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.

The temporal origin of Madagascar’s extraordinary endemic diversity is debated. A preference for Cenozoic dispersal origins has replaced the classical view of Mesozoic vicariance in the wake of molecular dating. However, evidence of ancient origins is mounting from arthropod groups. Using phylogenetic ‘tip-dating’ analysis with fossils, we show that a whirligig beetle species, Heterogyrus milloti, inhabiting forest streams in southeastern Madagascar is the last survivor of a once dominant and widespread Mesozoic group. With a Late Triassic to Early Jurassic origin (226–187 Ma) it is the hitherto oldest dated endemic lineage of animal or plant on Madagascar. Island biotas’ sensitivity to extinction is well known, but islands can also provide refuge from continental extinction. Heterogyrus milloti is an irreplaceable link to the freshwater biota of the Mesozoic and serves as a reminder of what may be lost without critical conservation efforts on Madagascar.

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The present paper is devoted to an overview on fossil Coleoptera studied from Inner Mongolia, Daohugou (Middle Jurassic, Jiulongshan Formation) and Liaoning (Upper Jurassic‐Lower Cretaceous, Yixian Formation) deposited in Chinese collections. As a result, species of the tribe Sperchopsini and Hydrophilini from Hydrophilidae, families and subfamilies Silphidae, Syndesinae from Lucanidae, Pleocomidae, Trogidae, Trogissitidae, Pyrochroidae, Diaperinae from Tenebrionidae, and Cerambycidae were first registered in the Mesozoic and some families were defined as new. It was shown that many superfamilies represented in the Recent Fauna were formed within the Middle Jurassic and Lower Cretaceous. The materials examined confirm the hypothesis that Cucujiformian beetles are a younger group than other infraordera of Polyphaga (Staphyliniformia and Elateriformia) and, therefore, they appeared in the fossil record only in the late Mesozoic. It was shown and confirmed that most superfamilies appeared in the fossil records before Cucujoidea. The synonymy of Notocupes Ponomarenko, 1964; Sinocupes Lin, 1976, syn. nov.; Amblomma Tan, Ren et Liu 2005, syn. nov.; Euryomma Tan, Ren et Shih, 2006, syn. nov., non Stein, 1899 and Ovatocupes Tan et Ren, 2006, syn. nov.; synonymy of Tetraphalerus Waterhouse, 1901 and Odontomma Tan, Ren et Ge 2006, syn. nov.; and synonymy of Priacmopsis Ponomarenko, 1966 and Latocupes Tan et Ren, 2006, syn. nov. are proposed. Sinorhombocoleus papposus Tan et Ren, 2009 is transferred from the family Rhombocoleidae to Schizophoridae. Cervicatinius complanus Tan, Ren et Shih, 2007 and Forticatinius elegans Tan, Ren et Shih, 2007 are transferred from the family Catiniidae (suborder Archostemata) to superfamily Cleroidea (suborder Polyphaga: first among the family Peltidae and second as a closely related group to the latter family). The family Parandrexidae is transferred from the superfamily Cucujoidea to Cleroidea. The ecological circumstances of the past ecosystems and hypotheses of historical development of the order Coleoptera are discussed. The age of faunas examined is considered. The list of the taxa described from Daohugou and Liaoning is compiled.