Abstract
Mutovia intercalaria gen. et sp. n. is described from the Upper Permian sediments of Isady in north-western Russia and placed withina new Palaeozoic cockroach family, Mutoviidae fam. n.. The forewing, with very basal, distinct separation of R1 and RS resemblingthe cockroach hind wing and also the forewing of mantises and eoblattids, and with a sharply convex clavus, is unique. The generalvein scheme is closely analogous to Cretaceous mantises (Baissomantis Gratshev et Zherikhin, 1994), but the two taxa are not directlyrelated. The new species comprises about 3% of all fossil insects and 10% of fossil cockroaches at the site, and thus is one of the dom-inant species at the site. Reorganization of the main veins within the taxon suggests the group was rapidly evolving, perhaps becauseof environmental stress. The high coefficient of variability for total number of veins meeting margin (14.5%) is characteristic fornewly emerging (early Palaeozoic and early Mesozoic) lineages and is much higher than for the same characteristic in advanced Pale-ozoic taxa. In spite of an apparently aerodynamic shape of the wing, it likely was not a good flier, as indicated by the extreme variationof veins in the anterior margin and narrow and plastic hind wing remigium. A sex ratio approaching 2:1 suggests that males were moreactive in flight. The new family represents a direct evidence for appearance of new taxa before the Permotriassic boundary and disability of these taxa to cross that crisis.References
Aristov, D.S. (2004a) Grylloblattids of the family Chaulioditidae (= Tomiidae syn. nov.) (Insecta: Grylloblattida) from the Upper Permian of the Orenburg Region. Paleontological Journal, 38 (Suppl. 2), 146–149.
Aristov, D.S. (2004b) Grylloblattids of the family Liomopteridae (Insecta: Grylloblattida) of the Upper Half of the Upper Permian. Paleontological Journal, 38 (Supplement 2), 150–157.
Aristov, D.S. (2008) New Tatarian Chaulioditinae (Insecta; Grylloblattida; Chaulioditidae) from European Russia. Paleontological Journal, 1, 33–36. (In Russian, translation in Paleontological Journal, 1, 32–35.)
Aristov, D.S. (2009) New Permian grylloblattids (Insecta; Grylloblattida) from Isady locality (Severodvinian Stage of Vologda Region). Russian Entomological Journal, 18 (1), 1–6.
Béthoux, O., Klass, K.-D. & Schneider, J.W. (2009) Tackling the “Protoblattoidea problem”: Revision of Protoblattinopsis stubblefieldi (Dictyoptera; Late Carboniferous). European Journal of Entomology, 106, 145–152.
Béthoux, O. & Wieland, F. (2009) Evidence for Carboniferous origin of the order Mantodea (Insecta: Dictyoptera) gained from forewing morphology. Zoological Journal of the Linnean Society, 156, 79–113.
Golubev, V.K. (2000) Permian and Triassic Chroniosuches and biostratigraphy of the Upper Tatarian sediments of Eastern Europe based on tetrapods. Nauka, Moscow, 174 pp. (in Russian)
Gratshev, V.G. & Zherikhin, V.V. (1993) New fossil mantids (Insecta, Mantida). Paleontological Journal, 27(1A), 148–165.
Grimaldi, D. (2003) A revision of Cretaceous mantises and their relationships, including new taxa (Insecta: Dictyoptera: Mantodea). American Museum Novitates, 3412, 1–47.
Jarzembowski, E.A. & Schneider, J.W. (2007) The stratigraphical potential of blattodean insects from the late Carboniferous of southern Britain. Geological Magazine 144 (3), 449–456.
Liang, J-H., Vršanský, P., Ren, D. & Shih, C. (2009) A new Jurassic carnivorous cockroach (Insecta, Blattaria, Raphidiomimidae) from the Inner Mongolia in China. Zootaxa 1974, 17–30.
Latreille, P.A. (1810) Considérations générales sur l’ordre naturel des animaux composant les classes des Crustacés, des Arachnides et des Insectes avec un tableau méthodique de leurs genres disposés en familles. Schoell, Paris, 444 pp.
Menning, M., Alekseev, A.S., Chuvashov, B.I., Davydov, V.I., Devuyst, F.-X., Forke, H.C., Grunt, T.A., Hance, L., Heckel, P.H., Izokh, N.G., Jin, Y.-G., Jones, P.J., Kotlyar, G.V., Kozur, H.W., Nemyrowska, T.I., Schneider, J.W., Wang, X.-D., Weddige, K., Weyer, D. & Work, D.M. (2006) Global time scale and regional stratigraphic reference scales of Central and West Europe, East Europe, Tethys, South China, and North America as used in the Devonian–Carboniferous–Permian Correlation Chart 2003 (DCP 2003), Palaeogeography, Palaeoclimatology, Palaeoecology, 240, 318–372.
Rasnitsyn, A.P. & Quicke, D.L.J. (Eds) (2002) History of insects. Kluwer Academic Publishers, Dordrecht, The Netherlands, 517 pp.
Shcherbakov, D.E. (2008) On Permian and Triassic insect faunas in relation to biogeography and the Permian–Triassic Crisis. Paleontological Journal, 42(1), 15–31.
Schneider, J. (1977) Zur Variabilität der Flügel paläozoischer Blattodea (Insecta), Teil I. Freiberger Forschungshefte, C 326, 87–105.
Schneider, J. (1978) Zur Variabilität der Flügel paläozoischer Blattodea (Insecta), Teil II. Freiberger Forschungshefte, C 326, 21–39.
Schneider, J. (1980) Zur Entomofauna des Jungpaläozoikums der Boskovicer Furche (ČSSR), Teil I: Mylacridae (Insecta, Blattodea). Freiberger Forschungshefte, C 357, 43–55.
Schneider, J. (1983) Die Blattodea (Insecta) des Paläozoikums, Teil 1: Systematik, Ökologie und Biostratigraphie. Freiberger Forschungshefte, C 382, 106–145.
Schneider, J. (1984) Die Blattodea (Insecta) des Paläozoikums, Teil II: Morphogenese der Flügelstrukturen und Phylogenie. Freiberger Forschungshefte, C 391, 5–34.
Schneider, J, Goretzki, J. & Rössler, R. (2005) Biostratigraphisch relevante nicht-marine Tiergruppen im Karbon der variscischen Vorsenke und der Innensenken. Courier Forschungsinstitut Senckenberg, 254, 103–118.
Schneider, J. & Werneburg, R. (2006) Insect biostratigraphy of the Euramerican continental Late Pennsylvanian and Early Permian. In: Lucas S.G., Cassinis G. & Schneider J.W. (Eds), Non-Marine Permian biostratigraphy and biochronology 265. Geological Society of London, Special Publications, London, pp. 325–336.
Sinitchenkova, N.D. & Aristov, D.S. (2010) New Permian stoneflies of the family Palaeonemouridae (Insecta: Perlida = Plecoptera) from the Isady locality. Paleontological Journal, 44, 49–52.
Storozhenko S.Y. (1997) Classification of order Grylloblattida (Insecta) with description of new taxa. Far Eastern Entomologist, 42, 1–20.
Svenson, G.J. & Whiting, M.F. (2009) Reconstructing the origins of praying mantises (Dictyoptera, Mantodea): The roles of Gondwanan vicariance and morphological convergence. Cladistics, 25, 1–47.
Vršanský, P. (1997) Piniblattella gen. nov. - the most ancient genus of the family Blattellidae (Blattodea) from the Lower Cretaceous of Siberia. Entomological Problems, 28 (1), 67–79.
Vršanský, P. (2000) Decreasing variability—from the Carboniferous to the Present! (validated on independent lineages of Blattaria). Palaeontological Journal, 34 (Supplement to No. 3), 374–379.
Vršanský, P. (2002) Origin and the Early Evolution of Mantises. Amba projekty, 6 (1), 1–16.
Vršanský, P. (2003) Unique assemblage of Dictyoptera (Insecta - Blattaria, Mantodea, Isoptera) from the Lower Cretaceous of Bon Tsagaan Nuur in Mongolia. Entomological Problems 33 (1–2), 119–151.
Vršanský, P. (2005) Mass mutations of insects at the Jurassic/Cretaceous boundary? Geologica Carpathica, 56 (6), 473–781.
Vršanský, P. (2008a) Mesozoic relative of the common synanthropic German cockroach (Blattodea). Deutsche Entomologische Zeitschrift. 55 (2), 215–221.
Vršanský, P. (2008b) New blattarians and a review of dictyopteran assemblages from the Lower Cretaceous of Mongolia. Acta Palaeontologica Polnica, 53 (1), 129–136.
Vršanský, P., Vishniakova, V.N. & Rasnitsyn, A.P. (2002) Order Blattida Latreille, 1810. In: Rasnitsyn, A.P. & Quicke, D.L.J. (Eds), History of insects. Kluwer Academic Publishers, Dordrecht, The Netherlands, pp. 263–270.
Vršanský P., Cifuentes-Ruiz P., Vidlička L., Čiampor F. J.R. & Vega, F.J. (2011) Afro-Asian cockroach from Chiapas amber and the lost Tertiary American entomofauna. Geologica Carpathica 62 (5), 463–475.
Vršanský, P., Liang J-H. & Ren, D. (2009) Advanced morphology and behaviour of extinct earwig-like cockroaches (Blattida: Fuziidae fam. n.). Geologica Carpathica, 60 (6), 449–462.
Vršanský, P., Vidlička Ľ., Čiampor. F. Jr, & Marsh, F. (2011) Derived, still living cockroach genus Cariblattoides (Blattida: Blattellidae) from the Eocene sediments of Green River in Colorado, USA. Insect Science 00, 1–10, DOI 10.1111/j.1744-7917.2010.01390.x
Wang, T.T., Liang J.H. & Ren, D. (2007) Variability of Habroblattula drepanoides gen. et sp. nov. (Insecta: Blattaria: Blattulidae) from the Yixian Formation in Liaoning, China. Zootaxa, 1443, 17–27.
Wang, T., Liang, J-H., Ren, D. & Shi C. (2007) New Mesozoic cockroaches (Blattaria: Blattulidae) from Jehol Biota of western Liaoning in China. Annales Zoologici, 57 (3), 483–495.
Zherikhin, V.V. (2002) 2.2.2.1.3 Mantida Latreille, 1802. The Mantises (=Mantodea Burmeister, 1838). In: Rasnitsyn, A.P. & Quicke, D.L.J. (Eds), History of insects. Kluwer Academic Publishers, Dordrecht, The Netherlands, pp. 273–276.