Eur. J. Entomol. 101 (1): 95-144, 2004 | DOI: 10.14411/eje.2004.018

Relationships among coleopteran suborders and major endoneopteran lineages: Evidence from hind wing characters

Jarmila KUKALOVÁ-PECK1, John F. LAWRENCE2,3
1 Earth Sciences, Carleton University, Ottawa, ON K1S 5B6, Canada; e-mail: JarmilaPeck@pigeon.carleton.ca
2 CSIRO Entomology, GPO Box 1700, Canberra, ACT 2601, Australia
3 130 Hartwig Road, Gympie, QLD 4570, Australia; e-mail: beetle@spiderweb.com.au

A phylogenetic analysis of the four coleopteran suborders (Polyphaga, Archostemata, Myxophaga and Adephaga), four other endoneopteran taxa (Strepsiptera, Neuropterida, Mecopterida and Hymenoptera) and three neopteran outgroups (Orthoneoptera, Blattoneoptera and Hemineoptera) is performed based on 63 characters of hind wing venation, articulation and folding patterns, with character states coded for the groundplan of each taxon (not for exemplar genera or species). The shortest tree found using Winclada with Nona exhibits the following topology: Orthoneoptera + (Blattoneoptera + (Hemineoptera + Endoneoptera: (Hymenoptera + ((Neuropterida + Mecopterida) + (Coleoptera + Strepsiptera))))). Homologization of the hind wing venation in Coleoptera is reviewed and updated, and comments are made concerning recent works on wing folding. Recent phylogenetic schemes proposed for the orders of Endoneoptera and suborders of Coleoptera are reviewed and their supporting evidence critically examined. The special role and influence of the hind wing anojugal lobe on the diversification of Neoptera and Endoneoptera is discussed. A scenario is proposed for the origin and evolution of the insect hind wing.

Keywords: Coleoptera suborders, Endopterygota, Neoptera, Endoneoptera lineages, insect wings, phylogeny, morphology

Received: August 19, 2002; Revised: October 9, 2003; Accepted: November 5, 2003; Published: March 25, 2004  Show citation

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KUKALOVÁ-PECK, J., & LAWRENCE, J.F. (2004). Relationships among coleopteran suborders and major endoneopteran lineages: Evidence from hind wing characters. EJE101(1), 95-144. doi: 10.14411/eje.2004.018
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References

  1. Abouheif E. & Wray G.A. 2002: Evolution of the gene network underlying wing polyphenism in ants. Science 297: 249-252 Go to original source...
  2. Averof M. & Cohen M. 1997: Evolutionary origin of insect wings from ancestral gills. Nature 385: 627-630 Go to original source...
  3. Baehr M. 1975: Skelett und Muskulatur des Thorax von Priacma serrata LeConte (Coleoptera, Cupedidae). Z. Morphol. Tiere 81: 55-101 Go to original source...
  4. Beutel M. & Gorb S.N. 2001: Ultrastructure of attachment specializations of hexapods (Arthropoda): evolutionary patterns inferred from a revised ordinal phylogeny. J. Zool. Syst. Evol. Res. 39: 177-207 Go to original source...
  5. Beutel R. & Haas F. 1998: Verwandtschaftsbeziehungen der Unterordnungen der Coleoptera (Insecta). In: Systematik im Aufbruch. 1. Jahrestagung der Gesellschaft fuer Biologische Systematik e. V. Zoologisches Forschungsinstitut und Museum Alexander Koenig, Bonn, pp. 22-23
  6. Beutel R. & Haas F. 2000: Phylogenetic relationships of the suborders of Coleoptera (Insecta). Cladistics 16: 103-141 Go to original source...
  7. Bocharova-Messner O.M. 1959: Development of the wing in the early postembryonic stages in the ontogeny of dragonflies (order Odonata). Trudy Inst. Morfol. Zhivot. Im. Severtsova 27: 187-200 [in Russian]
  8. Boudreaux H.B. 1979: Arthropod Phylogeny, with Special Reference to Insects. John Wiley & Sons, New York, viii + 320 pp
  9. Brodsky A.K: 1994: The Evolution of Insect Flight. Oxford University Press, Oxford, 229 pp. Buening J. 1998: The ovariole: structure, type and phylogeny. In: Harrison F.W. & Locke H. (eds): Microscopic Anatomy of Invertebrates 11C, Insecta. Wiley-Liss, New York, pp. 897-932
  10. Buening J. & Maddison D.R. 1998: Surprising ovary structures at the base of Coleoptera. In: Brunnhofer V. & Soldan T. (eds): Book of Abstracts, VIth European Congress of Entomology. Institute of Entomology, Ceske Budejovice, p. 182
  11. Carmean D. & Crespi B.J. 1995: Do long branches attract flies? Nature 373: 666 Go to original source...
  12. Carmean D., Kimsey L.S. & Berbee M.L. 1992: 18S rDNA sequences and holometabolous insects. Mol. Phylogenet. Evol. 1: 270-278 Go to original source...
  13. Carpenter F.M. 1992: Treatise on Invertebrate Palaeontology, Part R, Arthropoda 4. The Geological Society of America, Boulder, Colorado, 655 pp
  14. Caterino M.S., Shull V.L., Hammond P.M. & Vogler A.P. 2002: Basal relationships of Coleoptera inferred from 18S rDNA sequences. Zool. Scripta 31: 41-49 Go to original source...
  15. Chadwick L.E. 1959: Spinasternal musculature in certain insect orders. Smiths. Misc. Collect. 137: 117-156
  16. Chalwatzis N., Hauf J., van de Peer Y., Kinzelbach R. & Zimmermann F.K. 1996: 18S ribosomal RNA genes of insects: primary structure of the genes and molecular phylogeny of the Holometabola. Ann. Entomol. Soc. Am. 89: 788-803 Go to original source...
  17. Crowson R.A. 1955: The Natural Classification of the Families of Coleoptera. Nathaniel Lloyd, London, 187 pp
  18. Crowson R.A. 1960: The phylogeny of Coleoptera. Annu. Rev. Entomol. 5: 111-134 Go to original source...
  19. Crowson R.A. 1975: The evolutionary history of Coleoptera, as documented by fossil and comparative evidence. Atti del Congresso Nazionale Italiano di Entomologia, Sassari - 20-25 Maggio 1974, Firenze, Italy, pp. 47-90
  20. Forbes W.T.M. 1922: The wing-venation of the Coleoptera. Ann. Entomol. Soc. Am. 15: 328-345, pls. xxiv-xxv
  21. Garcia-Bellido A.P. 1975: Genetic control of wing disc development in Drosophila. In: Cell Patterning. Ciba Foundation Symposium No. 29, pp. 161-182 Go to original source...
  22. Goloboff P.A. 1999: NONA ver. 2.0. P.A. Goloboff, Tucuman, Argentina. Haas F. & Beutel R.G: 2001. Wing folding and the functional morphology of the wing base in Coleoptera. Zoology 104: 123-141 Go to original source...
  23. Haas F. & Kukalova-Peck J. 2001: Dermaptera hindwing structure and folding: new evidence for familial, ordinal and superordinal relationships within Neoptera (Insecta). Eur. J. Entomol. 98: 445-509 Go to original source...
  24. Haas F. & Wootton R.J. 1996: Two basic mechanisms in insect wing folding. Proc. R. Entomol. Soc. London 263: 1651-1658 Go to original source...
  25. Haas F., Gorb S. & Blickhan R. 2000: The function of resilin in beetle wings. Proc. R. Entomol. Soc. London 267: 1375-1381 Go to original source...
  26. Hamilton K.G.A. 1972: The insect wing, part II. Vein homology and the archetypal insect wing. J. Kansas Entomol. Soc. 45: 54-58
  27. Hennig W. 1969: Die Stammesgeschichte der Insekten. Waldemar Kramer, Frankfurt am Main, 436 pp
  28. Hennig W. 1981: Insect Phylogeny. John Wiley, New York, 514 pp
  29. Hoernschemeyer T. 1998: Morphologie und Evolution des Fluegelgelenks der Coleoptera und Neuropterida. Bonner Zool. Monogr. 43: 1-126
  30. Hoernschemeyer T. 2002: Phylogenetic significance of the wing-base of the Holometabola (Insecta). Zool. Scripta 31: 17-29 Go to original source...
  31. Huelsenbeck J.P. 1997: Is the Felstenstein zone a fly trap? Syst. Biol. 46: 69-74 Go to original source...
  32. Huelsenbeck J.P. 1998: Systematic bias in phylogenetic analysis: is the Strepsiptera problem solved? System. Biol. 47: 519-537
  33. Huelsenbeck J.P: 2001: A Bayesian perspective on the Strepsiptera problem. Tijdschr. Entomol. 144: 165-178
  34. Hwang U.W., Kim W., Tautz D. & Friedrich M. 1998: Molecular phylogenetics at the Felsenstein Zone: approaching the Strepsiptera problem using 5.85 and 28S rDNA sequences. Mol. Phylogenet. Evol. 9: 470-480 Go to original source...
  35. Kinzelbach R.K. 1971: Morphologische Befunden an Faecherfluegeln und ihre phylogenetische Bedeutung (Insecta: Strepsiptera). Zoologica 41: 1-256
  36. Kinzelbach R.K. 1990: The systematic position of Strepsiptera (Insecta). Am. Entomol. 36: 292-303 Go to original source...
  37. Kirejtshuk A.G. 1999: Sikhotealinia zhiltzovae (Lafer, 1996) - recent representative of the Jurassic coleopterous fauna (Coleoptera, Archostemata, Jurodidae). Proc. Zool. Inst. Russian Acad. Sci. 281: 21-26
  38. Klausnitzer B. 1975: Probleme der Abgrenzung von Unterordnungen bei den Coleoptera. Entomol. Abh. Staatl. Mus. Tierkd. Dresden 40: 269-275 Go to original source...
  39. Koenigsmann E. 1976: Das phylogenetische System der Hymenoptera I. Dt. Entomol. Z. (N.F.) 23: 253-279 Go to original source...
  40. Kristensen N.P. 1975: The phylogeny of hexapod "orders". A critical review of recent accounts. Z. Zool. Syst. Evolutionsforsch. 13: 1-44 Go to original source...
  41. Kristensen N.P. 1981: Phylogeny of insect orders. Annu. Rev. Entomol. 26: 135-157 Go to original source...
  42. Kristensen N.P. 1991: Phylogeny of extant hexapods. In: CSIRO (ed.): The Insects of Australia. A Textbook for Students and Research Workers. Second Edition, Vol. 1. Melbourne University Press, Carlton, pp. 125-140
  43. Kristensen N.P. 1995: Forty years' insects phylogenetic systematics. Zool. Beitr. (N. F.) 36: 83-124
  44. Kristensen N.P. 1997: The groundplan and basal diversification of the hexapods. In: Fortey R.A. & Thomas R.H. (eds): Arthropod Relationships. Chapman & Hall, London, pp. 281-293 Go to original source...
  45. Kristensen N.P. 1999: Phylogeny of endopterygote insects, the most successful lineage of living organisms. Eur. J. Entomol. 96: 237-253
  46. Kukalova-Peck J. 1978: Origin and evolution of insect wings and their relation to metamorphosis, as documented by the fossil record. J. Morphol. 156: 53-126 Go to original source...
  47. Kukalova-Peck J. 1983: Origin of the insect wing and wing articulation from the arthropodan leg. Can. J. Zool. 61: 1618-1669 Go to original source...
  48. Kukalova-Peck J. 1987: New Carboniferous Diplura, Monura, and Thysanura, the hexapod ground plan, and the role of the thoracic side lobes in the origin of wings (Insecta). Can. J. Zool. 65: 2237-2345 Go to original source...
  49. Kukalova-Peck J. 1991: Fossil history and the evolution of hexapod structures. In: C.S.I.R.O. (ed.): Insects of Australia. A Textbook for Students and Research Workers. Second Edition, Vol. 1. Melbourne University Press, Carlton, pp. 141-179
  50. Kukalova-Peck J. 1997: Arthropod phylogeny and "basal" morphological structures. In: Fortey R.A. & Thomas R.H. (eds): Arthropod Relationships. Chapman & Hall, London, pp. 249-268 Go to original source...
  51. Kukalova-Peck J. & Brauckmann C. 1990: Wing folding in pterygote insects, and the oldest Diaphanopterodea from the early Late Carboniferous of West Germany. Can. J. Zool. 68: 1104-1111 Go to original source...
  52. Kukalova-Peck J. & Lawrence J.F. 1993: Evolution of the hind wing in Coleoptera. Can. J. Zool. 125: 181-258 Go to original source...
  53. Kukalova-Peck J. & Richardson E.S. 1983: New Homoiopteridae (Insecta: Paleodictyoptera) with wing articulation from the Upper Carboniferous strata of Mazon Creek, Illinois. Can. J. Zool. 61: 1670-1687 Go to original source...
  54. Lafer G.Sh. 1996: Family Sikhotealiniidae. In: Ler P.A. (ed.): Key to the Insects of the Russian Far East. Vol. III, Part 3. Dal'nauka, Vladivostok, pp. 390-396
  55. Larsen O. 1966: On the morphology and function of the locomotor organs of the Gyrinidae and other Coleoptera. Opusc. Entomol. (Suppl.) 30: 1-242
  56. Lawrence J.F. 1989: Mycophagy in the Coleoptera: feeding strategies and morphological adaptations. In: Wilding N., Collins M., Hammond P.M. & Webber J.F. (eds): Insect-Fungus Interactions. R. Entomol. Soc. Symp. Ser. 14, pp. 1-23 Go to original source...
  57. Lawrence J.F. 1991: Order Coleoptera.. In: Stehr F.W. (ed.): Immature Insects. Vol. II. Kendall-Hunt, Dubuque, Iowa, pp. 144-298
  58. Lawrence J.F. 1999: The Australian Ommatidae (Coleoptera: Archostemata): new species, larva and discussion of relationships. Invertebr. Taxon. 13: 369-390 Go to original source...
  59. Lawrence J.F. 2001: A new genus of Valdivian Scirtidae (Coleoptera) with comments on Scirtoidea and the beetle suborders. In: Morimoto K. et al. (eds): Sukunahikona. Special Publication of the Japan Coleopterological Society No. 1. Japan Coleopterological Society, Osaka, pp. 351-361
  60. Lawrence J.F. & Newton A.F. Jr. 1995: Families and subfamilies of Coleoptera (with selected genera, notes and references, and data on family-group names). In: Pakaluk J. & Slipinski S.A. (eds): Biology, Phylogeny, and Classification of Coleoptera: Papers Celebrating the 80th Birthday of Roy A. Crowson. Muzeum i Instytut Zoologii, Polska Akademia Nauk, Warsaw, pp. 779-1006
  61. Lawrence J.F., Slipinski S.A. & Pakaluk J. 1995: From Latreille to Crowson: a history of the higher-level classification of beetles. In: Pakaluk J. & Slipinski S.A. (eds): Biology, Phylogeny, and Classification of Coleoptera: Papers Celebrating the 80th Birthday of Roy A. Crowson. Muzeum i Instytut Zoologii, Polska Akademia Nauk, Warsaw, pp. 87-154
  62. Lawrence J.F., Hastings A.M., Dallwitz M.J., Paine T.A. & Zurcher E.J. 1999: Beetles of the World: a Key and Information System for Families and Subfamilies. CD-ROM, version 1.0 for MS-Windows. C.S.I.R.O. Publishing, East Melbourne. Leschen R.A.B. 2003: Erotylidae (Insecta: Coleoptera: Cucujoidea): phylogeny and review. Fauna of New Zealand 47: 1-108
  63. Maddison D.R., Baker M.D. & Ober K.A. 1999: Phylogeny of carabid beetles as inferred from 18S ribosomal DNA. Syst. Entomol. 24: 103-138 Go to original source...
  64. Manton S.M. 1997: The Arthropoda. Clarendon Press, Oxford, 527 pp
  65. Martynov A.V. 1923: On two basic types of insect wings and their significance for the general classification of insects. In: Deryugina K.M. (ed.): Trudy Perv. Vserossij. S'ezda Zool., Anatomov i Gistologov 1: 88-89
  66. Matsuda R. 1970: Morphology and evolution of the insect thorax. Mem. Entomol. Soc. Can. 76: 1-431 Go to original source...
  67. Mickoleit G. 1971: Das Exoskelett von Notiothauma reedi McLachlan, ein Beitrag zur Morphologie und Phylogenie der Mecoptera (Insecta). Z. Morphol. Tiere 69: 318-362 Go to original source...
  68. Mickoleit G. 1973: Uber den Ovipositor der Neuropteroidea und Coleoptera und seine phylogenetische Bedeutung (Insecta, Holometabola). Z. Morphol. Tiere 76: 145-171 Go to original source...
  69. Nixon K.C. 1999: Winclada (BETA) ver. 0.9.9. K.C. Nixon, Ithaca, New York. Pashley D.P., McPheron B.A. & Zimmer E.A. 1993: Systematics of holometabolous insect orders based on 18S ribosomal RNA. Mol. Phylogenet. Evol. 2: 132-142 Go to original source...
  70. Ponomarenko A.G. 1969: The historical development of archostematan beetles. Trudy Paleontol. Inst. Akad. Nauk SSSR 125: 1-238 [in Russian]
  71. Ponomarenko A.G. 1985: Beetles from the Jurassic of Siberia and West Mongolia. In: Rasnitsyn A.P. (ed.): Jurassic Insects from Siberia and West Mongolia. Trudy Paleontol. Inst. Akad. Nauk SSSR 211: 47-87 [In Russian]
  72. Ponomarenko A.G. 1995: The geological history of beetles. In: Pakaluk J. & Slipinski S.A. (eds): Biology, Phylogeny, and Classification of Coleoptera: Papers Celebrating the 80th Birthday of Roy A. Crowson. Muzeum i Instytut Zoologii, Polska Akademia Nauk, Warsaw, pp. 155-171
  73. Ponomarenko A. G. 2002: 2.2.1.3.2. Superorder Scarabaeidea Laicharting, 1781. Order Coleoptera Linne, 1858. The Beetles. In: Rasnitsyn A.P. & Quicke D.L.J. (eds.): History of Insects. Kluwer Academic Publishers, Dordrecht, pp. 164-176
  74. Rohdendorf B.B. & Ponomarenko A.G. 1962: Order Coleoptera. Beetles. In: Rohdendorf B.B. (ed.): Fundamentals of Paleontology. Volume 9. Arthropoda, Tracheata, Chelicerata. Akademia Nauk SSSR, Moscow, pp. 241-267 [in Russian; translation 1991, D.R. Davis (ed.), Smithsonian Institution and National Science Foundation, Washington, D.C.]
  75. Rohdendorf B.B. & Rasnitsyn A.P. (eds) 1980: Historical development of the class Insecta. Trudy Paleontol. Inst. Akad. Nauk SSSR 175: 1-199
  76. Ross H.H. 1965: A Textbook of Entomology. 3rd Edition. John Wiley & Sons, New York, ix + 539 pp
  77. Schneider P. 1978: Die Flug- und Faltungstypen der Kaefer (Coleoptera). Zool. Jb. Anat. Ontog. Tiere 99: 174-210
  78. Sharov A.G. 1966: Basic Arthropodan Stock. Pergamon Press, Elmsford, N.Y., 271 pp
  79. Shear W.A. & Kukalova-Peck J. 1990: The ecology of Paleozoic terrestrial arthropods: the fossil evidence. Can. J. Zool. 68: 1807-1834 Go to original source...
  80. Shubin N., Tabin C. & Carroll S. 1997: Fossils, genes and the evolution of animal limbs. Nature 388: 639-648 Go to original source...
  81. Shull V.L., Vogler A.P., Baker M.D., Maddison D.R. & Hammond P.M. 2001: Sequence alignment of 18S ribosomal RNA and the basal relationships of adephagan beetles: evidence for monophyly of aquatic families and the placement of Trachypachidae. Syst. Biol. 50: 945-969 Go to original source...
  82. Siddall M.E. 1998: Success of parsimony in the four-taxon case: long-branch repulsion by likelihood in the Farris zone. Cladistics 14: 209-220 Go to original source...
  83. Siddall M.E. & Whiting M.F. 1999: Long-branch abstractions. Cladistics 15: 9-24 Go to original source...
  84. Snodgrass R.E. 1935: Principles of Insect Morphology. McGraw-Hill, New York, x + 667 pp
  85. Stenzhorn H.J. 1974: Experimentelle Untersuchungen zur Entwicklung des Lymantria dispar L. (Lepidoptera). Wilhelm Roux Arch. EntwMech. Org. 175: 65-86 Go to original source...
  86. Sulc K. 1927: Das Tracheensystem von Lepisma (Thysanura) und Phylogenie der Pterygogenea. Acta Soc. Sci. Nat. Moravicae 4(7), 108 pp
  87. Tower W.L. 1903: The origin and development of the wings of Coleoptera. Zool. Jb. Anat. 17: 517-572 Go to original source...
  88. Vilhelmsen L. 1996: The preoral cavity of lower Hymenoptera (Insecta): comparative morphology and phylogenetic significance. Zool. Scripta 25: 143-170 Go to original source...
  89. Vilhelmsen L. 2001: Phylogeny and classification of the extant basal lineages of the Hymenoptera (Insecta). Zool. J. Linn. Soc. 131: 393-442 Go to original source...
  90. Wallace F.L. & Fox R.C. 1980: A comparative morphological study of the hindwing venation of the order Coleoptera, Part II. Proc. Entomol. Soc. Wash. 82: 609-654
  91. Ward R.D. 1979: Metathoracic wing structures as phylogenetic indicators in the Adephaga (Coleoptera). In: Erwin T.L., Ball G.E., Whitehead D.R. & Halpern A.L. (eds): Carabid Beetles: Their Evolution, Natural History, and Classification. W. Junk, The Hague, pp. 113-180 Go to original source...
  92. Wheeler W.C., Whiting M., Wheeler Q.D. & Carpenter J.M. 2001: The phylogeny of the extant hexapod orders. Cladistics 17: 113-169 Go to original source...
  93. Whiting M.F. 1998: Phylogenetic position of the Strepsiptera: review of molecular and morphological evidence. Int. J. Insect Morphol. Embryol. 27: 53-60 Go to original source...
  94. Whiting M.F. 2002: Phylogeny of the holometabolous insect orders: molecular evidence. Zool. Scripta 31: 3-15 Go to original source...
  95. Whiting M.F. & Kathirithamby J. 1995: Strepsiptera do not share hind wing venational synapomorphies with Coleoptera: a reply to Kukalova-Peck and Lawrence. J. N. Y. Entomol. Soc. 103: 1-14
  96. Whiting M.F. & Wheeler W.C. 1994: Insect homeotic transformation. Nature 368: 696 Go to original source...
  97. Whiting M.F., Carpenter J.C., Wheeler Q.D. & Wheeler W.C. 1997: The Strepsiptera problem: phylogeny of the holometabolous insect orders inferred from 18S and 28S ribosomal DNA sequences and morphology. Syst. Biol. 46: 1-68 Go to original source...
  98. Willmann R. 1994: Raphidiodea aus dem Lias und die Phylogenie der Kamelhalsfliegen (Insecta: Holometabola). Palaeontol. Z. 68: 167-197 Go to original source...
  99. Willmann R. 1997: Advances and problems in insect phylogeny. In: Fortey R.A. & Thomas R.H. (eds): Arthropod Relationships. Chapman & Hall, London, pp. 269-279 Go to original source...
  100. Wootton R.J. 1976: The fossil record and insect flight. In: Rainey R.C. (ed.): Insect Flight. R. Entomol. Soc. London Symp. Ser. 7, pp. 235-254
  101. Wootton R.J. 1979: Function, homology and terminology in insect wings. Syst. Entomol. 4: 81-93 Go to original source...
  102. Wootton R.J. 1981: Support and deformability in insect wings. J. Zool. (London) 193: 447-468 Go to original source...
  103. Wootton R.J. 1990: The mechanical design of insect wings. Sci. Am. 262: 114-120 Go to original source...
  104. Wootton R.J. 1992: Functional morphology of insect wings. Annu. Rev. Entomol. 37: 113-140 Go to original source...
  105. Wootton R.J. 1995: Geometry and mechanics of insect hind wing fans: a modelling approach. Proc. R. Soc. London (B) 262: 181-187 Go to original source...
  106. Wootton R.J. 1996: Functional wing morphology in Hemiptera systematics. In: Schaefer C.W. (ed.): Studies on Hemipteran Phylogeny. Entomological Society of America, Lanham, Maryland, pp. 179-198 Go to original source...
  107. Wootton R.J. & Kukalova-Peck J. 2000: Flight adaptation in Palaeozoic Palaeoptera (Insecta). Biol. Rev. 75: 129-167 Go to original source...
  108. Zherikhin V.V. & Gratshev V.G. 1995: A comparative study of the hind wing venation of the superfamily Curculionoidea, with phylogenetic implications. In: Pakaluk J. & Slipinski S.A. (eds): Biology, Phylogeny, and Classification of Coleoptera: Papers Celebrating the 80th Birthday of Roy A. Crowson. Muzeum i Instytut Zoologii, Polska Akademia Nauk, Warsaw, pp. 633-777. APPENDIX 1. PARTIAL LISTS OF GENERA EXAMINED

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