Abstract
Most currently applied systematic methods use post-groundplan character states to reconstruct phylogenies in modern higher Insecta/Arthropoda taxa. But, this approach is unable to separate synapomorphies from frequently occurring homoplasies. Conflicting, unresolved and unrealistic higher-level phylogenies result. The reasons are analyzed. A contrasting “groundplan” method, long used in Vertebrata and found to be superior in resolving higher-level phylogenies, is described. This method, as used for insects, uses a highly diversified morphological organ system (such as limb/wing), identifies its homologues in all subphyla and classes, records the full history of its character transformation series in all lineages from the shared Paleozoic ancestor to modern times, pursues the full homologization of its character states in all modern orders, and verifies these data with evidence from other fields of biology. Only such an extremely broad dataset provides the complex information needed to identify and homologize the groundplan character states in modern orders and other higher taxa in the insect/arthropod fauna. After this is accomplished, the gate to recognizing higher-level synapomorphies is open. Only groundplan-level character states include distinct synapomorphies, since homoplasies are either absent or easily detectable. Examples are given. The interpretations of higher phylogenies and evolutionary processes in Hexapoda, based on the unpredictable and often misleading post-groundplan character states found in extant, Tertiary and Mesozoic fauna, are critically compared with those based on the evolution of organ systems, by using the groundplan method.
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Acknowledgements
Broad evolutionary-morphological contributions in insects are quite impossible without selfless and massive help from systematists specializing in individual orders. I am deeply obliged to many entomologists who, over the years, by their suggestions and their wonderful insect collections, have helped to clarify for me the theoretical and practical aspects of the groundplan method. This includes my past and present co-authors and friends, who patiently shared with me their extensive knowledge of modern and fossil entomofaunas. This essay would never be possible without the profound morphological analysis of arthropods by Edward Laidlaw Smith (The California Academy of Sciences, San Francisco), which I had the privilege to witness and participate in, and without my field assistant Karel Havlata, who organized the fossil excavations in Moravia. Because of my politically challenged origins, my international background in Paleozoic insects was possible only with the most generous support of Frank M. Carpenter, Harvard University, Cambridge, and Ernst Mayr, Museum of Comparative Zoology, Cambridge. With their grant I also studied, over multiple prolonged visits, all Paleozoic insects deposited at the Muséum d’Histoire Naturelle, Paris, at the Palentological Institute, Moscow (with B. B. Rohdendorf and A. G. Sharov), and at the Museum of Natural History, London. My work in Canada was supported by the Canadian NSERC grant agency, and by Stewart B. Peck, Carleton University, Ottawa. The grants made it possible for me to study extensively Paleozoic insects in Russia, Germany, on many occasions at the Field Museum, Chicago (with E. S. Richardson) and in numerous private collections in the Chicago area. Fossils and rare primitive hemipteroids were researched at The Australian Museum, Sydney (Australia), and at the Natal Museum, Pietermaritzburg (South Africa). Modern insects were collected extensively in North, Central, and South America, including Alaska, Yukon, Tierra del Fuego, Galápagos Islands, Puerto Rico, Dominican Republic, Jamaica, Cuba, Lesser Antilles, South Africa and Ghana, all around Australia, New Zealand, New Caledonia, Fiji, Lord Howe Island, Japan, and China. I studied extensively in several months-long visits, modern Ephemeroptera at the Florida A&M University, Tallahassee (with J. & J. Peters) and at the Czech Academy of Sciences, České Budějovice (Czech Republic) (with T. Soldán), modern Coleoptera (with J. F. Lawrence), orthopteroids (with D. Rentz) at CSIRO, Canberra, and orthopteroids, blattoids and Dermaptera at the Academy of Sciences, Philadelphia (with D. Otte and F. Haas). Orthopteroids and tropical Hemipteroids were studied in Prague (with P. Štys), at INBIO, Santo Domingo, Costa Rica, and at the Territorial Museum, Darwin, peloridiids in Sydney (Australia) (with J. Evans), DSIR Auckland (New Zealand) (with W. Kuschel), and in the Bishop Museum, Honolulu (with S. Miller). At the Smithonian Institution, Washington, D.C., I studied Mecoptera and Trichoptera (with O. Flint), orthopteroids (with D. Nickles), and Carboniferous insects (with C. Labandeira). Plecoptera were researched at the Max Planck Institute, Schlitz, Germany (with P. Zwick), Embioptera at the California Academy of Sciences, San Francisco (with E.S. Ross), and Neuroptera and Megaloptera mainly in Ottawa (on material delivered by K. Lambkin). Hymenoptera were researched in the collection of Agriculture, Canada, Ottawa (with M. Sharkey and G. Gibson). My friend Jan G. Peters, Florida A&M University, Tallahassee devoted many hours to reviewing this manuscript. G. S. Ball and B. S. Heming, University of Alberta, Edmonton, R.G. Beutel, Friedrich Schiller Universität, Jena, and V.A. Grebenikov, Agriculture and Agri-Food Canada, Ottawa provided critical remarks and important pointers in contemporary systematic methods. From G. Bechly, Museum für Naturkunde, Stuttgart, I received useful suggestions clarifying some terms. I also received invaluable editorial help from B. Hallgrimsson, and from two reviewers who devoted generously their time to improve the presentation of arguments. To all these colleagues, co-authors and friends, and others too many to be named, goes my sincere gratitude and thanks.
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Kukalová-Peck, J. Phylogeny of Higher Taxa in Insecta: Finding Synapomorphies in the Extant Fauna and Separating Them from Homoplasies. Evol Biol 35, 4–51 (2008). https://doi.org/10.1007/s11692-007-9013-4
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DOI: https://doi.org/10.1007/s11692-007-9013-4