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.

Lycopsid–arthropod associations and odonatopteran oviposition on Triassic herbaceous Isoetites

“…Ovipositional scars of endophytically inserted eggs were found on the leaf surfaces of the herbaceous lycopsids Isoetites madygensis and I. sixteliae from the Middle-Late Triassic Madygen Formation of Kyrgyzstan. Based on comparisons of ovipositional scar features of fossil and recent dragonflies, it is likely that the eggs were laid by a member of the Archizygoptera, an extinct suborder of dragonflies (order Odonatoptera) that included major Paleozoic taxa (Nel et al, 2012). This interpretation is supported by the occurrence of several fossil forms of the OdonatopteraArchizygoptera (Table 2) in the Madygen fossil biota.…”

“…The fossil record of Odonata from the Madygen Formation consists primarily of well-preserved wings of protozygopteran insects (Pritykina, 1981;Nel et al, 2001Nel et al, , 2002Nel et al, , 2005Shcherbakov, 2008). Based on the most recent taxonomic reevaluation of the order Odonatoptera proposed by Nel et al (2012), fossil dragonflies from Madygen are part of the extinct suborder Archizygoptera Handlirsch 1908. They were mainly represented by gracile, damselfly-like insects with petiolate wings (Nel et al, 2012), which are associated with relatively small egg sizes, such as the fossils described here.…”

“…Based on the most recent taxonomic reevaluation of the order Odonatoptera proposed by Nel et al (2012), fossil dragonflies from Madygen are part of the extinct suborder Archizygoptera Handlirsch 1908. They were mainly represented by gracile, damselfly-like insects with petiolate wings (Nel et al, 2012), which are associated with relatively small egg sizes, such as the fossils described here. Moreover, some ovipositional scars on the leaves of Isoetites madygensis ( Fig.…”

Bechala sommeri Ilger & Brauckmann, 2012 enlightens the Namurian griffenfly diversity (Insecta: Odonatoptera: Bechalidae)

“…Together with the Westphalian Protozygoptera Bechlya ericrobinsoni Jarzembowski & Nel, 2002, Bechala confirms the presence of damselfly-like Odonatoptera in the early Late Carboniferous belonging to different clades that probably were living in rather similar way as the modern Zygoptera, predating on small insects or other arthropods (in relation to its size) in forest environments (similarly to the recent small damselflies), unlike the larger Meganeuridae or Namurotypidae that certainly could not fly between the trunks of the trees and were probably hunting above the large lakes (Jarzembowski & Nel 2002;Nel et al 2012). The same ecological niche was successively occupied by the Late Carboniferous Bechala and Bechlya, later by other Protozygoptera (a group that was flourishing during the Permo-Triassic and still present in the Early Cretaceous), and by the Zygoptera after the Triassic.…”

How to date a dragonfly: Fossil calibrations for odonates

“…Some of these were closely related to modern Odonata. For example, the protozygopterous genera Bechlya (307 Ma) and Luiseia (299 Ma) (Jarzembowski and Nel, 2002;Nel et al, 2012), already had typical odonate wing venational characters like the pterostigma, nodus, and discoidal cell. However, 2 crown group Odonata are much younger, having diverged sometime after the late Triassic and greatly diversified during the Cretaceous according to the fossil evidence.…”