Abstract
Ipsvicia Tillyard, 1919, and Ipsviciopsis Tillyard, 1922, in Tillyard’s new family, Ipsviciidae, were two of the most impressive insects from the Denmark Hill locality of the Late Triassic (Norian) Blackstone Formation, Ipswich Coal Measures, south-eastern Queensland. Substantial new material, including several fragmentary body specimens, from the Dinmore locality in the same formation, has enabled a revision of the two genera, with the following results: Ipsvicia jonesi Tillyard, 1919 (= Ipsvicia maculata Tillyard, 1919, syn. nov., = Ipsvicia acutipennis Tillyard, 1919, syn. nov.), Ipsviciopsis elegans Tillyard, 1922 (= Ipsviciopsis magna Tillyard, 1922, syn. nov.). The tegmen of Ipsviciopsis is distinguished from that of Ipsvicia on its narrower shape and less convex costal margin, simpler surface sculpture, longer basal cell, sinuous R, only slightly angulate CuA, and sinuous base of 1A. The tegmina of both I. jonesi and I. elegans are noteworthy for their variability in apical venation and shape of the apex, with no two specimens quite the same. The tegmen of I. jonesi has extraordinary surface sculpture comprising patches of fine tubercles set in a coarser tuberculate/punctate groundmass, the patches extremely variable in shape and pattern, again with no two specimens the same. A new diagnosis of the Ipsviciidae has identified the unique form of the distal portion of Sc (dSc—a long groove running along the costal space and crossed by many costal veinlets), the complex and highly variable apical venation, and the simple CuA, as the most distinctive characters. Analysis of all previous taxa which have been referred to the Ipsviciidae restricts the family to the Middle to Late Triassic, with records from Australia, China, France, Germany, Japan and Kyrgyzstan, and with one Early Jurassic record from Kyrgyzstan. Ipsvicia langenbergensis Barth, Ansorge et Brauckmann, 2011, from the Late Triassic of Germany meets the diagnosis of Ipsviciopsis and is transferred as Ipsviciopsis langenbergensis (Barth, Ansorge et Brauckmann, 2011) comb. nov. All previous Permian records of the family have now been transferred to other families of the Scyctinopteroidea, and there are also numerous additional unconfirmed Triassic records. The new body specimens of both Ipsvicia and Ipsviciopsis show that the ipsviciids where robust hemipterans with tough, coriaceous, fairly flatly-folded tegmina, with a large, shield-like, highly sculptured pronotum covering all but the mesoscutellum, and apparently much of the head. The body form is not dissimilar to ground-dwelling cockroaches, or more especially some ground-dwelling moist environment Heteroptera, such as the nepomorph family Gelastocoridae. This body form, as well as their frequent occurrence among the rich Dicroidium-dominant flood plain and swamp flora of the Dinmore locality, where they represent nearly 20% of the preserved insect fauna (cockroaches represent 25%), suggest that the ipsviciids were ground-dwelling insects of moisture-rich floral environments.
References
Ansorge, J. (1994) Rhoeniella granulata gen. et sp. vov. (Insecta: Hemiptera) aus dem Oberen Buntsandstein der Rhӧn (Hessen). Senckenbergiana lethaea, 74, 325–327.
Barth, G., Ansorge, J. & Brauckmann, C. (2011) First record of the genus Ipsvicia (Hemiptera: Ipsviciidae) outside Gondwana—an Australian genus from the Upper Triassic of Germany. Polish Journal of Entomology, 80, 645–657.
https://doi.org/10.2478/v10200-011-0050-z
Becker-Migdisova, E.E. (1948) Permian cicadas of the family Scytinopteridae from the Soyana River. Trudy Paleontologicheskogo Instituta Akademiya Nauk SSSR, 15, 1–42. [in Russian]
Becker-Migdisova, E.E. (1961) Suborder Rhynchota, In: Rohdendorf, B.B., Becker-Migdisova, E.E., Martynova, O.M. & Sharov, A.G. (Eds.), Paleozoic insects of the Kuznetsk Basin, Trudy Paleontologicheskogo Instituta Akademiya Nauk SSSR, 85, pp 286–393. [in Russian]
Becker-Migdisova, E.E. (1962a) Superorder Rhynchota. Insects with proboscis. In: Rohdendorf, B.B. (Ed.), Fundamentals of Paleontology. Vol. 9. Arthropoda, Tracheata, Chelicerata. Akademiya Nauk SSSR, Moscow, pp. 216–317. [in Russian; English translation 1991, Smithsonian Institution Libraries and The National Science Foundation, Washington, D.C.]
Becker-Migdisova, E.E. (1962b) Nekotorye novye poluzhestkokrylye i senoedy [Some new Hemiptera and Psocoptera]. Paleontologicheskij zhurnal, 1962(1), 89–104 [In Russian].
Brauckmann, C. & Schlüter, T. (1993) Neue Insekten aus der Trias von Unter-Franken. Geologica et Palaeontologica, 27, 181–199.
Carpenter, F.M. (1992) Superclass Hexapoda. In: Kaesler, R.L. (Ed.), Treatise on Invertebrate Paleontology. Part R. Arthropoda 4. Vol. 3. The Geological Society of America Inc. Boulder, Colorado and The University of Kansas, and Lawrence, Kansas, pp. i–xxi + 1–277.
China, W.E. (1962) South American Peloridiidae. Transactions of the Royal Entomological Society of London, 114, 131–161.
https://doi.org/10.1111/j.1365-2311.1962.tb01077.x
Dijk, D.E. van & Geertsema, H. (1999) Permian insects from the Beaufort Group of Natal, South Africa. Annals Natal Museum, 40, 137–171. [https://hdl.handle.net/10520/AJA03040798_102]
Dunstan, B. (1923) Mesozoic insects of Queensland Part I.—Introduction and Coleoptera. Geological Survey of Queensland Publication, 273, i–v + 1–76.
Evans, J.W. (1943) Upper Permian Homoptera from New South Wales. Records of the Australian Museum, 21, 180–198.
https://doi.org/10.3853/j.0067-1975.21.1943.534
Evans, J.W. (1946) A natural classification of leaf-hoppers. Part 1. Transactions of the Entomological Society of London, 96, 47–60.
https://doi.org/10.1111/j.1365-2311.1946.tb00442.x
Evans, J.W. (1950) A re-examination of an Upper Permian insect, Paraknightia magnifica Ev. Records of the Australian Museum, 22, 246–250.
https://doi.org/10.3853/j.0067-1975.22.1950.605
Evans, J.W. (1956) Palaeozoic and Mesozoic Hemiptera (Insecta). Australian Journal of Zoology, 4, 165–158.
https://doi.org/10.1071/ZO9560165
Evans, J.W. (1963) The systematic position of the Ipsviciidae (Upper Triassic Hemiptera) and some new Upper Permian and Middle Triassic Homoptera from Australia (Insecta). Journal of the Entomological Society of Queensland, 2, 17–23.
https://doi.org/10.1111/j.1440-6055.1963.tb00384.x
Evans, J.W. (1964) The periods of origin and diversification of the Superfamilies of the Homoptera-Auchenorhyncha (Insecta) as determined by a study of the wings of Palaeozoic and Mesozoic fossils. Proceedings of the Linnean Society of London, 175, 171–181.
https://doi.org/10.1111/j.1095-8312.1964.tb00929.x
Fujiyama, I. (1973) Mesozoic insect fauna of east Asia Part I. Introduction and Upper Triassic faunas. Bulletin of the National Science Museum of Tokyo 16, 331–386.
Gall, J.-C. & Grauvogel-Stamm, L. (2005) The early Middle Triassic ‘Grès à Voltzia’ Formation of eastern France: a model of environmental refugium. Comptes rendus Palevol 4, 637–652.
https://doi.org/10.1016/j.crpv.2005.04.007
Gorochov, A.V. (2005) Review of Triassic Orthoptera with descriptions of new and little known taxa: Part 1. Paleontological Journal 39, 178–186.
Hamilton, K.G.A. (1992) Lower Cretaceous Homoptera from the Koonwarra fossil bed in Australia, with a new superfamily and synopsis of Mesozoic Homoptera. Annals of the Entomological Society of America, 85, 423–430.
https://doi.org/10.1093/aese/85.4.423
Handlirsch, A. (1906–1908) Die fossilen Insekten und die Phylogenie der rezenten Formen. Verlag von Wilhelm Engelmann, Leipzig, ix + 1430 pp + 51 plates.
https://doi.org/10.5962/bhl.title.34145
Handlirsch, A. (1939) Neue Untersuchungen über die fossilen Insekten mit Ergänzungen und Nachträgen sowie Ausblicken auf phylogenetische, palaeogeographische und allgemein biologische Probleme. II. Teil. Annalen des naturhistorischen Museums in Wien, 49, 1–240. [https://www.jstor.org/stable/pdf/41768379.pdf]
Hong, Y.-C. (1980) Granulidae, a new family of Homoptera from the Middle Triassic of Tongchuan, Shanxi Province. Acta Zootaxonomica Sinica, 5, 63–70.
Jeannel, R. (1947) Paleontologie et peuplement de la Terre. Introduction à l’entomologie, 1 (3), 1–101, 14 pls.
Jeannel, R. (1949) Les insectes. Classification et phylogénie. Les insectes fossiles. In: Grassé, P.-P. (Ed.), Traité de zoologie. Insectes (Paléontologie, géonémie, insectes inférieurs et Coléoptères). 9. Masson et Cie, Paris, pp. 1–110.
Jeannel, R. (1960) Introduction to entomology. Hutchison & Co., London, 344 pp.
Jeannel, R. (1979) Paléontologie et peuplement de la Terre. Deuxième edition. Société nouvelle des éditions Boubée & Cie, Paris, 101 pp., 14 pls.
Jell, P.A. (1993) Late Triassic homopterous nymph from Dinmore, Ipswich Basin. Memoirs of the Queensland Museum, 33, 360.
Jell, P.A. (2004) The fossil insects of Australia. Memoirs of the Queensland Museum, 50, 1–124.
https://doi.org/10.1082/j.2204-1478.50-1.2004.2004.01
Kramer, S. (1950) The morphology and phylogeny of auchenorrhynchous Homoptera (Insecta). Illinois Biological Monographs, 20 (4), i–vii + 1–111.
https://doi.org/10.5962/bhl.title.50340
Kukalová-Peck, J. (1991) Fossil history and the evolution of hexapod structures. In: Naumann, I. D. (Ed.), Insects of Australia A textbook for students and research workers. Melbourne University Press, Carlton, Victoria, pp. 141–179.
Lambkin, K.J. (2016) Revision of the Scytinopteridae (Hemiptera: Cicadomorpha: Scytinopteroidea) of the Queensland Triassic. Zootaxa, 4117 (4), 580–590.
https://doi.org/10.11646/zootaxa.4117.4.9
Lambkin, K.J. (2018) A new genus and species of the family Archipsyllidae (Insecta: Paraneoptera: Permopsocida) from the Late Triassic of Queensland. Zootaxa, 4382 (1), 192–194.
https://doi.org/10.11646/zootaxa.4382.1.9
Lambkin, K.J. (2020) Planthoppers (Insecta: Hemiptera: Fulgoromorpha: Fulgoroidea: Surijokocixiidae) of the Queensland Triassic. Memoirs of the Queensland Museum—Nature, 62, 123–131.
https://doi.org/10.17082/j.2204-1478.62.2020.2019-04
Lin, Q.B. (1982a) Insecta. In: Nanjing Institute of Geology and Mineral Resources (Ed.), Paleontological atlas of east China Shaanxi Part 3 Volume of Mesozoic and Cenozoic. Geological Publishing House, Beijing, pp. 148–155. [in Chinese]
Lin, Q.B. (1982b) Class Insecta. In: Xi’an Institute of Geology and Mineral Resources (Ed.), Paleontological atlas of northwest China Shaanxi Gansu Ningxia volume Part III Mesozoic and Cenozoic. Geological Publishing House, Beijing, pp. 70–83 [in Chinese]
Lin, Q.B. (1992) Late Triassic insect fauna from Toksun, Xinjiang. Acta Palaeontologica Sinica, 31, 313–335 [in Chinese with English summary]
Linnaeus, C. (1758) Systema naturae per regna tria naturae, secundum classes, ordines, genera, species, cum characteribus, differentiis, synonymis, locis. Tomus I. Editio decima, reformata. Laurentii Salvii, Holmiae, [4] + 824 pp.
https://doi.org/10.5962/bhl.title.542
Martins-Neto, R.G., Brauckmann, C., Gallego, O.F & Carmona, M. J. (2006) The Triassic insect fauna from Argentina.—Blattoptera, Glosselytrodea, Miomoptera, Auchenorrhyncha, and Coleoptera from the Los Rastros Formation (Bermejo Basin), Los Chañares locality (La Rioja Province). Clausthaler Geowissenschaften, 5, 1–9.
Papier, F., Nel, A., Grauvogel-Stamm, L. & Gall, J.-C. (1997) La plus ancienne sauterelle Tettigoniidae, Orthoptera (Trias, NE France): mimétisme ou exaptation? Die älteste Tettigoniidae, Orthoptera (Trias, NO-Frankreich): Mimikry oder Exaptation? Paläontologische Zeitschrift, 71, 71–77.
https://doi.org/10.1007/BF03022547
Popov, Y.A. (1971) Historical development of the infraorder Nepomorpha (Heteroptera). Trudy Paleontologicheskogo Instituta Akademiya Nauk SSSR, 129, 1–230. [in Russian]
Popov, Y.A. (1980) Superorder Cimicidea Laicharting, 1781. Order Cimicina Laicharting, 1781. Hemipterans, or rhynchotans. In: Rohdendorf, B.B. & Rasnitsyn, A.P. (Eds), Historical development of the class Insecta. Trudy Paleontologicheskogo Instituta Akademii Nauk SSSR, 175, pp. 58–69.
Purdy, D.J. & Cranfield, L.C. (2013) Ipswich Basin. In: Jell, P.A. (Ed.), Geology of Queensland. Geological Survey of Queensland, Brisbane, pp. 391–396.
Raven, R.J., Jell, P.A. & Knezour, R.A. (2015) Edwa maryae gen. et sp. nov. in the Norian Blackstone Formation of the Ipswich Basin – the first Triassic spider (Mygalomorphae) from Australia. Alcheringa, 39, 259–263.
https://doi.org/10.1080/03115518.2015.993300
Shcherbakov, D.E. (1984) Sistema i filogeniya permskikh Cicadomorpha (Cimicida, Cicadina). Palaeontologicheskiï Zhurnal, 18 (2), 87–97. [in Russian, publishjed in English as: Shcherbakov, D.E. (1984) A system and the phylogeny of Permian Cicadomorpha (Cimicida, Cicadina). Paleontological Zhurnal, 2, 89–101.]
Shcherbakov, D.E. (1996) Origin and evolution of the Auchenorrhyncha as shown by the fossil record. In: Schaefer, C.W. (Ed.), Studies on Hemipteran Phylogeny. Entomological Society of America, Lanham, Maryland, pp. 31–45.
Shcherbakov, D.E. (2002) the 270 million year history of Auchenorrhyncha (Homoptera). Denisia, 4, 29–35.
Shcherbakov, D.E. (2008) Madygen, Triassic Lagerstätte number one, before and after Sharov. Alavesia, 2, 113–124.
Shcherbakov, D.E. (2011) New and little-known families of Hemiptera Cicadomorpha from the Triassic of Central Asia – early analogs of treehoppers and planthoppers. Zootaxa, 2836 (1), 1–26.
https://doi.org/10.11646/zootaxa.2836.1.1
Shcherbakov, D.E. & Popov, Y.A. (2002) Superorder Cimicidea Laicharting, 1781 Order Hemiptera Linné, 1758. The bugs, cicadas, plantlice, scale insects, etc. (=Cimicida Laicharting, 1781, = Homoptera Leach, 1815 + Heteroptera Latreille, 1810). In: Rasnitsyn, A.P. & Quicke, D.L.J. (Eds.), History of Insects. Kluwer Academic Publishers, Dordrecht, pp. 143–157.
Szwedo, J., Bourgoin, T. & Lefebvre, F. (2004) Fossil planthoppers (Hemiptera: Fulgoromorpha) of the world An annotated catalogue with notes on Hemiptera classification. Studio 1, Warsaw, 199 pp., 8 pls.
Szwedo, J. (2018) The unity, diversity and conformity of bugs (Hemiptera) through time. Earth and Environmental Science Transactions of the Royal Society of Edinburgh, 107, 109–128.
https://doi.org/10.1017/S175569101700038X
Tillyard, R.J. (1916) Mesozoic and Tertiary insects of Queensland and New South Wales. Geological Survey of Queensland Publication, 253, 11–64.
Tillyard, R.J. (1919a) Mesozoic insects of Queensland. No. 7. Hemiptera Homoptera; with a note on the phylogeny of the Suborder. Proceedings of the Linnean Society of New South Wales, 44, 857–896.
Tillyard, R.J. (1919b) [Letter to Benjamin Dunstan, 10 July 1919], unpublished. In: Dunstan, B. & Tillyard, R.J. & National Library of Australia & Australian Joint Copying Project & State Library of New South Wales. [19--]. Papers of Benjamin Dunstan (as filmed by the AJCP) [microform]: [M2493], 1913–1923, Originals held by the Palaeontology Library, Natural History Museum, Cromwell Road, South Kensington, London. Available from: https://nla.gov.au/nla.obj-838028448 (accessed 27 September 2020)
Tillyard, R.J. (1919c) [Letter to Benjamin Dunstan, 23 October 1919], as above.
Tillyard, R.J. (1920) [Letter to Benjamin Dunstan, 19 March 1920], as above.
Tillyard, R.J. (1922) Mesozoic insects of Queensland. No. 9. Orthoptera, and additions to the Protorthoptera, Odonata, Hemiptera and Planipennia. Proceedings of the Linnean Society of New South Wales, 47, 447–470. [https://www.biodiversitylibrary.org/page/35116952]
Tillyard, R.J. (1923) Mesozoic insects of Queensland. No. 10. Summary of the Upper Triassic insect fauna of Ipswich, Q. (With an appendix describing new Hemiptera and Planipennia). Proceedings of the Linnean Society of New South Wales, 48, 481–498. [https://www.biodiversitylibrary.org/page/35067038]
Tillyard, R.J. (1926) The insects of Australia and New Zealand. Angus & Robertson, Sydney, xvi + 560 pp., 44pls.
Tillyard, R.J. & Dunstan, B. (1924) Mesozoic insects of Queensland. Queensland Geological Survey Publication, 273, i–v + 1–76 + 1–242.
Todd, E.L. (1960) The Gelastocoridae of Australia (Hemiptera). Pacific Insects, 2, 171–194. [http://hbs.bishopmuseum.org/pi/pdf/2(2)-171-194.pdf]
Zheng, D., Chang, S-C., Wang, H., Fang, Y., Wang, J., Feng, C., Xie, G., Jarzembowski, E. A., Zhang, H. & Wang, B. (2018) Middle-Late Triassic insect radiation revealed by diverse fossils and isotopic ages from China. Science Advances, 4, 1–7.
https://doi.org/10.1126/sciadv.aat1380