and morphological evidence.

Epeorus (Caucasiron) turcicus sp. nov. is described based on larvae from NE Turkey. The new species can be distinguished from other Caucasiron by a unique combination of several diagnostic characters: the presence of a rounded hypodermal medial femur spot, colouration of abdominal terga and sterna, narrow gill plate VII, fine hair-like setae on the surface of abdominal terga, and absence of postero-lateral projections on tergum X. In addition to morphological analysis, two single-locus analytical approaches are employed for delimiting the new species using COI sequences (Automatic Barcode Gap Discovery, ABGD; and General Mixed Yule Coalescent Model, GMYC). Both approaches unambiguously recognized E. (C.) turcicus sp. nov. as a distinct species. Our molecular dataset contains all Caucasiron species occurring in the Caucasus and the delimitation of individual species mostly follows the morphologically defined species. This study confirms the suitability of the GMYC approach for species delimitation within Caucasiron.

Diversity of Armenian mayflies (Ephemeroptera) with the description of a new species of the genus Ecdyonurus (Heptageniidae).

We provide the first commented checklist of Armenian mayflies, based on all relevant literature and recent extensive sampling of 72 localities throughout Armenia during 2011, 2014, and 2015. Altogether 46 species are listed, eight of them reported from Armenia for the first time. One new species, Ecdyonurus (Ecdyonurus) eurycephalus sp. nov. is described (larva and male imago) based on morphological and molecular (COI) data. The species is characterized by a unique head shape in male imago and by the presence of tracheal filaments on gill plate VII in the larval stage.

'Rolling' stoneflies (Insecta: Plecoptera) from mid-Cretaceous Burmese amber.

This contribution describes seven new species of fossil stoneflies from Cretaceous Burmese amber, all of which are dedicated to present and past members of the Rolling Stones. Two species-Petroperla mickjaggeri gen. nov. sp. nov. and Lapisperla keithrichardsi gen. nov. sp. nov.-are placed in a new family Petroperlidae within the stemline of Systellognatha. The first Cretaceous larval specimen of Acroneuriinae, Electroneuria ronwoodi gen. nov. sp. nov., is also described along with another four new species that are placed within the Acroneuriinae genus Largusoperla Chen et al., 2018: Largusoperla charliewattsi sp. nov., Largusoperla brianjonesi sp. nov., Largusoperla micktaylori sp. nov., and Largusoperla billwymani sp. nov. Additional specimens of Acroneuriinae are described without formal assignment to new species due to insufficient preservation. Implications for stonefly phylogeny and palaeobiogeography are discussed.