Permian Insects Research Articles

Evolution of Insect Diversity in the Permian and Triassic

The global warming that occurred during the Permo-Triassic transition, following the end of the Late Paleozoic glaciation, and the resulting responses of the biota to the changing environment, are considered important analogs for understanding rapid future warming scenarios. While there has been extensive research on the patterns and extent of diversity in plants, tetrapods, and marine invertebrates during the Permo-Triassic, the study of insect diversity and the evolution of their faunal composition has been relatively limited. The question of whether there were insect extinctions during this period continues to be a subject of debate. Here, we present a statistical study on taxonomic diversity of insects—at specific, generic and familial levels—throughout the Permian and Triassic, with subsampled context on the reported global occurrences. Our result show that more than one insect extinction events, accompanied by significant diversity drop and turnovers of faunal compositional, occurred in the Permian and Triassic. All the uncovered insect diversity crises exhibit strong correspondence with the well-known marine mass extinction events in the Middle Permian, Permo-Triassic transition, Carnian, and Rhaetian, whilst the marine correspondence with the Early Permian insect crisis is less pronounced. Insects, being a major component of terrestrial ecosystems, demonstrate varied diversity responses to climatic changes in Permian and Triassic. Our study sheds new light on the intricate interplay between insect diversity evolution and the changing environmental conditions during these critical geohistorical periods.

Latest Permian insects from Wapadsberg Pass, southern Karoo Basin, South Africa

AbstractThe Permian–Triassic global crisis was the only event to have a dramatic impact on insect diversity, with Palaeozoic insect clades disappearing and accompanied by the accelerated rise of modern lineages. To date, most Permian palaeoentomological work has focused on the upper Permian deposits of the Normandien Formation (Beaufort Group), KwaZulu–Natal Province. This paper describes the regional insect fauna preserved close to the Permian–Triassic boundary at Wapadsberg Pass, southern Karoo Basin of South Africa, which provides a rare glimpse into insect life immediately before this global crisis. Here, we describe six insect species from six families within five orders from the two Wapadsberg Pass localities. Mioloptera stuckenbergi (Grylloblattodea) and Permocicada sp. (Hemiptera) were recorded previously from the Lopingian KwaZulu–Natal Province. In addition, we detail the first potential occurrence of South African Permian Tettigoniida (Orthoptera: Tettigonioidea & Hagloidea) and Anthracoptilidae (Paoliida). An Auchenorrhynchan (Hemiptera) nymph and an undetermined insect (order uncertain) also are described.

New Dictyoneuridans (Insecta: Dictyoneurida Handlirsch = Palaeodictyoptera Goldenberg) from the Permian Deposits of the Vorkuta Basin

Two new species of the family Spilapteridae, Homaloneura brauckmanni sp. nov. and Baeoneura desperata sp. nov. are described from the Lower Permian insect bearing deposits of the Vorkuta Coal Basin. For Vorkutoneura variabilis Sinitshenkova, 1977 (Spilapteridae), the imaginal forewing and nymphal hindwings are described for the first time. The dictyoneuridan assemblage does not agree with the existing dating of these deposits as Late Permian, as the insect assemblage suggests an Early Permian age.

The engineering of the giant dragonflies of the Permian: revised body mass, power, air supply, thermoregulation and the role of air density.

An engineering examination of allometric and analogical data on the flight of giant Permian insects (Protodonata, Meganeura or griffinflies) indicates that previous estimates of the body mass of these insects are too low and that the largest of these insects (wingspan of 70 cm or more) would have had a mass of 100-150 g, several times greater than previously thought. Here, the power needed to generate lift and fly at the speeds typical of modern large dragonflies is examined together with the metabolic rate and subsequent heat generated by the thoracic muscles. This evaluation agrees with previous work suggesting that the larger specimens would rapidly overheat in the high ambient temperatures assumed in the Permian. Various extant mechanisms of thermoregulation are modelled and quantified, including behaviour, radiation and the constraints on convective respiration and evaporation imposed by air flow through spiracles. However, the effects of these on cooling an overheated insect are found to be limited. Instead, an examination of the heat budget in the flight medium indicates that, at about 1.6bar (160kPa), thermoregulation supply enters into equilibrium and, even at high ambient temperatures, overheating can be avoided and enough oxygen sourced. This approach indicates how fossil biology can be used to examine past atmospheres.

Evolution of a complex behavior: the origin and initial diversification of foliar galling by Permian insects.

A central notion of the early evolution of insect galling is that this unique behavior was uncommon to rare before the diversification of angiosperms 135 to 125m.yr. ago. However, evidence accumulated during recent years shows that foliar galls were diverse and locally abundant as early as the Permian Period, 299 to 252m.yr. ago. In particular, a diversity of leaf galling during the Early Permian has recently been documented by the plant-damage record of foliar galls and, now, our interpretation of the body-fossil record of culprit insect gallers. Small size is a prerequisite for gallers. Wing-length measurements of Permian insects indicate that several small-bodied hemipteroid lineages originated early during the Permian, some descendant lineages of which gall the leaves of seed plants to the present day. The earliest foliar gallers likely were Protopsyllidiidae (Hemiptera) and Lophioneuridae (Thripida). Much of the Early Permian was a xeric interval, and modern galls are most common in dry, extra-tropical habitats such as scrubland and deserts. Plant-damage, insect body fossils, and the paleoclimate record collectively support the ecological expansion of foliar galling during the Early Permian and its continued expansion through the Late Permian.

New Middle Permian insects from Salagou Formation of the Lodève Basin in southern France (Insecta: Pterygota)

SummaryNew insects are described from the Middle Permian (Guadalupian) of Salagou Formation in the Lodève Basin (Languedoc region, France). Palaeopterans are represented by new odonatopterans viz. Huangiopterum lodevense n. gen., n. sp. and Huangiopterum species B, attributed to clade Panodonata (Huangiopteridae n. fam.), already recorded through the family Saxonagrionidae in the same formation, and a diaphanopterodean Martynovia cf. halli Beckemeyer 2004 (Martynoviidae) known from the Lower Permian of Oklahoma. Archaeorthopterans are supplemented by Bethouxia ariegensis n. gen., n. sp., Paralongzhua elongata n. gen., n. sp., Lodevolongzhua incompleta n. gen., n. sp., the caloneurodean Jarmiloptera mouralensis n. gen., n. sp., and a cnemidolestodean cf. Tococladus sp. (Tococladidae). Permophyllum rotundatum n. sp. (?Tettigoniidae) demonstrates the first presence of Orthoptera with wings probably mimicking Peltaspermales reproductive organs in Late Palaeozoic possibly analogous to the known crypsis of roachoid tegmina with pteridosperm pinnules. Frequent occurrence of hardly identifiable grylloblattid wing fragments support their significant role in term of abundance and diversity of the Permian faunas. New taxa based on wing venation pattern as Genera and species undetermined A, B (Liomopteridae), Permobaharellus salagousensis n. gen., n. sp. (family uncertain), Lodevophlebia reticulata n. gen., n. sp. (Sylvaphlebiidae), Oborella lodevensis n. sp. (Euryptilonidae), and Genus and species undetermined C, were described. Paoliida is represented by a paoliid gen. et sp. indet. and three anthracoptilids Strephocladus gandi n. sp., Strephocladus mouralensis n. sp., Lodevocladus subtilis n. gen., n. sp. The first record of miomopteran Palaeomantis sp. (Paleomanteidae) from Salagou Formation is presented. Acercarian insects are represented by the first record of Thripida and diverse new hemipterans, namely Euroscytina lutevanorum n. gen., n. sp. (Cicadomorpha: Scytinopteridae), Prosbole dio n. sp., Prosbole garrici n. sp., Pervestigia cf. veteris Becker-Migdisova 1961 (Prosbolidae), Archescytina species A (Paleorrhyncha: Archescytinidae) and Permopsyllidium lesclansis n. sp. (Sternorrhyncha: Psylloidea: Protopsyllidiidae). Entomofauna from the Salagou Formation demonstrate highly diverse insect communities with some taxa linked to early and late Permian ones known from assemblages in Euramerica and Russia, such as Obora (Czech Republic), Wellington Formation (Kansas and Oklahoma, USA), and Tshekarda (Ural, Russia).

Plant–arthropod interactions in gymnosperm leaves from the Early Permian of Patagonia, Argentina

Evidence of plant–arthropod interactions in three orders of gymnosperms is documented in at least five localities from Lower Permian strata of the Río Genoa Formation (Patagonia, Argentina). Arthropod damages were identified in Glossopteris, Gangamopteris, Cordaites and Gingkoites leaves, corresponding to four interactive categories: margin feeding, hole feeding, surface feeding, and oviposition. In addition, a Permian insect trace not previously recorded is described on Glossopteris wilsonii leaves. The diversity of arthropod damage is consistent with the diverse flora of the Río Genoa Formation. In turn, the patterns of arthropod and plant–arthropod interaction provide constraints on the ecology and paleoclimate of Western Gondwana during the Early Permian.

Fossil insects of the middle and upper Permian of European Russia

Fossil insects of European Russia from the Urzhumian to Vyatkian stages are reviewed, new taxa are described, and dynamics of insect taxonomic diversity around the Permian-Triassic boundary in light of the Paleozoic-Mesozoic boundary global extinction problem is analyzed. Traces of interactions between arthropods and plants are analyzed. Insect-bearing deposits of the Late Paleozoic found in the northern and eastern areas of the East European Platform are unique on the global scale in their completeness and continuity, allowing us to trace especially comprehensively the biotic processes that occurred around the boundary described as the time of the greatest biotic catastrophe of the Phanerozoic. A total of 28 genera and 111 species are newly described. Within the range from the Urzhumian to the Permo-Triassic boundary, 15 representative successive assemblages, including 112 families, are recognized (seven in the area in question and eight in other regions of Asia, Australia, and Africa). New tools are developed for the analysis of the dynamics of diversity. These tools show an approximately equilibrium (slightly positive) dynamics in the Urzhumian and Severodvinian and a drop in diversity during the Vyatkian Age. It is shown that Permian insect assemblages acquired a substantially post-Paleozoic pattern much earlier than the end of the Paleozoic. The character of changes that took place in the Induan and Olenekian remains uncertain, but a large-scale extinction event did not occur here: most families that have not been recorded at the beginning of the Triassic are recorded again in the Middle and Upper Triassic. Nevertheless, a biotic crisis probably actually took place, but was reduced to reorganization of the biota’s structure, which provided enormous growth of biodiversity over subsequent hundreds of millions of years, rather than resulted in catastrophic extinction. This study is intended for entomologists, stratigraphers, and all readers interested in the biotic events that took place around the Permian-Triassic boundary.

New Middle Permian palaeopteran insects from Lodève Basin in southern France (Ephemeroptera, Diaphanopterodea, Megasecoptera)

Three new palaeopteran insects are described from the Middle Permian (Guadalupian) of Salagou Formation in the Lodève Basin (South of France), viz. the diaphanopterodean Alexrasnitsyniidae fam. n., based on Alexrasnitsynia permiana gen. et sp. n., the Parelmoidae Permelmoa magnifica gen. et sp. n., and Lodevohymen lapeyriei gen. et sp. n. (in Megasecoptera or Diaphanopterodea, family undetermined). In addition the first record of mayflies attributed to family Syntonopteridae (Ephemeroptera) is reported. These new fossils clearly demonstrate that the present knowledge of the Permian insects remains very incomplete. They also confirm that the Lodève entomofauna was highly diverse providing links to other Permian localities and also rather unique, with several families still not recorded in other contemporaneous outcrops.

On the history of Russian paleontology

The 130th anniversary of the birth of Andrei Vasil� ievich Martynov (1879–1938), one of the most active members of the Paleontological Society, renowned entomologist, founder of Russian entomology, profes� sor at the Universities of Warsaw and Rostov, member of staff of the Zoological Institute and the Geological Museum of the Academy of Sciences of the USSR, Head of the Entomology Department of the Paleonto� logical Institute. Andrei Vasilievich studied morphol� ogy and systematics of the Permian, Mesozoic, and extant insects, and of modern crustaceans. He amassed a uniquely rich collection of insects from the Permian deposits of various regions of the Russian Platform. He published his main conclusions in the monograph Sketches on the Geological History and Phylogeny of Insects. He also revised the section “Insecta” in the Russian edition of Zittel’s “Grundzuge der Palaontologie” He is the author of 172 publications, including 10 monographs (for more information, see Rodendorf and Ryabinin, 1945). The 120th anniversary of the birth of Aldona Fedor�

Permian fossil insects of North-East Europa: Revision of the family Atactophlebiidae (Ins. Gerarida, Atactophlebiidae)

Revision of family Atactophlebiidae is made. Olgaephilus emendatus gen. et sp. n. from Upper Permian of Arkhangelsk region is described. On the basis of wing-venation and structure of body family Atactophlebiidae must be transferred from order Grylloblattida (= Paraplecoptera) to the order Gerarida.

On Permian and Triassic insect faunas in relation to biogeography and the Permian-Triassic crisis

The taxonomic diversity dynamics of pterygote insects in the Permian and Triassic at the family/age level are considered. Different metrics of taxonomic diversity are compared. Biogeographic and taphonomic aspects of changes in the composition of insect faunas in the Permian and about the P-T transition are discussed. Some changes in the Permian insect faunas are of a biogeographic nature and do not indicate global changes in diversity. Insects with aquatic immatures were rather common in the Permian and Early Triassic, but these immatures are well represented in only few localities.

Pollen eaters and pollen morphology: co-evolution through the Permian and Mesozoic

In each of the sequential periods of seed-plant evolution, there were charismatic pollen grains that fascinated generations of palynologists. In the Permian, such peculiar pollen morphologies included taeniate pollen (with the wall split into bands, or taeniae) of Protohaploxypinus-Lunatisporites and Vittatina groups. One or more types of taeniate pollen prevailed in the pollen load of several species of Permian insects. The advent of the Mesozoic era was marked by a spread of smooth asaccate pollen of Ginkgocycadophytus type, virtually indistinguishable in the dominant orders of Mesozoic gymnosperms. Such pollen was never found in the guts. Charismatic through the Mesozoic was the rimulate pollen of Classopollis type. It constituted a monospecific load in the guts of large Jurassic katydids (Aboilus, Haglidae). The Eucommiidites group, related to gnetophytes, is peculiar in having lateral grooves parallel to the main sulcus. Recently this pollen type has been found in the gut compression of a xyelid, genus Ceroxyela. The reason why Palaeozoic and Mesozoic insects showed preferences for pollen with distinctive morphologies might be that, on the basis of the surface ornamentation of these different varieties, insects were able to distinguish edible pollen types. The distinctive pollen morphologies might have evolved primarily for this purpose.

The geological setting and palaeoenvironmental and palaeoecological reconstructions of the Upper Permian insect beds at Belmont, New South Wales, Australia

The geological setting and palaeoenvironmental and palaeoecological reconstructions of the Upper Permian insect beds at Belmont, New South Wales, Australia

<I>Permocoleus</I>, New Genus, the First Permian Beetle (Coleoptera) from North America

Permocoleus wellingtonensis, new genus and new species, is based on an elytron from the Permian Wellington Formation of Oklahoma and is the only Paleozoic record of the order Coleoptera from North America and the oldest record for the New World. Until now, Permian Coleoptera were known only from Europe, Australia, southern Africa, and South America but were conspicuously absent from North America, despite the Wellington Formation being among the most diverse deposits of Permian insects. Permocoleus provides evidence that early beetles were globally distributed.

The Permostridulidae fam. n. (Panorthoptera), a new enigmatic insect family from the Upper Permian of France

The unusual wing characters of the Permian insect Permostridulus brongniarti gen. n., sp. n. justifies the creation of a new family, the Permostridulidae fam. n., within the Panorthoptera. Phylogenetic relationships with the extinct order Caloneurodea, related to the Orthoptera (crickets, grasshoppers), are assumed. This assumption suggests an occurrence of the Permostridulidae at least since the Upper Carboniferous. The most prominent feature of the wing venation is a stridulatory apparatus, nonhomologous with those previously known in panorthopterid lineages. This is the oldest recorded sound-producing device of an animal.

The Permian Insect Fossils of Elmo, Kansas . By R oy J. B eckemeyer , 2000 (Emporia, KS: Emporia State University) The Kansas School Naturalist Vol. 46(1) [Pp. 15 with a reference section and numerous bw photographs and line drawings]. Price free (so long as copies last) but may involve a copying and postage charge. 0022-877X.

The Permian Insect Fossils of Elmo, Kansas . By R oy J. B eckemeyer , 2000 (Emporia, KS: Emporia State University) The Kansas School Naturalist Vol. 46(1) [Pp. 15 with a reference section and numerous bw photographs and line drawings]. Price free (so long as copies last) but may involve a copying and postage charge. 0022-877X.

Early Permian insect folivory on a gigantopterid-dominated riparian flora from north-central Texas

Quantitative analysis of 11,562 cm 2 of foliar surface area from an Early Permian flora from north-central Texas reveals that 2.6% was removed by insect folivores. Of the 1346 foliar items analyzed, 31.8% were attacked by insect folivores. This Permian insect herbivory approximated one-third the level in modern tropical to temperate forests. Three taxa of Gigantopteridaceae exhibited disproportionately high insect consumption levels, ranging from 3.1 to 4.4% of leaf area, and attack frequencies of 39.6 to 82.8% —levels comparable to taxon-specific values for modern woody dicotyledonous angiosperms. Qualitative analysis of folivory type also indicates that five of the eight categories of herbivore damage occurring on this flora were confined to or overwhelmingly represented on Gigantopteridaceae, suggesting preferential host specificity for Gigantopteridaceae by insect folivores. By contrast, other pteridophylls, conifers, cycadophytes, and two taxa of uncertain affinity, showed low levels of leaf area removal and attack frequencies. The Gigantopteridaceae, an enigmatic group of Permian plants with large, megaphyllous foliage, foreshadowed physiognomic attributes occurring in later angiosperms. Both groups had similar consumption patterns and intensities by insect folivores. In this moderately diverse Early Permian flora, insect folivory was significant, at least in one riparian environment outside the Euramerican coal-swamp ecosystem, indicating that modern modes and magnitudes of insect consumption of vascular plant tissue were in place during the Late Paleozoic.

Permian Pollen Eating

The Random Samples item “Permian pollen eaters” ([16 May, p. 1035][1]) provides a stimulating account of the discovery by Russian paleobiologists Alexander Rasnitsyn and Valentin Krassilov of identifiable pollen in the guts of Early Permian insects. Paleobotanist William Chaloner is quoted as saying that inadvertent consumption of pollen could explain the occurrence of this pollen. One of the preserved insects is a member of the Synomaloptilidae, ancestral to modern bark lice and a member of a Paleozoic insect clade that was postulated by Rasnitsyn to consume “plants by feeding primarily on their reproductive organs …” ( 1 , p. 27, translated from the Russian). In 1997, Rasnitsyn predicted that these and related taxa possessed head and mouthparts that consumed material nutritionally equivalent to pollen ( 2 , p. 65). It has been known that Late Carboniferous coal-ball permineralizations include fossilized fecal pellet assemblages containing histologically pristine plant tissues, some of which consist entirely of spores, pollen, and associated tissue ([3][2]). And insect consumption of wind-dispersed pollen has been demonstrated repeatedly in such consummate pollenivores as syrphid flies ([4][3]), bees ([5][4]), and other pollenivorous insects ([6][5]). The occurrence of pollen interpreted as wind-dispersed in the guts of Permian insects is thus unlikely to be accidental. A more parsimonious conclusion is that several lineages of Early Permian insects were actively consuming nutritionally rich pollen, regardless of the mode of dispersal. Such a diet was a necessary prelude to pollinator mutualisms between seed plants and insects that occurred subsequently in geologic time ([7][6]). 1. A. P. Rasnitsyn, Tr. Paleontol. Inst. 174 , 1 (1980). 2. \___|, Paleontol. J. 11 , 60 (1977). 3. [↵][7]R. W. Baxendale, Palaeontology 22 , 537 (1979); S. V. Meyen, Z. Geol. Wiss. 12 , 269 (1984). 4. [↵][8]P. Stelleman and A. D. J. Meeuse, Tijd. Entomol. 119 , 15 (1976). 5. [↵][9]J. Pojar, Am. Midl. Natl. 89 , 448 (1973). 6. [↵][10]O. Hagerup, Dan. Vidensk. Sels. Biol. Meddel. 18 (no. 4), 1 (1950). 7. [↵][11]C. C. Labandeira, Ann. Rev. Ecol. Syst. , in press. [1]: /lookup/doi/10.1126/science.276.5315.1035a [2]: #ref-3 [3]: #ref-4 [4]: #ref-5 [5]: #ref-6 [6]: #ref-7 [7]: #xref-ref-3-1 View reference 3 in text [8]: #xref-ref-4-1 View reference 4 in text [9]: #xref-ref-5-1 View reference 5 in text [10]: #xref-ref-6-1 View reference 6 in text [11]: #xref-ref-7-1 View reference 7 in text

Pollen in the guts of Permian insects: first evidence of pollinivory and its evolutionary significance

Uniquely preserved pollen was extracted from intestines of fossil insects from the Lower Permian of the Urals. A species of Hypoperlidae, an extinct family ancestral to bark-lice, bugs and plant-hoppers, and two species of Grylloblatida, a predominantly Permian group with a few extant representatives related to stoneflies, contain protosaccate taeniate grains of several pollen genera well known as dispersed microfossils and occasionally found in sporangia of conifers, peltasperms and glossopterids. This is so far the earliest direct evidence of pollinivory, a major factor of plant-insect coevolution. The partly digested pollen grains reveal infratectal reticulum and other structural details of evolutionary significance. It is suggested that the peculiar taeniate pollen of worldwide distribution in the Permian might simultaneously evolve in several groups of Paleozoic seed plants in relation to pollinivory that, by altering the rnicropyle load and thereby the pollen/ovule ratio, could also affect ovuliferous structures. Thus pollinivory might impel rapid diversification of gymnosperms in the Permian. The pollinivorous Hypoperlidae, which have evolved in the direction of ovulivory, might initiate insect pollination in the process.