Abstract
New specimens and a re-examination of their holotypes have clarified the status of five species of the extinct cicadomorphan family Hylicellidae from the Late Triassic (Norian) Mount Crosby Formation of south-eastern Queensland. All were found to be conspecific, resulting in the following synonymies: Mesothymbris Evans, 1956 (= Triassoscytina Evans, 1956, syn. nov. = (in part) Triassoscytinopsis Evans, 1956, syn. nov.), Mesothymbris perkinsi Evans, 1956 (= Mesothymbris woodwardi Evans, 1956, syn. nov. = Triassoscytina incompleta Evans, 1956, syn. nov. = Triassoscytinopsis stenulata Evans, 1956, syn. nov. = Triassoscytinopsis aberrans Evans, 1956, syn. nov.). The Hylicellidae is still poorly defined as is its generic composition. Mesothymbris, however, is clearly distinct from the other Mount Crosby hylicellids, Hylicella Evans, 1956, and Triassocotis, Evans, 1956, in the quite distal primary fork of R, the angled RA at the point of separation of RA1 with RA2 directed towards the apex of the tegmen, the upright RA1, the shape of the intra-medial cell, and CuA just distal to the basal cell strongly curved and very closely approximating the claval suture. The new synonymies further clarify the composition of the cicadomorphan fauna of the Mount Crosby Formation, which as a result of this and other recent revisions, now comprises 16 nominal species in the Dysmorphoptilidae, Hylicellidae and Mesojabloniidae, as well as three species incertae sedis. In the presence of Dysmorphoptilidae and Hylicellidae, the Mount Crosby cicadomorphan fauna is similar to that of the younger Late Triassic Blackstone Formation at nearby Denmark Hill and Dinmore. It differs significantly, however, in the absence, after 90 years of collecting of 100s of specimens, of any representatives of the Dunstaniidae, Mesogereonidae, or Tettigarctidae, families so characteristic of the Denmark Hill/Dinmore fauna. Whether this difference is biogeographical, ecological, or simply as a result of differential preservation is unknown.
References
Allen, R.J. (1961) The Kholo Sub-group of the Ipswich Coal Measures. Geological Survey of Queensland Publication, 300, 1–14.
Becker-Migdisova, E.E. (1962) 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.]
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.
Chen, J., Szwedo, J., Wang, B., Zheng, Y., Wang, Y., Wang, X. & Zhang, H. (2108) The first Mesozoic froghopper in amber from northern Myanmar (Hemiptera, Cercopoidea, Sinoalidae). Cretaceous Research, 85, 243–249.
https://doi.org/10.1016/j.cretres.2017.10.029
Evans, J.W. (1956) Palaeozoic and Mesozoic Hemiptera (Insecta). Australian Journal of Zoology, 4, 165–258.
https://doi.org/10.1071/ZO9560165
Evans, J.W. (1961) Some Upper Triassic Hemiptera from Queensland. Memoirs of the Queensland Museum, 14, 13–23.
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
Evans, J.W. (1971) Some Upper Triassic Hemiptera from Mount Crosby, Queensland. Memoirs of the Queensland Museum, 16, 145–152.
Fu, Y. & Huang, D. (2019a) New sinoalids (Insecta: Hemiptera: Cercopoidea) from Middle to Upper Jurassic strata at Daohugou, Inner Mongolia, China. Alcheringa: An Australasian Journal of Palaeontology, 43 (2), 246–256.
https://doi.org/10.1080/03115518.2018.1528509
Fu, Y. & Huang, D. (2019b) New Early Cretaceous hylicellids (Insecta: Hemiptera: Cicadomorpha) from southwest Beijing, China. Palaeoentomology, 2, 17–21.
https://doi.org/10.11646/palaeoentomology.2.1.4
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/aesa/85.4.423
Jell, P.A. (2004) The fossil insects of Australia. Memoirs of the Queensland Museum, 50, 1–124.
Lambkin, K.J. (2015) Revision of the Dysmorphoptilidae with emarginate tegmina (Hemiptera: Auchenorryncha: Cicadomorpha: Prosboloidea) of the Queensland Triassic. Zootaxa, 3936 (3), 357–374.
https://doi.org/10.11646/zootaxa.3936.3.3
Lambkin, K.J. (2016a) Revision of the Dysmorphoptilidae (Hemiptera: Cicadomorpha: Prosboloidea) of the Queensland Triassic—Part 2. Zootaxa, 4092 (2), 207–218.
https://doi.org/10.11646/zootaxa.4092.2.4
Lambkin, K.J. (2016b) 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. (2019) Mesodiphthera Tillyard, 1919, from the Late Triassic of Queensland, the oldest cicada (Hemiptera: Cicadomorpha: Cicadoidea: Tettigarctidae). Zootaxa, 4567 (2), 358–366.
https://doi.org/10.11646/zootaxa.4567.2.8
Martins-Neto, R.G., Gallego, O.F & Melchor, R.N. (2003) The Triassic insect fauna from South America (Argentina, Brazil and Chile): a checklist (except Blattoptera and Coleoptera) and descriptions of new taxa. Acta zoologica cracoviensia, 46, 229–256.
Nel, A., Prokop, J., Nel, P., Grandcolas, P., Huang, D-Y., Roques, P., Guilbert, E., Dostál, O. & Szwedo, J. (2012) Traits and evolution of wing venation pattern in paraneopteran insects. Journal of Morphology, 273, 480–506.
https://doi.org/10.1002/jmor.11036
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. (1988) New cicadas (Cicadina) from the later Mesozoic of Transbaikalia. Paleontological Journal, 1988, 52–63.
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. (2011) New and little-known families of Hemiptera Cicadomorpha from the Triassic of Central Asia—early analogs of treehoppers and planthoppers. Zootaxa, 2836, 1–26.
https://doi.org/10.11646/zootaxa.2836.1.1
Shcherbakov, D.E. (2012) A new family of Mesozoic Hylicellidae (Homoptera: Cicadomorpha). Russian Entomological Journal, 21, 441–444.
https://doi.org/10.15298/rusentj.21.4.10
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
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.
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. (1918) Mesozoic insects of Queensland. No. 4. Hemiptera Heteroptera: The family Dunstaniidae, with a note on the origin of the Heteroptera. Proceedings of the Linnean Society of New South Wales, 43, 568–592.
Tillyard, R.J. (1919) 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. (1921) Mesozoic insects of Queensland. No. 8. Hemiptera Homoptera (Contd). The genus Mesogereon; with a discussion of its relationship with the Jurassic Palaeontinidae. Proceedings of the Linnean Society of New South Wales, 46, 270–284.
https://doi.org/10.5962/bhl.part.14018
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.
Wang, B., Szwedo, J., Zhang, H. & Fang, Y. (2010) The major diversification of Cicadomorpha in the Jurassic (Insecta: Hemiptera). Earth Science Frontiers, 17, 186–187.