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Abstract and Figures
The stratigraphic distribution of blattoid (cockroach-like) insect genera in the type Pennsylvanian permits the zoning of lacustrine members in this sequence. When taken with the associated floras and the intercalated marine faunas, blattoid fossils permit fine correlation between the name-type Pennsylvanian of Pennsylvania, the goniatite-bearing Upper Carboniferous of Europe, the fusuline-bearing Pennsylvanian of Kansas, and other contemporaneous sequences, especially those of exclusively limnic facies such as the Morien, Pictou, and Narragansett Basins. Application of epochal ages, developed in Europe, to the type and other regions of the Pennsylvanian is now possible. The time distribution of the major genera is presented with a time-stratigraphic plot of data control points in the type region, and a documentation of the relevant systematics is appended.
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... Similar large post-Paleozoic forewings are only known from the Cretaceous of Mongolia, showing about five centimeters of forewing length [16] (Figure 4), and the modern Megaloblatta having wingspans up to 20 cm and corresponding forewing lengths of nearly 9 cm [17]. We focus on one of these large-winged genera, the Progonoblattina, previously considered as belonging to Archoblattinidae Schneider, 1983a [5], but now regarded as a member of the family Gyroblattidae Durden, 1969 [18]. Based on new finds and the revision of type specimens from the late 19th and early 20th centuries, some of the latter are figured for the first time as photographs and associated drawings. ...
... Order Blattodea Brunner von Wattenwyll, 1882 [29]. Durden, 1969 [18] Synonymy 1969 Phoberoblattidae-Durden, p. 1171 [18], syn. nov. ...
... Recently, the difference between Archoblattina, here considered as a new junior synonym of Progonoblattina, and Necymylacris were discussed [32] (p. 264-265), with some modifications made here: Progonoblattina is very similar in size and venation pattern to Necymylacris, the single genus of the family Necymylacridae Durden, 1969. Both genera differ mainly in the form of the CuA, which runs inclined and straight against the transition of the wing tip into the posterior wing border in Necymylacris (Figure 3) [5] (pl. ...
Large-winged blattoids of the Middle to Late Pennsylvanian reveal a striking appearance, diversification, and decline in the fossil record. Among them, the families Necymylacridae Durden, 1969, and Gyroblattidae Durden, 1969, as well as the mylacrid genus Opsiomylacris exhibit, the largest pre-Cenozoic blattoids with forewing lengths up to 7.5 cm. As finds from coal-bearing sedimentary basins in Europe, North Africa, and North America indicate, these giant insects started to spread around the Bashkirian–Moscovian transition and experienced a diversification in late Moscovian and Kasimovian times, until they disappeared in the middle Gzhelian. Whereas necymylacrids are only patchily reported and still lack distributional patterns, we disclose the occurrence and particular habitat preference of gyroblattids. Although appearing first in some vast North American basins, they became successively widespread only in small-sized basins of the European Variscan interior. Frequently found associated with enigmatic gymnosperms, they may have lived in well-drained hinterland areas from where they immigrated into the ever-wet basin centers only with increasing seasonality. Gyroblattids apparently followed meso- to xerophilous plants and likely colonized spaces offering a broader spectrum of edaphic conditions that resulted from the closeness of erosional and depositional areas. The presented analysis and revision of all gyroblattids aim to facilitate future more realistic biodiversity estimations based on fossil taxa.
... Handlirsch (1920) originally attributed Sooblattella to Archimylacridae, but Laurentiaux (1958) revised it and described a second species, Sooblattella ater (nomen nudum), from Kaiping. He indicated that Sooblattella is very similar to Etomylacris, a genus synonymized with Mylacris Scudder, 1868 in the Paleobiology Database and the GBIF database, after Durden's (1969Durden's ( : 1168 list of species in Mylacris even though this last author did not discuss the synonymization of the two genera. ...
The late Carboniferous (Pennsylvanian) witnessed the intensification of climatic gradients with the onset of the Late Paleozoic Ice Age. During this time, high latitudes were characterized by the spread of ice sheets, whereas the palaeotropics hosted the emergence of vast, verdant, coal-forming, swamp forests. Not only were the planet’s landscapes transformed through proliferation and diversification of vascular plants, but the evolution of diverse insect groups, including ‘primitive’ roachoids, reconfigured the terrestrial fauna. Here, we re-examine and describe new roachoid taxa from Moscovian strata of the Benxi Formation. The studied insect material was sourced from early 20th century collections held in the Swedish Museum of Natural History and the Royal Belgian Institute of Natural Sciences, and is co-preserved with a broad array of plant fossils—some with evidence of herbivory damage. Our findings introduce Tangshanblatta inexpectata gen. et sp. nov. (Phyloblattidae), Pseudaphtoromylacris paucinervis gen. et sp. nov. (in the family Pseudaphtoromylacridae, fam. nov.), Spilarchimylacris kaipingense sp. nov. (Archimylacridae), and Etomylacris straeleni sp. nov. and Sooblattella ater sp. nov. (both Mylacridae). We establish Pseudaphtoromylacridae on the basis of several new characters that clearly differentiate it from related families. Etomylacris is restored from synonymy with Mylacris. The co-preserved plant fossils reveal a rich palaeovegetation including pteridophytes, ‘pteridosperms’, sphenophytes, conifers, and arborescent lycophytes. The remains of these plants bear seven distinct damage types belonging to four functional feeding groups: hole feeding, margin feeding, piercing and sucking, and oviposition. This diversity of insects, plants, and herbivory strategies signifies a relatively complex food web in the Carboniferous tropical coal-forming forests of northeast China. This study also demonstrates the value of historical museum collections and the importance of inter-institutional collaboration for piecing together palaeobiodiversity and palaeoecological information from now inaccessible fossil sites.
... The probability of the calibrated node age being outside the defined range was set to 2.5%. The maximal root age of the tree was set to 323.2 million years in the MCMCTREE configuration file, which corresponds to the age of Mylacris anthracophila, the first roachoid fossil and stem Dictyoptera (Durden, 1969). Fossil ages were retrieved from the Paleobiology Database (https://paleobiodb.org/ accessed October 2023). ...
Termites are a lineage of social cockroaches abundant in tropical ecosystems where they are key decomposers of organic matter from wood to soil. Despite their ecological significance, only a handful of reference-quality termite genomes have been sequenced, which is insufficient to unravel the genetic mechanisms that have contributed to their ecological success. Here, we performed sequencing and hybrid assembly of 45 taxonomically and ecologically diverse termites and two cockroaches, resulting in haplotype-merged genome assemblies of 47 species, 22 of which were near-chromosome level. Next, we examined the link between termite dietary evolution and major genomic events. We found that Termitidae, which include ~80% of described termite species, have larger genomes with more genes and a higher proportion of transposons than other termites. Our analyses identified a gene number expansion early in the evolution of Termitidae, including an expansion of the repertoire of CAZymes, the genes involved in lignocellulose degradation. Notably, this expansion of genomes and gene repertoires coincided with the origin of soil-feeding in Termitidae and remained unchanged in lineages that secondarily reverted to a wood-based diet. Overall, our sequencing effort multiplied the number of available termite genomes by six and provided unprecedented insights into the genome evolution of the most ancient lineage of social insects.
... To evaluate the phylogenetic signal recoverable from forewing venational similarity and the associated confidence level, this study reconstructed phylogenies of the genera of well-recognized family-group taxa using only forewing characters. The results have implications for insect paleontological studies as well as geological studies because forewing fossils have been used for stratigraphic correlations (e.g., [27][28][29][30]). ...
A reliable character system is crucial to taxonomy and systematics, and it promises valid downstream inferences, e.g., estimates of diversity and disparity, reconstruction of evolutionary history, and even stratigraphic correlations. Modern taxonomy and systematics of extant cockroaches requires an integrative study involving multiple lines of evidence with emphasis on genital and reproductive characteristics and molecular data. In contrast, many fossil cockroach taxa published recently are based solely on forewings. Many studies have shown that forewing-based taxa are questionable. In order to find out how much of the phylogenetic signal we could ascertain from venational similarity, and how confident we could be, this study used forewing characters to reconstruct phylogenies of the genera of well-recognized family-group taxa. The intuitively reconstructed phylogeny of 75 extant genera failed to recover those taxa or their relationships. Parsimony analyses of various datasets all yielded strong polyphyly and chaotic relationships. In conclusion, the forewing of cockroaches is not a universally competent character system. The underlying causes are the complicated nature of veins and the limitations of current analytical techniques. The uncertainty in forewing-based taxonomy and systematics has been underestimated in the literature. Forewing-based fossil taxa warrant re-evaluation; some of them are herein deemed nomina dubia in their current state.
... Major strides have been made in understanding the spatial and temporal responses of plants to glacialinterglacial cycles during the Pennsylvanian, but the effects of short-term climate change on Pennsylvanian terrestrial arthropods is not well understood. Most insect fossils described from the Pennsylvanian have been collected above or below coal beds (Durden 1969;Schneider and Werneburg 2006) and often found incidentally during palaeobotanical fieldwork. Insect-bearing beds from important Middle and Late Pennsylvanian Lagerstätten, including Mazon Creek, USA (Clements et al. 2018), Piesberg, Germany (Leipner et al. 2021), Montceau-les-Mines, France (Perrier and Charbonnier 2014), and Commentry, France (Carpenter 1961) are associated with coals. ...
The Middle-Late Pennsylvanian Subperiod was marked by recurrent glacial-interglacial cycles superimposed on a longer-term trend of increasing aridity. Wetland and drought-tolerant floras responded with repeated migrations in the tropics, and a major plant turnover occurred in swamp ecosystems in parts of Euramerica near the Middle-Late Pennsylvanian boundary. However, the corresponding ecological and evolutionary responses of insects and other terrestrial arthropods are poorly understood. Here, we review the record of plant-arthropod interactions and analyse origination and extinction rates of insects during the Middle-Late Pennsylvanian. Although preliminary, plant-arthropod associations broadly persist through the Middle-Late Pennsylvanian boundary, and new damage types and host-plant associations first appear in the Late Pennsylvanian, possibly related to increased availability of accessible vascular and foliar tissues associated the shift from arborescent lycopsid to tree and seed fern dominance in Euramerican wetlands. Likewise, our analysis of the insect body fossil record does not suggest especially high rates of origination or extinction during this interval. Together, these results suggest that insects did not suffer major extinctions during the Middle-Late Pennsylvanian, despite short- and long-term changes in climate and environmental conditions.
Supplementary material at https://doi.org/10.6084/m9.figshare.c.6280586
... One year later, Handlirsch (1920) also named a genus †Boltonia, for a stem-dictyoptera "roachoid", designating †Gerablattina (Aphthoroblattina) sulcata Bolton 1911 as its type species. Currently, †Boltonia Handlirsch is a junior synonym of †Necymylacris Bolton 1917 (Blattodea) (Durden, 1969). Both Pruvost (1919) and Handlirsch (1920) named their genera after the paleoentomologist Herbert Bolton. ...
... At that time, his interregional correlations were already very convincing and close to modern ones. A later attempt by Durden (1969) to use North American and European blattoid insects for biostratigraphy failed because of a completely inadequate classification and oversplit taxonomy. The present insect-zone biostratigraphy of the Late Pennsylvanian and the early Permian (Schneider et al. 2020), based on the wings of blattoids, started with Schneider (1982). ...
For the biostratigraphy of mixed continental-marine and purely continental sections in the palaeotropical belt of Euramerica, 9 insect and 8 conchostracan zones are newly defined or improved. These zones encompass the time interval from the Early Pennsylvanian (middle Bashkirian) up into the early Permian (early Asselian) of the Euramercian biotic province. They are linked as much as possible to the marine Standard Global Chronostrigraphic Scale by common occurrences of insects and/or conchostracans with conodonts in mixed marine-continental sections as well as by the thus far available and reliable radioisotopic ages of associated volcanic rocks. This insect and conchostracan zonation is an alternative tool to the well-established macro-plant biostratigraphy of the Pennsylvanian. In contrast to the latter, only single specimens of insects or conchostracans, even if more rare than plant remains, allow biostratigraphic dating with a similar high temporal resolution.
... The present insect-zone biostratigraphy of the Late Pennsylvanian and the early Permian based on the wings of blattoid insects started with Schneider (1982). The earlier Pennsylvanian blattoid insect biostratigraphy of Durden (1969) has been shown to be inapplicable because of a completely inadequate classification and oversplit taxonomy. ...
Nonmarine biostratigraphic/biochronologic schemes have been created for all or parts of the late Carboniferous–Middle Triassic using palynomorphs, megafossil plants, conchostracans, blattoid insects, tetrapod footprints and tetrapod body fossils, and these provide varied temporal resolution. Cross correlation of the nonmarine biochronologies to the Standard Global Chronostratigraphic Scale has been achieved in some parts of the late Carboniferous–Middle Triassic in locations where nonmarine and marine strata are intercalated, the nonmarine strata produce biochronologically significant fossils and the marine strata yield fusulinids, conodonts and/or ammonoids. Other cross correlations have been aided by magnetostratigraphy, chemostratigraphy and a growing database of radioisotopic ages. A synthetic nonmarine biochronology for the late Carboniferous–Middle Triassic based on all available nonmarine index fossils, integrated with the Standard Global Chronostratigraphic Scale, is presented here. The focus is on the nonmarine biostratigraphy/biochronology of blattoid insects, conchostracans, branchiosaurid amphibians, tetrapod footprints and tetrapod body fossils within the biochronological framework of land-vertebrate faunachrons. Correlation to the Standard Global Chronostratigraphic Scale presented here is divided into seven time intervals: Pennsylvanian, Carboniferous–Permian boundary, Cisuralian, Guadalupian, Lopingian, Permian–Triassic boundary and Early to Middle Triassic. The insects, conchostracans and branchiosaurs provide robust nonmarine correlations in the Pennsylvanian–Cisuralian, and the footprints and tetrapod body fossils provide robust correlations of varied precision within the entire Pennsylvanian–Middle Triassic. Radioisotopic ages are currently the strongest basis for cross correlation of the nonmarine biostratigraphy/biochronology to the Standard Global Chronostratigraphic Scale, particularly for the Pennsylvanian–Cisuralian. Chemostratigraphy and magnetostratigraphy thus far provide only limited links of nomarine and marine chronologies. Improvements in the nonmarine-marine correlations of late Paleozoic–Triassic Pangea require better alpha taxonomy and stratigraphic precision for the nonmarine fossil record integrated with more reliable radioisotopic ages and more extensive chemostratigraphic and magnetostratigraphic datasets.
Here we describe the first discoveries of insects from the Gzhelian of the Reyran Basin (northern part of Esterel, South of France). These are a new mylacrid, Opsiomylacris sp. an undetermined genus and species of Phyloblattidae, two other roachoids of uncertain affinities, and a Palaeodictyoptera, all based on forewings. Although these discoveries demonstrate that insects were present in the corresponding paleoforest, they remain less diverse and rarer than the plant remains. The genus Opsiomylacris is currently supposed to have lived under xeric environments, possibly corresponding to the more xeric paleobiotas of the margin of the humid valley bottoms.
We introduce a novel paoliid, Carbonidelia gallica gen. et sp. nov., which may share close affinities with the ‘ideliid’ genus Sojanidelia Storozhenko, 1992. This new taxon is described from Gzhelian strata exposed at Tante Victoire in Var, France. Additionally, we present a remarkably well-preserved forewing from the same outcrop belonging to the phyloblattid genus and species Anthracoblattina ensifer cf. ‘variety desguini’. This species was previously known from the Gzhelian of Commentry in Allier, France. Therefore, this finding strengthens the Gzhelian age derived from the study of the palaeoflora at this outcrop. In this study, we also briefly delve into the inter- and intraspecific variability in the forewing venation of the ‘Anthracoblattina gigantea-ensifera group’. Furthermore, we challenge the classification of two Mesozoic roachoids, Sobytie tungusicum Vrsansky, 2010 and Pozabudnutie antiquorum Hinkelman and Vrsansky, 2022, which, in our view, do not belong to the Phyloblattidae family. Instead, they should be regarded as Dictyoptera of uncertain family affinities. A comprehensive revision of these species will be necessary to establish their precise taxonomic positions.
I am indebted to the labours of many workers for the opportunity of preparing this paper upon the remains of fossil insects discovered in the Welsh Coal Measures during the last fifteen years.
Some of the specimens are in the National Museum of Wales, having been discovered in the collection of Coal-Measure fossils given to that Institution by Mr. David Davies, of Gilfach Goch.
Dr. A. E. Trueman and Miss E. Dix, M.Sc., have lent insect-remains from the Coal Measures of the Swansea area.
To Dr. F. J. North I am much indebted, not only for the opportunity of retaining specimens for examination over a long period of time, but for information respecting horizons at which the insect-remains occur.
To the Geological Survey and to Mr. John Pringle I am under obligations for the loan of a blattoid wing found by him in shales associated with the Mynyddislwyn Vein at Maes-y-Cwmmer (Glamorgan), and for an Archimylacrid pronotum from the Gwernau Level, three-quarters of a mile south-east of Maes-y-cwmmer, at the same horizon.
The series of wings represents nearly a score of individuals, and constitutes the largest group of Coal-Measure insects yet recorded at one time from any British coalfield. They are, with two exceptions, referable to the Blattoid families Archimylacridsæ, Handlirsch, and the transitional ‘Tribe’ named Heraimylacridia, Pruvost. Only two of the specimens are referable to the most archaic group of fossil insects, the Palæodictyoptera.
As we shall see later, the Hemimylacridian forms now known to occur in the South
Studies on the phylogeny of the Blattiform Orthoptera. Unpublished Ph.D. dis-sertation
- C J Durden
DURDEN, C. J. 1968. Studies on the phylogeny of the Blattiform Orthoptera. Unpublished Ph.D. dis-sertation, Yale Univ., New Haven, Conn., pp. 1-626.
Neue Insektenfunde aus dem Saarbriicker Steinkohlengebirge und die Bedeu-tung der Dictyoneuriden fiir die Stratigraphie des Saarkarbons
- P Guthorl
GUTHORL, P. 1951. Neue Insektenfunde aus dem Saarbriicker Steinkohlengebirge und die Bedeu-tung der Dictyoneuriden fiir die Stratigraphie des Saarkarbons. Bergbau-Archiv, 15, pp. 99-107.
FULGORlNA Goldenberg, 1869 (= Megablattina Brongniart, 1885, nec Sellards, 1903), with closely related genera in the Clyde Formation, Wichita Group of Texas, and in the Alikaevsk Member, Nidnebalakonsk Formation of the Kuznetsk Basin
- Family
- Fulgorinidae
Family FULGORINIDAE nov. (9) FULGORlNA Goldenberg, 1869 (= Megablattina Brongniart, 1885, nec Sellards, 1903), with closely related genera in the Clyde Formation, Wichita Group of Texas, and in the Alikaevsk Member, Nidnebalakonsk Formation of the Kuznetsk Basin, U.S.S.R. F. klieveri Goldenberg, 1869: 166, type species, Got-telborner Schichten, Ottweiler Gruppe, (Stephanian A), of Michelsberg b. Wemmetsweiler, Saar, Germany. Fulgorina sp. a, assemblage 8b.
1885:36, type species (= S. perforata Handlirsch, 1906) and the nominal species S. triassica, guttata, marginata Scudder, 1885, and S. fenestrata Handlirsch, 1906, from the Pony Springs Member, Maroon Formation
- S Scudder
S. gardineri Scudder, 1885:36, type species (= S.
perforata Handlirsch, 1906) and the nominal
species S. triassica, guttata, marginata Scudder,
1885, and S. fenestrata Handlirsch, 1906, from the
Pony Springs Member, Maroon Formation, 3f
miles (5.6 km) southeast of Fairplay, Park County,
Colorado.
1851:86) and S. dohrnii (Scudder, 1879:66), from the Stephanian C of Wettin
- S Germar
S. euglyptica (Germar, 1851:86) and S. dohrnii
(Scudder, 1879:66), from the Stephanian C of
Wettin, Saale, Germany.
Pedee Group, of Havercamp Farm
- Iaxa Sellards
- S Sellardsi
- Lawrenciana Handlirsch
S. maledicta, arcuata, curvata, Iaxa Sellards, 1908,
and S. sellardsi and lawrenciana Handlirsch, 1906,
from the Weston Shale, Pedee Group, of Havercamp Farm, (Lonestar Lake Damsite), 3 miles
(4.8 km) southeast of Lawrence, Douglas County,
Kansas.
Spiloblattina expansa Bolton, 1925, (= Etoblattina
voisine de E. balteata, Brongniart, 1893), from the
Stephanian A of Commentry, Allier, France.