1034
Journal of Paleontology
Upper Triassic (Norian) Cow Branch Formation at the Solite Quarry Lagerstätte near the North Carolina-Virginia boundary, USA. It represents the first orthopteran described from the Triassic of North America. The pterostigma-like structure on the wing of C. virginiana n. gen. n. sp. was analyzed using scanning electron microscopy (SEM) and energy dispersive X-ray spectro- scopy (EDS). Similar to pterostigma in modern insects, the pterostigma-like structure in elcanids was probably thickened and sclerotized, but it differs from true pterostigmae on the outer wings of other insects in that it is traversed by more crossveins. Elcanids may have evolved a particular flight mechanism distinct from those of other orthopterans.
Acknowledgments
This research was supported by the National Natural Science Foundation of China (41572010, 41622201, 41688103, 41730317), Chinese Academy of Sciences (XDPB05), U.S. National Science Foundation (EAR-1528553), Clay Mineral Society, Geological Society of America, Sigma Xi Scientific Research Society, and Virginia Tech Department of Geo- sciences. Analyses were conducted at the VT-ICTAS-NCFL with technical assistance from A. Giordani, S. McCartney, and C. Winkler. We thank N.C. Fraser for giving us permission to study this specimen, D.R. Zheng for discussion, and S. Chatzi- manolis, B. Hunda, and three anonymous reviewers for their constructive comments on an earlier draft of the paper.
References
Bechly, G., 1995, Morphologische Untersuchungen am Flügelgeäder der rezenten Libellen und deren Stammgruppenvertreter (Insecta; Pterygota; Odonata): unter besonderer Berücksichtigung der Phylogenetischen Systematik und des Grundplanes der Odonata: Petalura, Special Volume, 1, p. 1–341.
life by a Late Triassic lake margin, in Fraser, N.C., and Sues, H.-D., eds., Terrestrial Conservation Lagerstätten: Windows into the Evolution of Life on Land: Edinburgh, UK, Dunedin Academic Press, p. 105–129.
Gorochov, A.V., 1990, Orthopterans, Gryllida, in Pozdne-Mezozoyskie Nasekomye Vostochnogo Zabaykal’ya. Akademiya Nauk SSSR: Trudy Paleontologicheskogo Instituta, v. 239, p. 210–214.
Gorochov, A.V., 1995, System and evolution of the suborder Ensifera (Orthoptera) (in 2 parts): Proceedings of the Zoological Institute, Russian Academy of Science, v. 260, p. 1–224. 1–213. [In Russian].
Gorochov, A.V., Jarzembowski, E.A., and Coram, R.A., 2006, Grasshoppers and crickets (Insecta: Orthoptera) from the Lower Cretaceous of southern England: Cretaceous Research, v. 27, p. 641–662.
Grimaldi, D., and Engel, M.S., 2005, Evolution of the Insects: New York, Cambridge University Press, 772 p.
Grimaldi, D., Shmakov, A., and Fraser, N., 2004, Mesozoic thrips and early evolution of the order Thysanoptera (Insecta): Journal of Paleontology, v. 78, p. 941–952.
Handlirsch, A., 1906, Die fossilen Insekten und die Phylogenie der rezenten Formen, Parts I–IV: Leipzig, Verlag von Wilhelm Engelmann, p. 1–640.
Kearns, S.L., and Orr, P.J., 2009, Charge contrast imaging of exceptionally- preserved fossils: Palaeontology, v. 52, p. 673–680.
Linnaeus, C., 1758, Systema natura per regna tria naturae secundum classes, ordines, genera, species, cum characteribus, differentiis, synonymis, locis, Tenth edition., Laurentius Salvius, Holmiae, 824 p.
Liutkus, C.M., Beard, J.S., Fraser, N.C., and Ragland, P.C., 2010, Use of fine- scale stratigraphy and chemostratigraphy to evaluate conditions of deposi- tion and preservation of a Triassic Lagerstätte, south-central Virginia: Journal of Paleolimnology, v. 44, p. 645–666.
Liutkus-Pierce, C.M., Fraser, N.C., and Heckert, A.B., 2014, Stratigraphy, sedimentology, and paleontology of the Upper Triassic Solite Quarry, North Carolina and Virginia, in Bailey, C.M., and Coiner, L.V., eds., Elevating Geoscience in the Southeastern United States: New Ideas about Old Terranes—Field Guides for the 2014 GSA Southeastern Section Meeting, Blacksburg, Virginia: Boulder, CO, Geological Society of America, p. 255–269.
Muscente, A.D., and Xiao, S., 2015, Resolving three-dimensional and sub- surficial features of carbonaceous compressions and shelly fossils using backscattered electron scanning electron microscopy (BSE-SEM): Palaios, v. 30, p. 462–481.
Norberg, R.A., 1972, The pterostigma of insect wings an inertial regulator of wing pitch: Journal of Comparative Physiology, v. 81, p. 9–22.
Olivier, G.A., 1789, Histoire naturelle. Entomologie, ou histoire naturelle des crustacés, des arachnides et des insectes: Encyclopédie méthodique, v. 4, p. 12–16. Paris, Chez Panckoucke.
Bechly, G., 1998, New fossil dragonflies from the Lower Cretaceous Crato Formation of north-east Brazil (Insecta: Odonata): Stuttgarter Beiträge zur Naturkunde, Series B, v. 264, p. 1–66.
Béthoux, O., and Nel, A., 2001, Venation pattern of Orthoptera: Journal of Orthoptera Research, v. 10, p. 195–198.
Béthoux, O., and Nel, A., 2002, Venational pattern and revision of Orthoptera sensu nov and sister group. Phylogeny of Orthoptera sensu nov: Zootaxa, v. 96, p. 1–88.
Blagoderov, V., Grimaldi, D.A., and Fraser, N.C., 2007, How time flies for flies: diverse Diptera from the Triassic of Virginia and early radiation of the order: American Museum Novitates, v. 3572, p. 1–39.
Carle, F.L., and Wighton, D.C., 1990, Odonata, in Grimaldi, D.A., ed., Insects from the Santana Formation, Lower Cretaceous of Brazil: Bulletin of American Museum of Natural History, v. 195, p. 41–68.
Chapman, R.F., 1998, The Insects: Structure and Function, (4th edition): Cambridge, Cambridge University Press, 770 p.
Criscione, J., and Grimaldi, D., 2017, The oldest predaceous water bugs (Insecta, Heteroptera, Belostomatidae), with implications for paleolimnology of the Triassic Cow Branch Formation: Journal of Paleontology, v. 91, p. 1166–1177.
Fang, Y., Zhang, H.C., Wang, H., Wang, B., Zheng, D.R., Zhang, Q., Li, S., and Liu, Q., 2015, New Cretaceous Elcanidae from China and Myanmar (Insecta, Orthoptera): Cretaceous Research, v. 52, p. 323–328.
Fraser, N.C., Grimaldi, D.A., Olsen, P.E., and Axsmith, B., 1996, A Triassic Lagerstätte from eastern North America: Nature, v. 380, p. 615–619.
Fraser, N.C., Grimaldi, D.A., Axsmith, B.J., Heckert, A.B., Liutkus-Pierce, C., Smith, D., and Dooley, A.C. Jr., 2017, The Solite Quarry—a window into
Olsen, P.E., Froelich, A.J., Daniels, D.L., Smoot, J.P., and Gore, P.J.W., 1991, Rift basins of early Mesozoic age, in Horton, W., ed., Geology of the Carolinas: Knoxville, TN, University of Tennessee Press, p. 142–170.
Orr, P.J., Kearns, S.L., and Briggs, D.E.G., 2002, Backscattered electron imaging of fossils exceptionally preserved as organic compressions: Palaios, v. 17, p. 110–117.
Peñalver, E., and Grimaldi, D.A., 2010, Latest occurrences of the Mesozoic family Elcanidae (Insecta: Orthoptera), in Cretaceous amber from Myanmar and Spain: Annales de la Societé Entomologique de France, v. 46, p. 88–99.
Sharov, A.G., 1968, Phylogeny of orthopteroid insects: Proceedings of the Paleontological Institute: Russian Academy of Sciences, v. 118, p. 1–217. [In Russian].
Shcherbakov, D.E., 2013, Permian ancestors of Hymenoptera and Raphi- dioptera: ZooKeys, v. 358, p. 45–67.
Tang, Q., Pang, K., Yuan, X., and Xiao, S., 2017, Electron microscopy reveals evidence for simple multicellularity in the Proterozoic fossil Chuaria: Geology, v. 45, p. 75–78.
Tillyard, R.J., 1937, Kansas Permian Insects. Part 17. The order Megasecoptera and additions to the Palaeodictyoptera, Odonata, Protoperlaria, Copeog- natha, and Neuroptera: American Journal of Science, Series 5, v. 33, p. 81–110.
Zessin, W., 1988, Neue Saltatoria (Insecta) aus dem Oberlias Mitteleuropas: Freiberger Forschungshefte C, v. 419, p. 107–121.
Accepted 13 March 2018
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