Alexander G. Kirejtshuk

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.

The most ancient bark beetle known: a new tribe, genus and species from Lebanese amber (Coleoptera, Curculionidae, Scolytinae)

Abstract Cylindrobrotus pectinatusgen. et sp.n., a new scolytine species from Cretaceous Lebanese amber, is described. A new tribe, Cylindrobrotini trib.n., is proposed for this unique species, which demonstrates an unusual combination of some archaic and many advanced characters. This finding suggests that the Scolytinae became a distinct lineage of Curculionoidea from the Lower Cretaceous. Fossil records are reviewed, and some remarks on the origin and taxonomic position of bark and ambrosia beetles are made. Some comments on the various phylogenetic interpretations of the last 30 years are given, particularly in respect of their correspondence with the fossil record. The early appearance of Scolytinae in the fossil record before other Curculionidae (which appeared in the Upper Cretaceous) can be used as evidence against the hypothesis of bark beetles as offspring of weevils. The question of the taxonomic rank of bark beetles (separate subfamily or family) and their placement among other groups of the superfamily remains unsolved.

Current knowledge of Coleoptera (Insecta) from the Lower Cretaceous Lebanese amber and taxonomical notes for some Mesozoic groups

This paper overviews more than 39 families of fossil Coleoptera from Lower Cretaceous Lebanese amber from nine outcrops. Lebanese amber contains the oldest representatives of the families Scydmaenidae (considered by some as a subfamily of Staphylinidae), Ptiliidae, Elodophalmidae, Clambidae, Throscidae, Lebanophytidae fam. n., Ptilodactylidae, Cantharidae, Melyridae, Dasytidae, Dermestidae, Ptinidae, Kateretidae, Erotylidae, Latridiidae, Laemophloeidae, Salpingidae, Anthicidae, Melandryidae, Aderidae, Curculionidae (Scolytinae). The families Chelonariidae and Scraptiidae are known from both Lebanese amber and Baissa, with both sites having a comparable age. The subfamilies Trechinae (Carabidae), Euaesthetinae (Staphylinidae) and Liparochrinae (Hybosoridae) first appear in the fossil record in Lebanese amber. The Coleoptera in Lebanese amber mostly belong to groups with arboreal habits (as found today in wood and tree fungi). Eochelonarium belle gen. et sp. n., Rhizophtoma synchrotronica sp. n., Rhizobactron marinae gen et sp. n. and Atetrameropsis subglobosa gen. et sp. n. are described from Lebanese amber. A new subfamily in the family Cerophytidae is proposed for Aphytocerus communis Zherichin, 1977 (Aphytocerinae subfam. n.) and new genus Baissopsis gen.nov. is erected for Baissophytum amplus Chang, Kirejtshuk et Ren, 2011. Also a new interpretation of the taxon “Lasiosynidae” is provided by placing it as a subfamily in the family Eulichadidae with two genera ( Lasiosyne Tan, Ren et Shih, 2007 and Bupredactyla Kirejtshuk, Chang, Ren et Shih, 2010), while the other genera initially regarded as “Lasiosynidae” were tentatively transferred into Eulichadinae sensu n. ( Mesodascilla Martynov, 1926; Tarsomegamerus Zhang, 2005; Brachysyne Tan et Ren, 2009; Anacapitis Yan, 2009; Parelateriformius Yan et Wang, 2010 and Cretasyne Yan, Wang et Zhang, 2013) with the new synonymy of Tarsomegamerus and Parelateriformius syn. n. The genus Mesaplus Hong, 1983 described in the family Triaplidae is also transvered to Eulichadinae. The genera Artematopodites Ponomarenko, 1990; Dzeregia Ponomarenko, 1985 and Glaphyropteroides Handlirsch, 1906 proposed for species known only by separate elytra and recently included in the “family” Lasiosynidae (Yan et al., 2013) are regarded as Elateriformia incertae sedis . The first insect from the newly discovered outcrops of Nabaa Es-Sukkar – Brissa: Caza (District) Sir Ed-Danniyeh, Mouhafazet (Governorate) Loubnan Esh-Shimali (North Lebanon) is described and the first general description of this outcrop is made.

Family Lasiosynidae n. fam., new palaeoendemic Mesozoic family from the infraorder Elateriformia (Coleoptera: Polyphaga)

Abstract Lasiosynidae n. fam. is proposed for the genera Lasiosyne Tan, Ren & Chih 2007 (transferred from Archostemata to Polyphaga), Anacapitis Yan 2009 Tarsomegamerus Zhang 2005 (proposed in the superfamily Chrysomeloidea) and Bupredactyla n. gen. The new family is regarded in composition of the infraorder Elateriformia without a more detailed attribution, because it demonstrates a mixture of characters of different families and superfamilies, i.e. somehow an intermediate position between the superfamilies Dascilloidea, Elateroidea, Buprestoidea and Byrrhoidea with most resemblance to Dascillidae, Schizopodidae, Eulichadidae, Ptilodactylidae and Callirrhipidae and probable more close relationship to the last three families. Four new fossil species of the genus Lasiosyne: L. daohugouensis n. sp., L. fedorenkoi n. sp., L. gratiosa n. sp., L. quadricollis n. sp., and also Bupredactyla magna n. sp. are described from the Middle Jurassic Jiulongshan Formation of eastern Inner Mongolia, China. A probable generic composition of the new family is considered. The synonymy of generic names Anacapitis Yan 2009 and Brachysyne Tan & Ren 2009, n. syn. as well as synonymy of species names Lasiosyne euglyphea Tan, Ren & Chih 2007, Pappisyne eucallus Tan & Ren 2009, n. syn. and Pappisyne spathulata Tan & Ren 2009, n. syn. are proposed.

Evolution of the elytral venation and structural adaptations in the oldest Palaeozoic beetles (Insecta: Coleoptera: Tshekardocoleidae)

The oldest definitive beetle, Coleopsis archaica gen. et sp. nov., is described from the earliest Permian (Asselian or early Sakmarian) of Germany (Grügelborn/Saarland). Its elytral venation is typical of the Early Permian Tshekardocoleidae. The elytral venation pattern of the type of Moravocoleus permianus Kukalová, 1969 is reconsidered in order to clarify structural peculiarities of the type genus of Moravocoleidae Kukalová-Peck & Beutel, 2011, herein regarded as a junior synonym of Tshekardocoleidae. The new discovery allows reinterpretation of the elytral venation, abdominal laterosternites, and other structures, making it possible to identify a ‘subelytral space’. Tshekardocoleoidea and Labradorocolidae are confirmed as Coleoptera, while Umenocoleidae sensu nov. and Umenocoleoidea are restored to Holometabola as a potential sister group of all Coleoptera. The holotype of Umenocoleus sinuatus Chen & T’an, 1973 needs to be revised in more detail. ‘Moravocoleus’ perditus Kukalová, 1969, regarded by Ponomarenko as a member of the genus Avocoleus, is excluded from Tshekardocoleidae and preliminarily considered as Archostemata incertae sedis. Oborocoleidae is considered as a taxon of more dubious position (order uncertain for Oborocoleus Kukalová, 1969, while Liberocoleus Kukalová, 1969 probably belongs to Archostemata (family uncertain)). Protocoleoptera and Paracoleoptera are considered as synonyms of Protelytroptera, while Archecoleoptera is dismissed as a paraphyletic group. Adiphlebia lacoana Scudder, 1885, previously proposed as the oldest beetle, is not a member of Coleoptera, as confirmed by Kukalová-Peck & Beutel (2012). The composition of beetle suborders and the origin of the order, which probably took place concurrently with the initial diversification of other holometabolan insects, are briefly discussed. http://zoobank.org/urn:lsid:zoobank.org:pub:2044EEFE-B88B-4552-B1F9-983E6F7A583D

First Fossil Click Beetles from the Middle Jurassic of Inner Mongolia, China (Coleoptera: Elateridae)

Abstract. A new genus with two new species and one new species of fossil elaterids are described: Paradesmatus baiae sp. nov., Paradesmatus ponomarenkoi sp. nov., Protagrypnus robustus sp. nov. These findings document the fossils from the Middle Jurassic Jiulongshan Formation of Eastern Inner Mongolia, China. Fossil elaterids of Mesozoic with large triangular plates of metacoxae have been discovered only from the Upper Jurassic strata of Karatau so far, Paradesmatus baiae with the unique feature will expand our knowledge on the early diversification of elaterids.

New beetles of the suborder Polyphaga from the Lowermost Eocene French amber (Insecta: Coleoptera)

Abstract The paper deals with the descriptions of three new genera, one newsubgenus and eleven new species of the superfamilies Scirtoidea, Cleroidea, and Cucujoidea (suborder Polyphaga) originated from the Lowermost Eocene amber, namely: the Scirtidae: Cyphon gallicus nov. sp. and Cyphon lobanovi nov. sp.; the Melyridae: Malachiinae Colotes constantini nov. sp. and C. impexus nov. sp.; the Nitidulidae: Cybocephalinae Pastillocenicus polyaki nov. gen., nov. sp., and P. grandiclavis nov. gen., nov. sp. and P. longifrons nov. gen., nov. sp.; the Kateretidae: Hetherelus expressus nov. sp. and Eoceniιretes yantaricus nov. gen., nov. sp.; the Smicripidae: Smicrips europeus nov. sp.; and the Anthicidae: Eurygeniinae Oisegenius antiquus nov. gen., nov. sp. The systematic positions of these new taxa and hypotheses on their bionomy are discussed.

New genera and new species of leaf beetles (Coleoptera: Polyphaga: Chrysomelidae) from Lowermost Eocene French amber

Abstract Three new monotypical genera of leaf beetles are described: Aoriopsis eocenicus n. gen, n. sp. and Acolaspoides longipes n. gen., n. sp. from the subfamily Eumolpinae, and also Crepidocnema yantarica n. gen., n. sp. from the subfamily Alticinae. Short reviews of known fossil records of the mentioned subfamilies are given and position of the new genera discussed.

New Archostemata (Insecta: Coleoptera) from the French Paleocene and Early Eocene, with a note on the composition of the suborder

Abstract The paper deals with descriptions of four new species of Archostemata from the Paleocene and Earliest Eocene amber, Cupes manifestus n. sp., C. orbiculatus n. sp., C. ponomarenkoi n. sp. and Micromalthus eocenicus n. sp., the latter representing the oldest member of genus Micromalthus Leconte 1878 (Micromalthidae). The position, composition and origin of the genera Cupes and Micromalthus are considered. Cupes seemed to appear during Cretaceous or very close to this period, while Micromalthus could originate somewhat later. The synonymy of Cupedini and Priacmini is grounded. The genus Crowsoniella Pace 1975 (Crowsoniellidae) could be excluded from the suborder Archostemata and transferred into the infraorder Cucujiformia (Polyphaga).

Current knowledge of Coleoptera (Insecta) from the Lowermost Eocene Oise amber

The paper provides an overview on fossil Coleoptera from the Lowermost Eocene Oise amber of the Paris Basin (circa 53 million years old). At present 45 beetle families have been found from this source. This fossil “fauna” has some peculiarities in comparison with other amber deposits. Some of these families are not recorded in older outcrops (Pselaphidae, Smicripidae, Coccinellidae, Ciidae). Some subfamilies and tribes of other groups have their oldest representatives recorded in Oise, i.e., Eurygeniinae (Anthicidae), Inopeplinae (Salpingidae), Trinodini (Dermestidae), Megatominae (Dermestidae), Attageninae (Dermestidae), Brontinae (Silvanidae), Synchitini (Zopheridae) and Opatrini (Tenebrionidae). The genera defined in the “Oise fauna” show various geographical links with relatives in the recent fauna. These alternative links support that the faunistic composition of the early Eocene had a weak zonal differentiation. Scirtes circumcisus sp.n., Boleopsidae fam.n., Boleopsis polinae gen. et sp.n. and Antiphloeus stramineus gen. et sp.n. are described. The synonymy of the genera Scirtes Latreille, 1796 and Eohelodites Hong, 2002 is proposed.