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Meganisoptera: Meganeurida: Meganeura monyi BROGNIART, 1893, holotype. Late Pennsylvanian, Commentry, France. Muséum national d'Histoire naturelle, Paris. Muscle attachments seen through skeleton marked by circles. Proxalaria (PR) free, preserved only as shapeless "bumps". All axalaria (AX) fused to fulcalaria (F). Anterior plate and posterior plate similar to those in Figs. 5 and 6A, B, row-sclerites above blood pathways fused with sutures.-After KUKALOVÁ-PECK (1991, fig. 6.15D), updated. Total wing length about 30 cm.
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A b s t r a c t Three extremely rare fossil protodonatoid dragonfly nymphs are described from the middle Pennsylvanian (Moscovian) of Mazon Creek, Illinois: Dragonympha srokai n. gen., n. sp. (Meganisoptera), a large, nearly com-plete young nymph with an extended labial mask and uplifted wing pads; Alanympha richardsoni n. gen., n. sp. (Meganisopte...
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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, M...
Phylogenetic relationships of basal insects remain a matter of discussion. In particular, the relationships among Ephemeroptera, Odonata and Neoptera are the focus of debate. In this study, we used a next-generation sequencing approach to reconstruct new mitochondrial genomes (mitogenomes) from 18 species of basal insects, including six representat...
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Citations
... The Meganeuridae certainly had aquatic larvae, as known at Mazon Creek (Moscovian) (Kukalová-Peck, 2009). In some cases (Laveineopteris or Alethopteris), the rachis was probably still alive when the insect spawned, because shows a plant reaction ring (Fig. 7). ...
Plant-animal interactions shed light on the ecology of the rich insect community from the middle Pennsylvanian basin of Northern France (Nord-Pas-de-Calais coal basin, Hauts-de-France). The data set derives from coal bed layers locally named the Bruay Formation and were collected on the slag heap. This work is a meta-analysis of the different types of damages caused by arthropods, especially insects, consisting of endophytic oviposition scars, holes, galls, bulges, and undetermined traces. These damage traces are associated with host plants belonging to Lycopsida, Medullosales, Pinopsida, and Polypodiopsida. The paleoenvironmental framework and mutualism within each community are also discussed. Based on the present discoveries in Northern France, these interactions may have occurred stratigraphically from Westphalian B (Anzin Formation) until to Westphalian C (Bruay Formation). Despite the limited fossilization potential and sampling difficulties due to slag heap bias, occurrences may be compared with other European localities of the Carboniferous age.
... Aquatic or semiaquatic larvae with abdominal tracheal gills occur in both extant lineages of the clade Palaeoptera, comprising dragonflies (Odonata) and mayflies (Ephemeroptera). Their stem groups and earliest representatives recorded since the late Carboniferous or early Permian are also characterized by aquatic larvae 16,17 . Though there is a Paleozoic record of aquatic immatures for several polyneopteran lineages, like the stem group of Plecoptera or some enigmatic "Grylloblattida" (Lemmatophoridae), their higher phylogenetic relationships, especially in respect of the crown group of Plecoptera remain unresolved 10,18,19 . ...
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.
... These considerations on the large size of Meganisoptera, however, only refer to their flight and not on the growth during the larval stage that may have been influenced by hyperoxia (Verbeck et al., 2011). The maximum lengths of larval stages of the Meganisoptera are poorly known (Kukalová-Peck, 2009), as are the effects of predation which may have selected for larger larvae (and consequently) adults at the greatest size that the environment could support. It is worth noting that 'giant' beetles (i.e. ...
Adult Odonatoptera are among the most efficient flying predators. They have retained many physical characteristics over an immense period stretching from the Carboniferous to the present. Over this time they have greatly varied in size and mass, as shown in the fossil record and in particular by the length, shape, and structure of their wings. A fossil of Meganeurites gracilipes indicates that this large ‘griffenfly’ had a ‘hawker’ hunting behavior similar to certain extant species, with long periods of flight in which power, thermoregulation, and respiration would therefore tend to a ‘steady state’ equilibrium, allowing oxygen requirements and tracheole volumes to be projected and compared to extant ‘hawkers’. Comparing these values with standard pO2 models allows paleo-atmospheric density estimates to be derived. The results suggest that paleo-air pressure has varied from over two bars in the Late Carboniferous, Late Permian, and Middle to Late Jurassic, with lower values in the Early Triassic and Early Jurassic.
... The best-known examples of Paleozoic palaeopteran insects are giant predators such as griffenflies (Meganisoptera), which reached wingspans of approximately 70 cm; however, the majority of Carboniferous and Permian species had dimensions comparable to those of their modern relatives. A recent analysis incorporating fossil species of major palaeopteran groups recovered a monophyletic Hydropalaeoptera, including Bojophlebiidae + (Odonatoptera + Panephemeroptera), as well as supporting Palaeoptera as a clade (163), corroborating estimates based on morphological and genomic analyses (19,77,92,157). The Late Paleozoic Palaeodictyopterida was resolved as monophyletic and recognized as a sister group to Neoptera. ...
... Small damselfly-like Odonatoptera with subpetiolate or petiolate wings appeared during the Carboniferous convergently in several lineages, such as Enigmapteridae, Bechlyidae, or even the meganisopteran Bechalidae (98). Fossil larvae are rare, but evidence from the early instar meganisopteran Dragonympha srokai from Mazon Creek shows the presence of a labial mask and filamentous lateral gill-like appendage on abdominal segments I-VII comparable to the extant Euphaeidae and Polythoridae (77). These lateral gill-like projections are considered to be possible homologs of tergalia in mayflies and presumably also abdominal lateral outgrowths (flaps) in palaeodictyopterans (129, 168). ...
While Mesozoic, Paleogene, and Neogene insect faunas greatly resemble the modern one, the Paleozoic fauna provides unique insights into key innovations in insect evolution, such as the origin of wings and modifications of postembryonic development including holometaboly. Deep-divergence estimates suggest that the majority of contemporary insect orders originated in the Late Paleozoic, but these estimates reflect divergences between stem groups of each lineage rather than the later appearance of the crown groups. The fossil record shows the initial radiations of the extant hyperdiverse clades during the Early Permian, as well as the specialized fauna present before the End Permian mass extinction. This review summarizes the recent discoveries related to the documented diversity of Paleozoic hexapods, as well as current knowledge about what has actually been verified from fossil evidence as it relates to postembryonic development and the morphology of different body parts.
... Entomologists working on different lineages of Pterygota sometimes use subtle modifications of the Comstock-Needham-Redtenbacher vein nomenclature, but overall the fundamentals of the present system differ little from that established by Comstock and Needham. Kukalová-Peck and colleagues (1978, 1983, 1991, 1997, 2009Kukalová-Peck and Richardson, 1983;Riek and Kukalová-Peck, 1984;Kukalová-Peck and Brauckmann, 1992;Haas and Kukalová-Peck, 2001;Kukalová-Peck and Lawrence, 2004;Kukalová-Peck et al., 2009) have provided the most extensive recent revision of the Comstock-Needham system as well as other hexapod appendicular structures, attempting to incorporate considerable, albeit controversial (Béthoux and Briggs, 2008;, evidence from paleontological data. The Kukalová-Peck venational modification effectively considers all the longitudinal sectors to have been paired in the ancestral insect wing, and relies on an archetype with some hypothetical veins not present in any modern or fossil wing (refer to Discussion, below). ...
We present a phylogeny of Chrysopidae inferred from combining molecular and morphological data. Apochrysinae were recovered as sister to the rest of the family (Nothochrysinae and Chrysopinae). The monogeneric tribe Nothancylini is confirmed as sister to the remaining Chrysopinae. The other four tribes are grouped in two clades: Belonopterygini + Leucochrysini and Ankylopterygini + Chrysopini. The Nineta-group is herein transferred from Chrysopini to Ankylopterygini. Within the diverse and species-rich Chrysopini we recognize five genus-group clades: Chrysopa, Chrysoperla, Chrysopodes, Eremochrysa and Meleoma generic groups. The mapping of characters, such as the parameres, the tignum, the im cell or the tibial spurs, on the phylogeny provides insights into the evolution of these traits in the family. In addition, we propose the following taxonomic changes to the classification of the family: the inclusion of Chrysopidia, Nineta and Tumeochrysa in the tribe Ankylopterygini, and the synonymization of Furcochrysa with Chrysopa.
... The labial mask itself exhibits considerable detail including the claw-like paraglossae, the rimmed and laterally attached labial palps with outwardly directed teeth, and almost all of the articulatory sclerites of the mentum, suggesting a modern-aspect strike apparatus. Kukalová-Peck (2009) suggests that Dragonympha was semiaquatic and that the Labial Mask mouthparts originated as a capture device for terrestrial prey, later co-opted to secure prey in aquatic environments. Interestingly, a similar extrudable labial mask device has independently arisen in stenine rove beetles (Coleoptera: Staphylinidae) (Schmitz 1943), but the mechanism of extension is hydraulically based and slingshot in mode, rather than rapid extension of a multi-jointed apparatus (Weinreich 1968). ...
The mouthparts of insects are a phenomenal example of a multi-element, modular, feeding apparatus that repeatedly has been modified structurally to perform every feeding function imaginable in the terrestrial and freshwater realms, a process that began in the Early Devonian. Insect mouthparts have been structured to chew, pierce and suck, siphon, lap, sponge, bore, and mine on and within a wide variety of tissues, as well as filter, sieve, and collect particulate food such as plankton and pollen. Thirty-seven fundamental mouthpart classes perform these roles in the modern and fossil record, a result that has been expanded somewhat from earlier, phenetic cluster analyses of modern insect mouthparts. A broad survey of fossil insect mouthparts, in conjunction with the phenetic mouthpart analysis, revealed patterns of mouthpart innovation occurring in bursts of cladogenesis separated from intervals of rather static mouthpart morphology. For the Paleozoic Era, based on direct (body fossil) and indirect (trace fossil) evidence, and commencing during the Devonian Period, the four earliest mouthpart classes were present, accounting for 11.4% of all mouthpart classes in the fossil record. In the succeeding Mississippian Subperiod, no mouthparts are documented; the four mouthparts from the Devonian continue into the succeeding Pennsylvanian Subperiod. During Pennsylvanian time, there was a spectacular burst of new mouthpart classes, coincident with the appearance of approximately 15 major insect lineages. By the end of the period, 29.7% of all insect classes had appeared. The following Permian Period added another seven mouthpart classes, particularly those from early hemimetabolous and holometabolous lineages, resulting in 48.6% of all mouthpart classes present. The profound ecological crisis at the end of the Permian notably saw the near extirpation of only one mouthpart class, the Robust Beak of piercing-and-sucking paleodictyopteroid insects, which eventually was extinguished sometime during the ensuing Triassic Period. For the Mesozoic Era, the Triassic Period added another seven mouthpart classes, particularly involving aquatic naiads and larvae, and early dipteran mouthparts, resulting in 67.6% of all mouthpart classes at the end of the period. During the Jurassic, the Mesozoic Lacustrine Revolution had begun, reaching a peak in the invasion of freshwater ecosystems that commenced during the Late Triassic, but undergoing a major diversification of mouthparts in terrestrial lineages, resulting in 83.3% of all mouthpart classes present, notably before the ecological expansion of angiosperms in the subsequent Early Cretaceous. The Jurassic also was a time for the origin and initial innovation of mouthpart design in early Siphonaptera, and a largely parallel diversification event among hematophagous Diptera; both processes continued into the Early Cretaceous. The Cretaceous Period exhibits a considerable diversity in compression deposits and especially amber deposits, preserving relict lineages that bore mouthparts at a Permian and Triassic stage of evolution as well as new lineages with bizarre mouthpart structures that are difficult to place among existing mouthpart classes. During the Cretaceous, three new mouthparts classes are added, yielding 97.1% of all mouthparts at the end of the period. For the Cenozoic Era, no mouthpart classes are added during the Paleogene Period, and only one mouthpart class, lacking a fossil record, is added during the Neogene Period. During this time, there is modification and expansion of mouthpart classes established during the mid Mesozoic and the development of special mouthpart elements involved in leaf mining, blood feeding, and pollination.
... While the fossil record of hexapods extends to the Early Devonian, the first tangible evidence of aquatic insect specializations is documented from the Early Permian in some stem mayflies of Permoplectoptera and stoneflies (Plecoptera) [1][2][3]. Although the prior records of stem-group representatives of Ephemeroptera and Odonata with putatively aquatic larvae, are documented since the Late Carboniferous [4], the evidence is uncertain, perhaps with the exception of a meganeurid griffenfly larva Dragonympha srokai bearing lateral abdominal tracheal gills from the Late Carboniferous of the Mazon Creek 'Konservat-Lagerstätte' [5]. Based on their morphological specializations, most aquatic Permian immature and adult insects indicate lotic palaeoenvironmental conditions while the evidence from lentic habitats is lacking prior to the late Permian [6,7]. ...
... 6: 190460 This issue is made all the more interesting as the phylogenetic relationships of the main pterygote insect lineages remain controversial, often dubbed the 'Palaeoptera problem' [24]. Palaeodictyopterida are traditionally considered as an early diverging group of Pterygota, either as the sister group to a putatively monophyletic Palaeoptera (the group comprising extant dragonflies and mayflies and their fossil relatives) [5,25], or resolved as sister group to Neoptera (all flying insects with the ability to fold their wings over the abdomen) on the basis of a recent phylogenetic analysis [26]. Interestingly, Palaeodictyopterida have not been recovered as sister group to all other Pterygota, i.e. to a Palaeoptera + Neoptera clade. ...
The Late Palaeozoic insect superorder Palaeodictyopterida exhibits a remarkable disparity of larval ecomorphotypes, enabling these animals to occupy diverse ecological niches. The widely accepted hypothesis presumed that their immature stages only occupied terrestrial habitats, although authors more than a century ago hypothesized they had specializations for amphibious or even aquatic life histories. Here, we show that
different species had a disparity of semiaquatic or aquatic specializations in larvae and even the supposed retention of abdominal tracheal gills by some adults. While a majority of mature larvae in Palaeodictyoptera lack unambiguous lateral tracheal gills, some recently discovered early instars had terminal appendages with prominent lateral lamellae like in living damselflies, allowing support in locomotion along with respiratory function. These results demonstrate that some species of Palaeodictyopterida had aquatic or semiaquatic larvae during at least a brief period of their post-embryonic development. The retention of functional gills or gill sockets by adults indicates their amphibious lifestyle and habitats tightly connected with a water environment as is analogously known for some modern Ephemeroptera or Plecoptera. Our study refutes an entirely terrestrial lifestyle for all representatives of the early diverging pterygote group of Palaeodictyopterida, a greatly varied and diverse lineage which probably encompassed
many different biologies and life histories.
... On account of the critical role in maneuvering flights, the wing base structure has been well studied and documented for the extant Odonata (Sargent, 1937;Chao, 1953;Asahina, 1953;Pfau, 1986Pfau, , 1991Genet et al., 2013;. Due to preservation, the wing bases in the fossil taxa were rarely described except for several protodonatans: Alanympha richardsoni, Eugeropteron sp., Eugeropteron lunatum and Meganeura monyi (Riek & Kukalov a-Peck, 1984;Kukalov a-Peck, 1991, 2009Kukalov a-Peck et al., 2009). However, the wing bases of fossil Odonata have never been described hitherto. ...
A new fossil burmaeshnid, Proaeschna zhangi gen. et sp. nov., is described from the mid-Cretaceous Burmese amber. Its well-preserved wing base structure allows us to illustrate the morphological details of wing base in the fossil Odonata for the first time. The proximal costal plate in the new species consists of the anterior and posterior parts which are fused together with a suture. The anterior proximal costal plate in P. zhangi and extant Odonata is homologized to the basalare of Protodonata. In addition, P. zhangi has developed the semi-detached scutal plate that is similar as the extant Aeshnidae. The traces of venal basivenalia in weakly sclerotized axillary plate contribute useful information to interpret the venation of wing base within Odonata.
... Entomologists working on different lineages of Pterygota sometimes use subtle modifications of the Comstock-Needham-Redtenbacher vein nomenclature, but overall the fundamentals of the present system differ little from that established by Comstock and Needham. Kukalová-Peck and colleagues (1978, 1983, 1991, 1997, 2009Kukalová-Peck and Richardson, 1983;Riek and Kukalová-Peck, 1984;Kukalová-Peck and Brauckmann, 1992;Haas and Kukalová-Peck, 2001;Kukalová-Peck and Lawrence, 2004;Kukalová-Peck et al., 2009) have provided the most extensive recent revision of the Comstock-Needham system as well as other hexapod appendicular structures, attempting to incorporate considerable, albeit controversial (Béthoux and Briggs, 2008;, evidence from paleontological data. The Kukalová-Peck venational modification effectively considers all the longitudinal sectors to have been paired in the ancestral insect wing, and relies on an archetype with some hypothetical veins not present in any modern or fossil wing (refer to Discussion, below). ...
The wings of insects are one of their most prominent features and embody numerous characters and modifications congruent with the variety of their lifestyles. However, despite their evolutionary relevance, homology statements and nomenclature of wing structures remain understudied and sometimes confusing. Early studies on wing venation homologies often assumed Neuropterida (the superorder comprising the orders Raphidioptera, Megaloptera, and Neuroptera: Snakeflies, alderflies and dobsonflies, and lacewings) to be ancient among Pterygota, and therefore relied on their pattern of venation for determining groundplans for insect wing venation schemata and those assumptions reciprocally influenced the interpretation of lacewing wings. However, Neuropterida are in fact derived among flying insects and thus a reconsideration of their wings is crucial. The identification of the actual wing venation of Neuropterida is rendered difficult by fusions and losses, but these features provide systematic and taxonomically informative characters for the classification of the different clades within the group. In the present study, we review the homology statements of wing venation among Neuropterida, with an emphasis on Chrysopidae (green lacewings), the family in which the highest degree of vein fusion is manifest. The wing venation of each order is reviewed according to tracheation, and colored schemata of the actual wing venation are provided as well as detailed illustrations of the tracheation in select families. According to the results of our study of vein tracheation, new homology statements and a revised nomenclature for veins and cells are proposed.
... The morphology of various body parts such as the external genitalia and, notably, the pattern of wing venation in Palaeodictyoptera correspond in some traits to those of extant mayflies (Ephemeroptera) and dragonflies (Odonata) (Figure 1). While extant immatures of these latter groups show derived traits such as dorsally placed wing pads or their rotation (in Odonata), their Paleozoic stem-group relatives such as griffenflies (Protodonata) appear to have also possessed articulated wing pads [22], implying that this form in protodonatans and palaeodictyopterans is plesiomorphic. ...
The appearance of wings in insects, early in their evolution, has been one of the more critical innovations
contributing to their extraordinary diversity. Despite the conspicuousness and importance of wings, the origin of these structures has been difficult to resolve and represented one of the ‘‘abominable mysteries’’ in evolutionary biology. More than a century of debate has boiled the matter down to two competing alternative —one of wings representing an extension of the thoracic notum, the other stating that they are appendicular derivations from the lateral body wall. Recently, a dual model has been supported by genomic and developmental data, representing an amalgamation of elements from both the notal and pleural hypotheses. Here, we reveal crucial information from the wing pad joints of Carboniferous palaeodictyopteran insect nymphs using classical and high-tech techniques. These nymphs had three pairs of wing pads that were medially articulated to the thorax but also broadly contiguous with the notum anteriorly and posteriorly (details unobservable in modern insects), supporting their overall origin from the thoracic notum as well as the expected medial, pleural series of axillary sclerites. Our study provides support for the formation of the insect wing from the thoracic notum as well as the already known pleural elements of the arthropodan leg. These results support the unique, dual model for insect wing origins and the convergent reduction of notal fusion in more derived clades, presumably due to wing rotation during development, and they help to bring resolution to this long-standing debate.
