Günter Bechly

Phylogeny and classification of the Stenophlebioptera (Odonata: Epiproctophora)

Abstract The Juraheterophlebiidae, new family of the “heterophlebioid” lineage, the Henrotayiidae, new family of the “anisopteroid” lineage, the Prostenophlebiidae and the Liassostenophlebiidae, new families of the Stenophlebioptera, and three new genera and species of the Stenophlebiidae are described from the Mesozoic of Germany, Spain, England, Kazakhstan, and Mongolia. The phylogenetic positions of the families Erichschmidiidae and Gondvanogomphidae are discussed. A tentative phylogenetic analysis of the Anisopteromorpha is proposed. This significantly extends our knowledge on the palaeogeographical distribution of the Stenophlebioptera and the Epiproctophora (“dragondamselflies”).

The enigmatic Mesozoic insect taxon Chresmodidae (Polyneoptera): New palaeobiological and phylogenetic data, with the description of a new species from the Lower Cretaceous of Brazil

The morphology of the enigmatic, Mesozoic, aquatic insect family Chresmodidae is redescribed and its phylogenetic affinities among the polyneopterous orders discussed. Study of the complete venation of both fore- and hind wings observed in some specimens from the Spanish Barremian, permit us to postulate the hypothesis that the family belongs to the Archaeorthoptera, thus to the orthopteroid lineage rather than to crown-group Phasmatodea or to the more inclusive group Holophasmatodea (sensu GRIMALDI &E NGEL, 2005). New specimens from Spain, Lebanon, Brazil, and Germany permit a new re-description of some chresmodid body structures with concomitant implications for the phylogenetic position of the family. Chresmoda neotropica n. sp. is described from the Aptian-Albian of the Crato Formation (northeast Brazil). The functional morphology proposed for some of their specialized structures suggest a new hypothesis of Chresmoda palaeo- biology, and related to this some implications for the localized palaeoenvironment as well as global palaeoclimate. The problematic Sternarthron spp. from the Upper Jurassic of Solnhofen were described as probable palpigrades (Arachnida: Palpigradi), based on type material originally thought to be fossil insects. The affinities of Sternarthron HAASE, 1890 have been questioned. Our restudy of HAASEs types clearly confirmed earlier assumptions that these fossils represent nymphal specimens of chresmodids. Consequently, Sternarthron has to be considered as an invalid junior synonym of the fossil insect genera Propygolampis WEYENBERGH, 1874 and Chresmoda GERMAR, 1839.

Coxoplectoptera, a new fossil order of Palaeoptera (Arthropoda: Insecta), with comments on the phylogeny of the stem group of mayflies (Ephemeroptera)

Mickoleitia longimanus gen. et sp.n. is described from the Lower Cretaceous limestone of the Crato Formation in Brazil. It is attributed to a new family Mickoleitiidae and a new fossil insect order Coxoplectoptera within the palaeopterous Ephemerida, based on the presence of an elongated costal brace. This fossil insect exhibits a very peculiar combination of derived characters like specialized forelegs with strongly elongated, free coxae, single-clawed pretarsus, and distinctly skewed pterothorax as in dragonflies. On the other hand, several plesiomorphies are present that exclude this taxon from modern Ephemeroptera, namely large hind wings with widened anal area and numerous cross veins that separate the elongate costal brace from the costal margin. Fossil larvae described by Willmann as larval Cretereismatidae are herein attributed to Mickoleitiidae fam.n., based on the shared presence of broad hind wing buds with distinctly broadened anal area, wing bud venation similar to the adult holotype, and subchelate forelegs with elongate free coxae. These larvae are also highly autapomorphic in the structure of their abdominal gills and laterally flattened body with vertically oval section that is unique within Ephemerida. On the other hand they possess plesiomorphic lateral wing pads with pronounced articulation like Palaeozoic pterygote larvae, while wing pads in modern insects are always secondarily fused to the tergum. A similar fossil larva from the Jurassic of Transbaikals was earlier described as Mesogenesia petersae and classified within modern mayflies. It is herein attributed to Mickoleitiidae fam.n. Coxoplectoptera are recognized as putative sister group of modern Ephemeroptera based on the shared presence of only 7 pairs of abdominal gills, while Permoplectoptera still have retained 9 pairs of gills. The phylogenetic reclassification of the mayfly stem group by Willmann is critically discussed and modified.

New fossil insect order Permopsocida elucidates major radiation and evolution of suction feeding in hemimetabolous insects (Hexapoda: Acercaria)

With nearly 100,000 species, the Acercaria (lice, plant lices, thrips, bugs) including number of economically important species is one of the most successful insect lineages. However, its phylogeny and evolution of mouthparts among other issues remain debatable. Here new methods of preparation permitted the comprehensive anatomical description of insect inclusions from mid-Cretaceous Burmese amber in astonishing detail. These “missing links” fossils, attributed to a new order Permopsocida, provide crucial evidence for reconstructing the phylogenetic relationships in the Acercaria, supporting its monophyly, and questioning the position of Psocodea as sister group of holometabolans in the most recent phylogenomic study. Permopsocida resolves as sister group of Thripida + Hemiptera and represents an evolutionary link documenting the transition from chewing to piercing mouthparts in relation to suction feeding. Identification of gut contents as angiosperm pollen documents an ecological role of Permopsocida as early pollen feeders with relatively unspecialized mouthparts. This group existed for 185 million years, but has never been diverse and was superseded by new pollenivorous pollinators during the Cretaceous co-evolution of insects and flowers. The key innovation of suction feeding with piercing mouthparts is identified as main event that triggered the huge post-Carboniferous radiation of hemipterans, and facilitated the spreading of pathogenic vectors.

New predatory cockroaches (Insecta: Blattaria: Manipulatoridae fam.n.) from the Upper Cretaceous Myanmar amber

Abstract We describe a new extinct lineage Manipulatoridae (new family) of cockroaches from the Upper Cretaceous (Cenomanian) amber of Myanmar. Manipulator modificaputis gen. et sp. n. is a morphologically unique extinct cockroach that represents the first (of a total of 29 known worldwide) cockroach family reported exclusively from the Myanmar amber. This family represents an early side branch of the stem group of Mantodea (most probably a sister group of Eadiidae within Blattaria/Corydioidea) because it has some synapomorphies with the Mantodea (including the stem group and Eadiidae). This family also retains symplesiomorphies that exclude a position in the crown group, and furthermore has unique autapomorphies that exclude a position as a direct ancestor of Mantodea. The unique adaptations such as strongly elongated extremities and freely movable head on a long neck suggest that these animals were pursuit predators. Five additional specimens (including two immatures) reported from the Myanmar amber suggest that this group was relatively rare but belonged to the indigenous and autochthonous inhabitants of the ancient amber forest of the Myanmar region.

Systematics and Evolution of Paleozoic And Mesozoic Damselfly-Like Odonatoptera of the ‘Protozygopteran’ Grade

Abstract The Paleozoic to Mesozoic grade ‘Protozygoptera’ is revised. It appears to be composed of two main lineages, namely the superfamily Permagrionoidea, and the Archizygoptera. The latter taxon forms a monophyletic group together with Panodonata ( = crown-Odonata plus their closest stem-relatives). Therefore, the ‘Protozygoptera’ as previously understood is paraphyletic. Diagnostic characters of the ‘Protozygoptera’, Permagrionoidea, and Archizygoptera are re-evaluated. The Permolestidae is considered as a junior synonym of the Permagrionidae. The following new taxa are described: Permolestes sheimogorai new species, Permolestes soyanaiensis new species, Epilestes angustapterix new species, Solikamptilon pectinatus new species (all in Permagrionidae); Lodeviidae new family (for Lodevia); Luiseiidae new family (including Luiseia breviata new genus and species); Kennedya azari new species, Kennedya pritykinae new species, Kennedya ivensis new species, Progoneura grimaldii new species (all in Kennedyidae); Engellestes chekardensis new genus and species (in Bakteniidae); and Azaroneura permiana new genus and species (in Voltzialestidae). The Kaltanoneuridae and Oboraneuridae are revised. The evolution of protozygopteran Odonatoptera during the transition from the Permian to the Triassic is discussed. The larger taxa of the permagrionoid lineage apparently did not cross through the Permian–Triassic boundary, unlike the more gracile Archizygoptera. This last group shows a remarkable longevity from the late Carboniferous to the Early Cretaceous. It also presents a great taxonomic and morphological stability, with genera ranging from the Permian to the Triassic, and a wing venation pattern nearly unchanged from the late Carboniferous to the Late Triassic. The mass extinction at the end of the Permian period seemingly had a minor effect on these tiny and delicate insects.

Revision and phylogenetic affinities of the Jurassic Steleopteridae Handlirsch, 1906 (Odonata: Zygoptera)

The Jurassic odonate family Steleopteridae is revised. Two new genera and species Parasteleopteron guischardi and Euparasteleopteron viohli are described. The phylogenetic affinities of this group are discussed. The Steleopteridae are excluded from the Epiproctophora and transferred into the Zygoptera (stemgroup). Euphaeopsis multinervis is redescribed and transferred to Epiproctophora: Isophlebioidea, and the genus Pseudoeuphaea with its four species is considered as a nomen dubium in Odonata incertae sedis.

Preliminary phylogenetic analysis of the Protanisoptera (Insecta: Odonatoptera)

Abstract The Permian suborder Protanisoptera (Insecta: Odonatoptera) is revised and a new phylogenetic hypothesis proposed after analyses based on wing venation and different outgroups. After our study the families Camptotaxineuridae and Kaltanoneuridae are excluded from the Protanisoptera. After a new phylogenetic analysis, the family Permaeschnidae is redefined and the families Pholidoptilidae, Polytaxineuridae, Callimokaltaniidae and Hemizygopteridae are restored, as already proposed for the latter three families by Bechly (1996). The new genus Proditaxineura is described. The genus Gondvanoptilon ROSLER et al., 1981 is excluded from the Meganisoptera: Erasipteridae and re-included in the Permaeschnidae, as already proposed by Bechly (1998). Permaeschna proxima MARTYNOV, 1931 is considered as a junior synonym of Permaeschna dolloi MARTYNOV, 1931. Pholidoptilon camense ZALESSKY, 1931 is excluded from Permaeschna MARTYNOV, 1931 and the genus Pholidoptilon Zalessky, 1931a , Zalessky, 1931b is restored. Ditaxineurella stigmalis MARTYNOV, 1940 is excluded from the Hemizygopteridae and considered as a Protanisoptera Incertae sedis.

First Record of Anisoptera (Insecta: Odonata) from mid-Cretaceous Burmese Amber

The fossil dragonfly Burmalindenia imperfecta gen. et sp. nov. is described from mid-Cretaceous Burmese amber as the first record of the odonate suborder Anisoptera for this locality and one of the few records from amber in general. The inclusion comprises two fragments of the two hind wings of a dragonfly. The fossil can be attributed to a new genus and species of the family Gomphidae, presumably in the subfamily Lindeniinae, and features a strange teratological phenomenon in its wing venation.

Revision of the giant pterygote insect Bojophlebia prokopi Kukalová-Peck, 1985 (Hydropalaeoptera: Bojophlebiidae) from the Carboniferous of the Czech Republic, with the first cladistic analysis of fossil palaeopterous insects

The holotype is redescribed of the giant pterygote insect Bojophlebia prokopi Kukalová-Peck, 1985 from the Pennsylvanian of the Czech Republic. Multiple errors in the original description are documented and corrected. Bojophlebia prokopi has neither any visible traces of a costal brace nor an anal brace, but it does show triadic branchings of MA, MP, CuA, and even, as rare a plesiomorphy, of CuP. It is therefore rejected as a fossil stem mayfly and attributed as sister group of all other Hydropalaeoptera. The first cladistic analysis of fossil palaeopterous insects, including different palaeodictyopterid groups, is presented. A revised phylogeny of Hydropalaeoptera and the stem line of Ephemeroptera are suggested. Palaeodictyopterida is recognized as sister group of Neoptera; thus Palaeoptera s.l. is rejected as a paraphyletic taxon. Four new higher taxa – Paranotalia, Euhydropalaeoptera, Neopterygota and Litophlebioidea superfam. nov. – are introduced, as well as the new family Lithoneuridae. http://zoobank.org/urn/urn:lsid:zoobank.org:pub:1B16D6AB-2B29-4891-A61D-7A9212A85493