Abstract
†Tunguskagyrus, a Late Permian beetle fossil, was assigned to the adephagan family Gyrinidae in the original description. This placement was challenged in a recent study, where the genus was considered as part of a broad new concept of the extinct family †Triaplidae and transferred from Adephaga to Archostemata. Even though some structural features deviate from the typical pattern found in extant Gyrinidae, we maintain the placement of †Tunguskagyrus in the stemgroup of this family as the most plausibe interpretation. The suggested taxonomic changes concerning †Triaplidae (and †Schizophoroidea) are invalidated by an insufficient phylogenetic approach and several misinterpretations of the fossil imprints. The arguments are based on 1) general habitual similarity, 2) ubiquitous characters, 3) plesiomorphies, or 4) apomorphic conditions (compared to the Coleopteran groundplan) occurring in many beetle groups, strongly affected by homoplasy. The taxonomic shift of †Tunguskagyrus and the new concept of †Schizophoroidea and †Triaplidae are discarded due to the lack of a defined phylogenetic methodology and synapomorphies.
References
Archangelsky, M., Beutel, R.G. & Komarek, A. (2016) 12. Hydrophiloidea Latreille, 1802. In: Beutel, R.G. & Leschen, R.A.B. (Vol. Eds.) Handbook of Zoology, Vol. IV Arthropoda: Insecta. Part 38. Coleoptera, Vol. 1: Morphology and Systematics (Archostemata, Adephaga, Myxophaga, Polyphaga (partim). 2nd Ed. Walter De Gruyter, Berlin, New York, pp. 231–272.
Belkaceme, T. (1986) Skelet und Muskulatur der Hinterhüfte von Haliplus lineatocollis Mrsh. (Haliplidae, Coleoptera). Stuttgarter Beiträge zur Naturkunde Serie (A), 393, 1–12.
Belkaceme T. (1991) Skelet und Muskulatur des Kopfes und Thorax von Noterus laevis Sturm. Ein Beitrag zur Morphologie und Phylogenie der Noteridae (Coleoptera: Adephaga). Stuttgarter Beiträge zur Naturkunde Serie A, 462, 1–94.
Beutel, R.G. (1986) Skelet und Muskulatur des Kopfes und Thorax von Hygrobia tarda (Herbst). Ein Beitrag zur Klärung der phylogenetischen Beziehungen der Hydradephaga (Insecta: Coleoptera). Stuttgarter Beiträge zur Naturkunde (A), 388, 1–54.
Beutel, R.G. (1989) The head of Spanglerogyrus albiventris Folkerts (Coleoptera: Gyrinidae). Contribution towards clarification of the phylogeny of Gyrinidae and Adephaga. Zoologische Jahrbücher für Anatomie, 118, 431–461.
Beutel, R.G. (1992) Phylogenetic analysis of thoracic structures of Carabidae (Coleoptera). Journal of Zoological Systematics and Evolutionary Research, 30, 53–74.
https://doi.org/10.1111/j.1439-0469.1992.tb00390.x
Beutel, R.G. (1997) Über Phylogenese und Evolution der Coleoptera (Insecta), insbesondere der Adephaga. Verhandlungen des Naturwissenschaftlichen Vereins Hamburg (NF), 31, 1–164.
Beutel, R.G. & Haas, F. (2000) Phylogenetic relationships of the suborders of Coleoptera (Insecta). Cladistics, 16, 103–141.
https://doi.org/10.1111/j.1096-0031.2000.tb00350.x
Beutel, R.G., Ge, S.Q. & Hörnschemeyer, T. (2008) On the head morphology of Tetraphalerus, the phylogeny of Archostemata and the basal branching events in Coleoptera. Cladistics, 24, 270–298.
https://doi.org/10.1111/j.1096-0031.2007.00186.x
Beutel, R.G., Leschen, R.A.B. & Friedrich, F. (2009) Darwin, Beetles and Phylogenetics. Naturwissenschaften, 96 (11), 1293–1312.
https://doi.org/10.1007/s00114-009-0601-2
Beutel, R.G. & Roughley, R.E. (1988) On the systematic position of the family Gyrinidae (Coleoptera: Adephaga). Journal of Zoological Systematics and Evolutionary Research, 26, 380–400.
https://doi.org/10.1111/j.1439-0469.1988.tb00324.x
Beutel, R.G., Wang, B., Tan, J.J., Ge, S.Q., Dong, R. & Yang, X.K. (2013) On the phylogeny and evolution of Mesozoic and extant lineages of Adephaga (Coleoptera, Insecta). Cladistics, 29, 147–165.
https://doi.org/10.1111/j.1096-0031.2012.00420.x
Beutel, R.G, Yan, E., Richter, A., Büsse, S., Miller, K.B., Yavorskaya, M. & Wipfler, B. (2017) The head of Heterogyrus milloti (Coleoptera: Gyrinidae) and its phylogenetic implications. Arthropod Systematics and Phylogeny, 75, 261–280.
Beutel, R.G., Yan, E., Yavorskaya, M., Büsse, S., Gorb, S.N. & Wipfler, B. (2018) On the thoracic anatomy of the Madagascan Heterogyrus milloti and the phylogeny of Gyrinidae (Coleoptera). Systematic Entomology, 44 (2), 336–360.
https://doi.org/10.1111/syen.12325
Bils, W. (1976) Das Abdomenende weiblicher, terrestrisch lebender Adephaga (Coleoptera) und seine Bedeutung für die Phylogenie. Zoomorphology, 84 (2), 113–193.
https://doi.org/10.1007/BF00999711
Burmeister, E.G. (1976) Der Ovipositor der Hydradephaga (Coleoptera) und seine phylogenetische Bedeutung unter besonderer Berücksichtigung der Dytiscidae. Zoomorphology, 85 (3), 165–257.
https://doi.org/10.1007/BF00993515
Crowson, R.A. (1975) The evolutionary history of Coleoptera, as documented by fossil and comparative evidence. Atti del X Congresso Nazionale Italiano di Entomología, Sassari (1975), 47–90.
Crowson, R.A. (1982) Computers versus imagination in the reconstruction of phylogeny. In: Joysey, K.A. & Friday, A.E. (Eds.), Problems of phylogenetic reconstruction. Academic Press, London, pp. 245–255.
Crowson, R.A. (1991) A critique of current cladistics. Journal of Natural History, 25 (1), 3–5.
https://doi.org/10.1080/00222939100770021
Fikáček, M., Beutel,R.G., Cai,Ch.Y., Lawrence,J.F., Newton,A.F., Solodovnikov,A., Ślipiński,A., Thayer, M.K. & Yamamoto, S.. (In press) Reliable placement of beetle fossils via phylogenetic analyses—Triassic Leehermania as a case study (Staphylinidae or Myxophaga?). Systematic Entomology,
Friedrich, F, Farrell, B.D. & Beutel, R.G. (2009) The thoracic morphology of Archostemata and the relationships of the extant suborders of Coleoptera (Hexapoda). Cladistics, 25, 1–37.
https://doi.org/10.1111/j.1096-0031.2008.00233.x
Hennig, W. (1950) Grundzüge einer Theorie der phylogenetischen Systematik. Deutscher Zentralverlag, Berlin, 370 pp.
Hennig, W. (1966) Phylogenetic Systematics. University of Illinois Press, Urbana, Illinois, 263 pp.
Kirejtshuk, A.G. (1995) System, evolution of the way of life, and phylogeny of the order Coleoptera. Entomological Reviews, 74 (3), 12–31.
Kirejtshuk, A.G. & Prokin, A.A. (2018) The position of the Palaeozoic genus Tunguskagyrus Yan, Beutel et Lawrence in the family Triaplidae sensu n. (Coleoptera, Archostemata: Schizophoroidea). Entomological Review, 98 (7), 872–882.
https://doi.org/10.1134/S0013873818070084
Kjer, K.M., Simon, C., Yavorskaya, M. & Beutel, R.G. (2016)Progress, pitfalls, and parallel universes: A history of insect phylogenetics. Journal of the Royal Society Interface, 13, 20160363.
https://doi.org/10.1098/rsif.2016.0363
Kukalová-Peck, J. & Lawrence, J.F. (2004) Relationships among coleopteran suborders and major endoneopteran lineages: evidence from hind wing characters. European Journal of Entomology, 101, 95–144.
https://doi.org/10.14411/eje.2004.018
Lawrence, J.F. (1999) The Australian Ommatidae (Coleoptera: Archostemata): new species, larva and discussion of relationships. Invertebrate Taxonomy, 13, 369–390.
https://doi.org/10.1071/IT99008
Lawrence, J.F. & Newton, A.F. (1982) Evolution and classification of beetles. Annual Review of Ecology and Systematics, 13, 261–290.
https://doi.org/10.1146/annurev.es.13.110182.001401
Lawrence, J.F., Ślipiński, S.A., Seago, A.E., Thayer, M.K., Newton, A.F. & Marvaldi, A.E. (2011) Phylogeny of the Coleoptera based on morphological characters of adults and larvae. Annales Zoologici, 61, 1–271.
https://doi.org/10.3161/000345411X576725
Maddison, W.P. & Maddison, D.R. (2011) Mesquite: a modular system for evolutionary analysis. Version 2.75. Available from: https://www.mesquiteproject.org/ (Accessed 28 Aug. 2019)
Minelli, A. (1993) Biological Systematics: The State of the Art. Chapman & Hall, London, 387 pp.
Misof, B., Liu, Sh., Meusemann, K., Peters, R.S., Donath, A., Mayer, C., Frandsen, P.B., Ware, J., Flouri, T., Beutel, R.G., Niehuis, O., Petersen, M., Izquierdo-Carrasco, F., Wappler, T., Rust, J., Aberer, A.J., Aspöck, U., Aspöck, H., Bartel, D., Blanke, A., Berger, S., Böhm, A., Buckley, T., Calcott, B., Chen, J. Friedrich, F., Fukui, M., Fujita, M., Greve, C., Grobe, G., Gu, Sh., Huang, Y., Jermiin, L.S., Kawahara, A.Y., Krogmann, L., Kubiak, M., Lanfear, R., Letsch, H. Li, Y., Li, Zh., Li, J., Lu, H., Machida, R., Mashimo, Y., Kapli, P., McKenna, D.D., Meng, G., Nakagaki, Y., Navarrete-Heredia, J.L., Ott, M., Ou, Y., Pass, G., Podsiadlowski, L., Pohl, H., Reumont, B.M. v., Schütte, K., Sekiya, K., Shimizu, Sh., Slipinski, A., Stamatakis, A., Song, W., Su, X., Szucsich, N.U., Tan, M., Tan, X., Tang, M., Tang, J., Timelthaler, G., Tomizuka, Sh., Trautwein, M., Tong, X., Uchifune, T., Walzl, M.G., Wiegmann, B.M., Wilbrandt, J., Wipfler, B., Wong, T.K.F., Wu, Q., Wu, G., Xie, Y., Yang, Sh., Yang, Q., Yeates, D.K., Yoshizawa, K., Zhang, Q., Zhang, R., Zhang, W., Zhang, Y., Zhao, J., Zhou, Ch., Zhou, L., Ziesmann, T., Zou, Sh., Li, Y., Xu, X., Zhang, Y., Yang, H., Wang, J., Wang, J., Kjer, K.M. & Zhou, X. (2014) Phylogenomics resolves the timing and pattern of insect evolution. Science, 346, 763–767.
https://doi.org/10.1126/science.1257570
Niehuis, O., Hartig, G., Grath, S., Pohl, H., Lehmann, J.,Tafer, H., Donath, A., Krauss, V., Eisenhardt, C., Hertel, J., Petersen, M., Mayer, C., Meusemann, K., Peters, R.S., Stadler, P.F., Beutel, R.G., Bornberg-Bauer, E., McKenna, D.D. & Misof, B. (2012)Genomic and morphological evidence converge to resolve the enigma of Strepsiptera. Current Biology, 22 (14), 1309–1313.
https://doi.org/10.1016/j.cub.2012.05.018
Patterson, C. (1980) Cladistics. Biologist, 27, 234–240.
Peters, R.S., Meusemann, K., Petersen, M., Wilbrandt, J., Ziesmann, J., Donath, A., Mayer, C., Kjer, K.M., Aspöck, U., Aspöck, H., Aberer, A., Stamatakis, A., Friedrich, F., Hünefeld, F., Niehuis, O., Beutel, R.G. & Misof, B. (2014) The evolutionary history of holometabolous insects inferred from transcriptome-based phylogeny and comprehensive morphological data. BMC Evolutionary Biology, 14 (1), 52.
https://doi.org/10.1186/1471-2148-14-52
Ponomarenko, A.G. (1968) Archostematan beetles from the Jurassic of Karatau (Coleoptera, Archostemata). In: Rohdendorf, B.B. (Ed.), Jurassic Insects of Karatau. Nauka, Moscow,pp. 118–138. [In Russian.].
Ponomarenko, A.G. (1969) The historical development of archostematan beetles. Trudy Paleontologicheskogo Instituta Akademii Nauk SSSR, 125, 1–238 [In Russian].
Ponomarenko, A.G. (1977) Suborder Adephaga. In: Arnoldy, L.V., Jerikin, V.V., Nikritin, L.M. & Ponomarenko, A.G. (Eds.), Historical development of archostematan beetles Trudy Paleontologicheskogo Instituta Akademii Nauk SSSR, 161, 3–104.
Popper, K. (1957) Philosophy of science: a personal report. In: Mace, C.A. (Ed.), British Philosophy in the mid-Century. George Allen and Unwin, London, pp. 182–183.
Prokin, A.A. (2017) New data about evolution of aquatic beetles (Coleoptera). XV Congress of the Russian Entomological Society, Novosibirsk. Materials of the Congress, 576 pp.
Vasilikopoulos, A., Balke, M., Beutel, R.G., Donath, A., Podsiadlowski, L., Pflug, J.M., Waterhouse, R.M., Meusemann, K., Peters, R.S., Escalona, H.E., Mayer, Ch., Liu, Sh., Hendrich, L., Alarie, Y., Bilton, D.T., Jia, F., Zhou, X., Maddison, D.R., Niehuis, O. & Misof, B. (2019) Phylogenomics of the superfamily Dytiscoidea (Coleoptera: Adephaga) with an evaluation of phylogenetic conflict and systematic error. Molecular Phylogenetics and Evolution, 135, 270–285.
https://doi.org/10.1016/j.ympev.2019.02.022
Volkov, A.N. (2013) New species of Triaplidae from the Babii Kamen’ locality (Kuznetsk Basin). Paleontological Journal, 47, 94–97.
https://doi.org/10.1134/S0031030113010140
Yan, E.V., Beutel, R.G. & Ponomarenko, A.G. (2017a) †Ademosynidae (Insecta: Coleoptera)—a new concept for a coleopteran key taxon and its phylogenetic affinities to the extant suborders. Paleontologia Electronica, 20.2.31A.
Yan, E.V., Lawrence, J.F., Beattie, R. & Beutel, R.G. (2017b) At the dawn of the great rise: †Ponomarenkia belmonthensis (Insecta: Ceoleoptera), a new remarkable late Permian beetle from the Southern hemisphere. Journal of Systematic Paleontology, 16 (7), 611–619.
https://doi.org/10.1080/14772019.2017.1343259
Yan, E.V., Beutel, R.G. & Lawrence, J.F. (2018a) Whirling in the late Permian: ancestral Gyrinidae show early radiation of beetles before Permian-Triassic mass extinction. BMC Evolutionary Biology, 18 (1), 33.
https://doi.org/10.1186/s12862-018-1139-8
Yan, E.V., Beutel, R.G., Beattie, R. & Lawrence, J.F. (2018b) Ponomarenkium gen. nov., a replacement name for the stem group beetle Ponomarenkia Yan et al., 2017 (Insecta: Coleoptera). Paleontological Journal, 52 (2), 219.