Skip to main content Skip to main navigation menu Skip to site footer
Article
Published: 2017-10-03

Cleonini (Coleoptera: Curculionidae: Lixinae) are monophyletic and flightless: tribe overview, rampant adult homoplasy and illustrated global diversity

apt. 36, bul’var Komarova 13/1, Rostov-on-Don, 344092, Russia.
CFIA, 960 Carling Ave., Ottawa, ON, K1A 0C6, Canada.
Coleoptera Lixini Hennigian comb beet pests

Abstract

We summarize knowledge of the weevil tribe Cleonini worldwide, including its monophyly, relationships, distribution, biology, immature stages, economic significance and paleontology. We score adult morphological characters for 79 of a total of 96 extant genus-group Cleonini taxa considered valid to date. The resulting matrix contains 121 parsimoniously informative characters scored for 145 ingroup (Cleonini) and 29 outgroup terminals. Maximum Parsimony (MP) and Bayesian Inference (BI) analyses consistently recover monophyletic Lixinae and Cleonini. Relationships within the latter remain unresolved with either 47 (BI) or 37 (MP) branches radiating from the tribe’s most recent common ancestor. Most of the speciose genera of Cleonini emerge as monophyletic in both BI and MP analyses (generic names followed by the number of terminals, then by BI posterior probability / MP bootstrap): Adosomus (5, 94/77), Asproparthenis (6, 99/98), Chromonotus (6, 98/85), Cleonis (3, 64/76), Coniocleonus (10, 95/41), Conorhynchus (5, 95/51), Cyphoclenus (4, 65/76), Maximus (4, 84/68), Mecaspis (4, 95/91), Scaphomorphus (4, 90/84), Temnorhinus (8, 99/62) and Xanthochelus (6, 84/71). The genera Pseudocleonus (6, -/26) and Stephanocleonus (22, -/23) are not recovered in BI and weakly supported in MP. No genera are here added to, or removed from, Cleonini. We suggest that adult morphology of Cleonini was subject to widespread homoplasy obscuring the phylogenetic signal of morphological characters. Unlike the rest of Lixinae, all extant Cleonini are hypothesised to be flightless, even though often being macropterous. All 145 ingroup terminals are illustrated in three standard views; images of the type species of 15 of the 17 genus-group taxa that are not represented in our analysis are provided.

 

References

  1. Alonso-Zarazaga, M.A. & Lyal, C.H.C. (1999) A world catalogue of families and genera of Curculionoidea (Insecta: Coleoptera) (excepting Scolytidae and Platypodidae). Entomopraxis, Barcelona, 315 pp.

    Alonso-Zarazaga, M.A. & Lyal, C.H.C. (2002) Addenda and corrigenda to “A World Catalogue of Families and Genera of Curculionoidea (Insecta: Coleoptera)”. Zootaxa, 63 (1), 1–37.

    https://doi.org/10.11646/zoootaxa.63.1.1

    Aslam, N.A. (1963) On the genera of Indo-Pakistan Cleoninae and Hylobiinae (Coleoptera: Curculionidae). Bulletin of the British Museum (Natural History), Entomology, 13, 47–66.

    Anderson, R.S. (1988) Systematics, phylogeny and biogeography of New World weevils traditionally of the tribe Cleonini (Coleoptera: Curculionidae; Cleoninae). Quaestiones Entomologicae, 23, 431–709. [1987]

    Anderson, R.S. (2002) 131. Curculionidae Latreille 1802. In: Arnett, R.H., Thomas, M.T., Skelley, P.E. & Frank, J.H. (Eds.), American Beetles. Polyphaga: Scarabaeoidea through Curculionoidea. Vol. 2. CRC Press, Boca Raton, pp. 722–815.

    Arzanov, Yu. G. (2003) Use of the endophallus characters in the systematics of the Rhynchophorous beetles (Coleoptera, Curculionoidea). Entomological Review, 83, 930–944.

    Arzanov, Yu. G. (2005a) To cognition of systematic of the genus Conorhynchus sensu lato (Coleoptera: Curculionidae: Lixinae). Caucasian Entomological Bulletin, 1, 57–64. [in Russian]

    Arzanov, Yu. G. (2005b) Towards the knowledge of systematic of the tribe Cleonini (sensu lato). 1. Genera Pseudocleonus Chevrolat, 1873, Cleonis Dejean, 1821, Adosomus Faust, 1904, Cyphocleonus Motschulsky, 1860 and Xeradosomus gen. n. (Coleoptera, Curculionidae). Caucasian Entomological Bulletin, 1, 129–149. [in Russian]

    Arzanov, Yu. G. (2005c) Taxonomic notes on weevils of the tribe Cleonini (Coleoptera: Curculionidae: Lixinae). Caucasian Entomological Bulletin, 1, 150. [in Russian]

    Arzanov, Yu. G. (2006a) To the knowledge of weevil of the genus Chromonotus Motschulsky (sensu lato) (Coleoptera: Curculionidae: Lixinae). Proceedings of the Russian Entomological Society, 77, 8–17. [in Russian]

    Arzanov, Yu. G. (2006b) To the knowledge of weevils of the genus Coniocleonus Motschulsky (s. lato). Caucasian Entomological Bulletin, 2, 109–126. [in Russian]

    Arzanov, Yu. G. (2006c) Materials to the study of the tribe Cleonini (Coleoptera: Curculionidae: Lixinae). The genera Asproparthenis Gozis, 1886, Chromosomus Motschulsky, 1860 and Temnorhinus Chevrolat, 1873. Caucasian Entomological Bulletin, 2, 187–198. [in Russian]

    Arzanov, Yu. G. (2006d) Borisocleonus gen. n. – a new genus of weevils from the tribus of Cleonini (Coleoptera: Curculionidae: Lixinae). Russian Entomological Journal, 15, 63–66. [in Russian]

    Arzanov, Yu. G. (2006e) To the knowledge of weevils of the tribe Cleonini (sensu lato) with coneform rostrum. Reports of the Southern Scientific Centre of Russian Academy of Sciences, 2, 82–85. [in Russian]

    Arzanov, Yu. G. (2007a) Features of the morphological structure of rostrum, and trophic preferences of weevils of the subfamily Lixinae (Coleoptera, Curculionidae). Caucasian Entomological Bulletin, 3, 51–62. [in Russian]

    Arzanov, Yu. G. (2007b) The genus Ephimeronotus Faust, 1904 and its systematic position in tribe Cleonini (Coleoptera: Curculionidae: Lixinae). Reports of the Southern Scientific Centre of Russian Academy of Sciences, 3, 101–102. [in Russian]

    Arzanov, Yu. G. (2008a) Position of the genus Leucophyes Marshall, 1946 in system of the weevils tribe Cleonini sensu lato (Coleoptera: Curculionidae: Lixinae) with the description of new species Leucophyes from East India. Caucasian Entomological Bulletin, 4, 93–99. [in Russian]

    Arzanov, Yu. G. (2008b) Some morphological patterns of the superfamily Curculionoidea (Coleoptera) Caucasian Entomological Bulletin, 4, 337–347. [in Russian]

    Arzanov, Yu. G. (2009a) A new genus of the weevil subfamily Lixinae (Coleoptera: Curculionidae) from Arabian Peninsula. Caucasian Entomological Bulletin, 5, 253–255. [in Russian]

    Arzanov, Yu. G. (2009b) Position and system subfamily Lixinae in family Curculionidae (Coleoptera). Reports of the Southern Scientific Centre of Russian Academy of Sciences, 5, 102–111. [in Russian]

    Arzanov, Yu. G. (2014) Review of Palaearctic species of the genus Ammocleonus Bedel, 1907 (Coleoptera: Curculionidae) and place on the genus in the system of weevils of the Cleonini. Caucasian Entomological Bulletin, 10, 261–270. [in Russian]

    Arzanov, Yu. G. & Korotyaev, B.A. (2011) Taxa of Coleoptera described by M.E. Ter-Minassian. Caucasian Entomological Bulletin, 7, 177–182. [in Russian]

    Arzanov, Yu. G. & Stradomsky, B.V. (2015) Experience of phylogenetic reconstruction of the weevil subfamily Lixinae (Coleoptera: Curculionidae) using molecular-genetic analysis. Caucasian Entomological Bulletin, 11, 63–70. [in Russian]

    Arzanov, Yu. G. & Valov, G.V. (2011) Elytral scales of weevils of the tribe Cleonini sensu lato (Coleoptera: Curculionidae: Lixinae) – try of typification. Caucasian Entomological Bulletin, 7, 197–209. [in Russian]

    Ballerio, A. & Grebennikov, V.V. (2016) Rolling into a ball: phylogeny of the Ceratocanthinae (Coleoptera: Hybosoridae) inferred from adult morphology and origin of a unique body enrollment coaptation in terrestrial arthropods. Arthropod Systematics and Phylogeny, 74, 23–52.

    Bouchard, P., Bousquet, Y., Davies, A. & Sikes, D. (2013) Checklist of beetles (Coleoptera) of Canada and Alaska. ZooKeys, 360, 1–44.

    https://doi.org/10.3897/zookeys.360.4742

    Cassis, G. & Schuh, R.T. (2010) Systematic methods, fossils, and relationships within Heteroptera (Insecta). Cladistics, 26, 262–280.

    https://doi.org/10.1111/j.1096-0031.2009.00283.x

    Chevrolat, A. (1873) Mémoire sur les Cléonides. Mémoires de la Société royale des sciences de Liège, 2, viii + 118 pp.

    https://doi.org/10.5962/bhl.title.47131

    Darwin, C. (1859) On the Origin of Species by Means of Natural Selection, or Preservation of Favored Races in the Struggle for Life. John Murray, London, ix + 502 pp.

    Dobzhansky, T. (1937) Genetics and the Origin of Species. Columbia University Press, New York, 364 pp.

    EPPO (2015) Global Database - European and Mediterranean Plant Protection Organization. Curculionidae. Available from: http://gd.eppo.int/taxon/1CURCF (accessed 3 November 2016)

    Farris, J.S. (1989) Hennig86: a PC-DOS program for phylogenetic analysis. Cladistics, 5, 163.

    Faust, J. (1904) Revision der Gruppe Cléonides vrais. Deutsche Entomologische Zeitschrift, 1904, 177–284.
    https://doi.org/10.1002/mmnd.48019040120

    Felsenstein, J. (1985a) Phylogenies and the comparative method. The American Naturalist, 125, 1–15.

    https://doi.org/10.1086/284325

    Felsenstein, J. (1985b) Confidence-limits on phylogenies-an approach using the bootstrap. Evolution, 39, 783–791.

    https://doi.org/10.2307/2408678

    Fikáček, M., Prokin, A.A., Angus, R.B., Ponomarenko, A.G., Yue, Y., Ren, D. & Prokop, J. (2012) Revision of Mesozoic fossils of the helophorid lineage of the superfamily Hydrophiloidea (Coleoptera: Polyphaga). Acta Entomologica Musei Nationalis Pragae, 52, 89–127.

    Gillett, C.P.D.T., Crampton-Platt, A., Timmermans, M.J.T.N., Jordal, B.H., Emerson, B.C. & Vogler, A.P. (2014) Bulk de novo mitogenome assembly from pooled total DNA elucidates the phylogeny of weevils (Coleoptera: Curculionoidea). Molecular Biology and Evolution, 31, 2223–2237.

    https://doi.org/10.1093/molbev/msu154

    Goloboff, P.A. (1999) NONA, V. 2.0. Available from: http://www.cladistics.com/aboutNona.htm (accessed 15 June 2015)

    Goloboff, P.A., Farris, J.S. & Nixon, K.C. (2008) TNT, a free program for phylogenetic analysis. Cladistics, 24, 774–786.

    https://doi.org/10.1111/j.1096-0031.2008.00217.x

    Grebennikov, V.V. (2017) Phylogeography and sister group of Lupangus, a new genus for three new flightless allopatric forest litter weevils endemic to the Eastern Arc Mountains, Tanzania (Coleoptera: Curculionidae, Molytinae). Fragmenta entomologica, 49, 37–55.

    Gültekin, L. (2008) Taxonomic review of the stem-inhabiting trehala-constructing Larinus Dejean, 1821 (Coleoptera: Curculionidae): New species, systematics and ecology. Zootaxa, 1714, 1–18.

    Gültekin, L. (2012) New data on Conorhynchus hololeucus (Coleoptera: Curculionidae). Florida Entomologist, 95, 503–505.

    https://doi.org/10.1653/024.095.0241

    Gültekin, L. & Fremuth, J. (2013) Tribe Lixini. In: Löbl, I. & Smetana, A. (Eds.), Catalogue of Palaearctic Coleoptera. Vol. 8. Curculionoidea II. Brill, Leiden, pp. 456–472.

    Gültekin, L. & Shahreyary-Nejad, S. (2015) A new trehala-constructing Larinus Dejean (Coleoptera: Curculionidae) from Iran. Zoology in the Middle East, 61, 246–251.

    https://doi.org/10.1080/09397140.2015.1069240

    Hooper, G.R. & Holdaway, B.F. (2007) Observations on an association between the weevil Scaphomorphus trivittatus (Say) and two Astragalus species on selenium-rich soils in Carbon County, Utah. Western North American Naturalist, 67, 605–608.

    https://doi.org/10.3398/1527-0904(2007)67[605:ooaabt]2.0.co;2

    Huber, J.T. & Vayssieres, J.F. (1990) Life cycle and host specificity of the Heliotrope weevil, Pachycerus cordiger (=madius auct.) [Col.: Curculionidae]. Entomophaga, 35, 475–484.

    https://doi.org/10.1007/bf02375273

    Hundsdoerfer, A.K., Rheinheimer, J. & Winka, M. (2009) Towards the phylogeny of the Curculionoidea (Coleoptera): Reconstructions from mitochondrial and nuclear ribosomal DNA sequences. Zoologischer Anzeiger, 248, 9–31.

    https://doi.org/10.1016/j.jcz.2008.09.001

    Isart, J. (1972) Observaciones bioecologicas sobre Cleonus mendicus (Gyllanhal, 1834). Graellsia, 28, 177–204.

    Kingsolver, J.M. (1962) Notes on fossil Cleoninae (Coleoptera: Curculionidae). Psyche, 69, 47–49.

    https://doi.org/10.1155/1962/93916

    Kobayashi, C., Okuyama, Y., Kawazoe, K. & Kato, M. (2012) The evolutionary history of maternal plant-manipulation and larval feeding behaviours in attelabid weevils (Coleoptera; Curculionoidea). Molecular Phylogenetics and Evolution, 64, 318–330.

    https://doi.org/10.1016/j.ympev.2012.04.006

    Kojima, H. & Morimoto, K. (2004) An online checklist and database of the Japanese weevils (Insecta: Coleoptera: Curculionoidea) (excepting Scolytidae and Platypodidae). Bulletin of the Kyushu University Museum, 2, 33–147.

    Korotyaev, B.A. (2010) On the 100th Anniversary of Margarita Ervandovna Ter-Minassian. Entomological Review, 90, 951–953.

    https://doi.org/10.1134/s0013873810070110

    Krell, F.T. (2000) The fossil record of Mesozoic and Tertiary Scarabaeoidea (Coleoptera: Polyphaga). Invertebrate Taxonomy, 14, 871–905.
    https://doi.org/10.1071/IT00031

    Kuzmina, S., Elias, S., Matheus, P., Storer, J.E. & Sher, A. (2008) Paleoenvironmental reconstruction of the Last Glacial Maximum, inferred from insect fossils from a tephra buried soil at Tempest Lake, Seward Peninsula, Alaska. Palaeogeography, Palaeoclimatology, Palaeoecology, 267, 245–255.

    https://doi.org/10.1016/j.palaeo.2008.07.007

    Lemic, D., Benítez, H.A., Püschel, T.A., Gašpari, H.V., Šatvar, M. & Bažok, R. (2016) Ecological morphology of the sugar beet weevil Croatian populations: Evaluating the role of environmental conditions on body shape. Zoologischer Anzeiger, 260, 25–32.
    https://doi.org/10.1016/j.jcz.2015.11.003

    Lewis, P.O. (2001) A likelihood approach to estimating phylogeny from discrete morphological character data. Systematic Biology, 50, 913–925.

    https://doi.org/10.1016/j.jcz.2015.11.003

    Marshall, A.K. (1944) New injurious Curculionidae from Africa (Col.). Bulletin of Entomological Research, 35, 43–48.

    https://doi.org/10.1080/106351501753462876

    McKenna, D.D., Sequeira, A.S., Marvaldi, A.E. & Farrell, B.D. (2009) Temporal lags and overlap in the diversification of weevils and flowering plants. Proceedings of the National Academy of Sciences, 106 (17), 7083–7088.

    https://doi.org/10.1073/pnas.0810618106

    McKenna, D.D., Scully, E.D., Pauchet, Y., Hoover, K., Kirsch, R., Geib, S.M., Mitchell, R.F., Waterhouse, R.M., Ahn, S., Arsala, D., Childers, A.K., Benoit, J.B., Blackmon, H., Bledsoe, T., Bowsher, J., Busch, A., Calla Zalles, B., Chao, H., Childers, C., Clark, D.J., Cohen, L., Demuth, J.P., Dinh, H., Doddapaneni, H., Dolan, A., Duan, J.J., Dugan, S., Friedrich, M., Glastad, K.M., Goodisman, M.D., Haddad, S., Han, Y., Hughes, D.T., Ioannidis, P., Vargas Jentzsch, I.M., Johnston, J., Jones, J.W., Kuhn, L.A., Lance, D.R., Lee, C., Lee, S.L., Lin, H., Lynch, J.A., Moczek, A.P., Murali, S.C., Muzny, D.M., Nelson, D.R., Palli, S.R., Panfilio, K.A., Pers, D., Poelchau, M.F., Quan, H., Qu, J., Ray, A.M., Rinehart, J.P., Robertson, H.M., Roehrdanz, R.L., Rosendale, A.J., Shin, S., Silva, C., Torson, A., Werren, J.H., Worley, K.C., Yocum, G.D., Zdobnov, E.M., Gibbs, R.A. & Richards, S. (2016) Genome of the Asian longhorned beetle (Anoplophora glabripennis), a globally significant invasive species, reveals key functional and evolutionary innovations at the beetle–plant interface. Genome Biology, 17, 227.

    https://doi.org/10.1186/s13059-016-1088-8

    McKenna, D.D., Wild, A.L., Kanda, K., Bellamy, C.L., Beutel, R.G., Caterino, M.S., Farnum, C.W., Hawks, D.C., Ivie, M.A., Jameson, M.L., Leschen, R.A.B., Marvaldi, A.E., McHugh, J.V., Newton, A.F., Robertson, J.A., Thayer, M.K., Whiting, M.F., Lawrence, J.F., Slipinski, A., Maddison, D.R. & Farrell, B.D. (2015) Beetles survived the end Permian mass extinction to diversify during the Cretaceous terrestrial revolution. Systematic Entomology, 40, 835–880.

    https://doi.org/10.1111/syen.12132

    Meregalli, M. (2002) Notes on the genus Pachycerus Schoenherr 1823 with description of a new species from Somalia (Coleoptera Curculionidae Lixinae). Tropical Zoology, 15, 233–242.

    https://doi.org/10.1080/03946975.2002.10531177

    Meregalli, M. (2004) Pseudeumecops n. gen., a new genus of Cleonini from East Africa (Coleoptera, Curculionidae, Lixinae). Italian Journal of Zoology, 71, 153–163.

    https://doi.org/10.1080/11250000409356567

    Meregalli, M. (2005) Eurycleonus talamellii n. sp. of Cleonine weevil from the Moroccan desert (Coleoptera: Curculionidae: Lixinae: Cleonini). Zootaxa, 1053, 23–34.

    Meregalli, M. (2008) Taxonomic relationships between Pachycerus and Rhabdorrhynchus (Coleoptera: Curculionidae: Lixinae), with descriptions of two new species of Rhabdorrhynchus from the Arabian Peninsula. Zoological Journal of the Linnean Society, 152, 25–37.

    https://doi.org/10.1111/j.1096-3642.2007.00335.x

    Meregalli, M. (2009) Revision of the Indo-African Pachycerus Schoenherr, 1823, with a description of four new species (Coleoptera: Curculionidae: Lixinae). Zoological Journal of the Linnean Society, 157, 295–325.

    https://doi.org/10.1111/j.1096-3642.2008.00506.x

    Meregalli, M. (2014) 3.7.6 Lixinae Schoenherr, 1823. In: Leschen, R.A.B. & Beutel, R.G. (Eds.), Handbook of Zoology, Arthropoda: Insecta: Coleoptera. Volume 3: Morphology and Systematics (Phytophaga). Walter de Gruyter, Berlin, pp. 523–529.

    Meregalli, M. (2017) World catalogue of the Curculionidae: Lixinae: Cleonini. Available from: http://weevil.info/content/world-catalogue-curculionidae-lixinae-cleonini# (accessed 19 July 2017)

    Meregalli, M. & Fremuth, J. (2013) tribe Cleonini. In: Löbl, I. & Smetana, A. (Eds.), Catalogue of Palaearctic Coleoptera. Vol. 8. Curculionoidea II. Brill, Leiden, pp. 437–456.

    Meregalli, M. & Silvestro, D. (2008) Assessing relationships between Mediterranean and Afrotropical taxa of Cleonini (Coleoptera: Curculionidae: Lixinae): a survey based on Bayesian analysis and lineage geohistory reconstruction. Biogeographia, 29, 117–132.

    https://doi.org/10.21426/b6110181

    Meregalli, M. & Talamelli, F. (2009) Revision of the genus Epexochus Reitter, with description of three new species (Coleoptera: Curculionidae: Lixinae: Cleonini). Zootaxa, 2011, 47–68.

    Meregalli, M. & Talamelli, F. (2017) A review of the Pseudocleonus subgenus Phryganocleonus Arnol’di, with the description of a new species from Afghanistan (Coleoptera, Curculionidae: Lixinae). Entomological Review, 97, 82–89.

    https://doi.org/10.1134/S0013873817010109

    Motschulsky, V. (1860) Coléoptères rapportés en 1859 par M. Sévertsef des steppes méridionales des Kirghises, et enumerés par V. de Motschulsky. Bulletin Physico–Mathématique de l’Academie Impériale des Sciences de St. Petersburb, 2, 513–544.

    Nixon, K.C. (2002) WinClada V. 1.00.08. Published by the author, Ithaca, NY, USA.

    Oberprieler, R.G., Marvaldi, A.E. & Anderson, R.S. (2007) Weevils, weevils, weevils everywhere. Zootaxa, 1668, 491–520.

    O’Brien C.W. & Marshall, G.B. (1987) Unusual larval “sand tubes” constructions in the weevil genus Cleonidius in West Texas. The Southwestern Entomologist, 12, 357–360.

    Omland, K.E., Cook, L.G. & Crisp, M.D. (2008) Tree thinking for all biology: the problem with reading phylogenies as ladders of progress. BioEssays, 30, 854–867.

    https://doi.org/10.1002/bies.20794

    Parker, J. & Grimaldi, D.A. (2014) Specialized myrmecophily at the ecological dawn of modern ants. Current Biology, 24, 2428–2434.

    https://doi.org/10.1016/j.cub.2014.08.068

    Pomerinke, M.A., Thompson, D.C. & Clason, D.L. (1995) Bionomics of Cleonidius trivittatus (Coleoptera: Curculionidae): native biological control of Purple Locoweed (Rosales: Fabaceae). Environmental Entomology, 24, 1696–1702.

    https://doi.org/10.1093/ee/24.6.1696

    Rambaut, A., Suchard, M.A., Xie, D. & Drummond, A.J. (2013) Tracer v1.5. Available from: http://beast.bio.ed.ac.uk/Tracer (accessed 21 December 2016)

    Ronquist, F., Teslenko, M., van der Mark, P., Ayres, D.L., Darling, A., Höhna, S., Larget, B., Liu, L., Suchard, M.A. & Huelsenbeck, J.P. (2012) MrBayes 3.2: efficient Bayesian phylogenetic inference and model choice across a large model space. Systematic Biology, 61, 539–542.

    https://doi.org/ 10.1093/sysbio/sys029

    Shelef, O., Helman, Y., Friedman, A.L.L., Behar, A. & Rachmilevitch, S. (2013) Tri-party underground symbiosis between a weevil, bacteria and a desert plant. PLoS ONE, 8 (11), e76588.
    https://doi.org/10.1371/journal.pone.0076588

    Sher, A.V., Kuzmina, S.A., Kuznetsova, T.V. & Sulerzhitsky, L.D. (2005) New insights into the Weichselian environment and climate of the East Siberian Arctic, derived from fossil insects, plants, and mammals. Quaternary Science Reviews, 24, 533–569.

    https://doi.org/10.1016/j.quascirev.2004.09.007

    Skuhrovec, J., Stejskal, R., Trnka, F. & Gosik, R. (2014) Digital-Weevil-Determination for Curculionoidea of West Palaearctic. Tranaslpina: Cleonini. Snudebiller, 15, 1–18.

    Stinson, C.S.A., Schroeder, D. & Marquardt, K. (1994) Investigations on Cyphocleonus achates (Fahr.) (Col., Curculionidae), a potential biological control agent of spotted knapweed (Centaurea maculosa Lam.) and diffuse knapweed (C. diffusa Lam.) Compositae in North America. Journal of Applied Entomology, 117, 35–50.

    https://doi.org/10.1111/j.1439-0418.1994.tb00705.x

    Stejskal, R. & Trnka, F. (2014) Curculionidae: Lixinae. Icones insectorum Europae centralis. Folia Heyrovskyana, 20, 1–17.

    Stejskal, R., Trnka, F. & Skuhrovec, J. (2014) Biology and morphology of immature stages of Coniocleonus nigrosuturatus (Coleoptera: Curculionidae: Lixinae). Acta Entomologica Musei Nationalis Pragae, 54, 337–354.

    Story, J.M., Callan, N.W., Corn, J.G. & White, L.J. (2006) Decline of spotted knapweed density at two sites in western Montana with large populations of the introduced root weevil, Cyphocleonus achates (Fahraeus). Biological Control, 38, 227–232.

    https://doi.org/10.1016/j.biocontrol.2005.12.018

    Swofford, D.L. (2003) PAUP: Phylogenetic Analysis Using Parsimony (*and Other Methods), Version 4. Sinauer Associates, Sunderland, Massachusetts.

    Ter-Minassian, M.E. (1967) Weevils of the subfamily Cleoninae of the fauna of USSR: tribe Lixini. Opredeliteli po faune SSSR, izdavaemye Zoologicheskim Institutom Akademii Nauk SSSR, 95, 1–142. [in Russian]

    Ter-Minassian, M.E. (1974) A short review of the Palaearctic fauna of the genus Stephanocleonus Motschulsky (Coleoptera, Curculionidae). Insects of Mongolia, 2, 211–227. [in Russian]

    Ter-Minassian, M.E. (1979) Review of the weevil genus Stephanocleonus Motsch. (Coleoptera, Curculionidae). Insects of Mongolia, 6, 184–342. [in Russian]

    Ter-Minassian, M.E. (1984) Weevils of the subfamily Cleoninae (Coleoptera, Curculionidae) in the fauna of Mongolia. Insects of Mongolia, 10, 393–412. [in Russian]

    Ter-Minassian, M.E. (1988) Weevils of the subfamily Cleoninae of the fauna of USSR: tribe Cleonini. Opredeliteli po faune SSSR, izdavaemye Zoologicheskim Institutom Akademii Nauk SSSR, 155, 1–235. [in Russian]

    Tóth, M., Furlan, L. & Campagna, G. (2007) Attractant for the sugar-beet weevil Conorrhynchus (Cleonus) mendicus (Col.: Curculionidae). Journal of Applied Entomology, 131, 569–572.

    https://doi.org/10.1111/j.1439-0418.2007.01215.x

    Trnka, F., Stejskal, R. & Skuhrovec, J. (2015) Biology and morphology of immature stages of Adosomus roridus (Coleoptera: Curculionidae: Lixinae). Zootaxa, 4021 (3), 433–446.

    https://doi.org/10.11646/zootaxa.4021.3.3

    Van Dam, M.H. (2014) A simple, rapid technique for the inflation of the endophallus, with particular focus on the Curculionoidea (Coleoptera). The Coleopterists Bulletin, 6, 263–268.

    Volovnik, S.V. (2010) Weevils Lixinae (Coleoptera, Curculionidae) as gall formers. Entomological Review, 90, 585–590.

    https://doi.org/10.1134/s0013873810050052

    Wang, J.-W., Luo, Y.-Q., Zong, S.-X., Gao, B.-Q. & Li, Y.-H. (2009) Characterization of larval instars of Artemisia weevil, Adosomus sp. Forestry Studies in China, 11, 20–23.

    Winter, S., Friedman, A.L.L., Astrin, J.J., Gottsberger, B. & Letsch, H. (2017) Timing and host plant associations in the evolution of the weevil tribe Apionini (Apioninae, Brentidae, Curculionoidea, Coleoptera) indicate an ancient co-diversification pattern of beetles and flowering plants. Molecular Phylogenetics and Evolution, 107, 179–190.

    https://doi.org/10.1016/j.ympev.2016.10.015

    Yamamoto, S., Maruyama, M. & Parker, J. (2016) Evidence for social parasitism of early insect societies by Cretaceous rove beetles. Nature Communications, 7, 13658.

    https://doi.org/10.1038/ncomms13658

    Zachos, F.E. (2014) Paraphyly—again!? A plea against the dissociation of taxonomy and phylogenetics. Zootaxa, 3764 (5), 594–596.

    https://doi.org/10.11646/zootaxa.3764.5.8

    Zherikhin, V.V. & Egorov, A.B. (1990) Weevils (Coleoptera, Curculionidae) of the Far East of Russia (with review of subfamilies and description of new taxa). Vladivostok, Far East Brunch of the Academy of Sciences of USSR, 164 pp. [in Russian]

    Zotov, A.A. (2011) Morphology of the preimaginal stages of weevils of the tribe Cleonini sensu lato (Coleoptera: Curculionidae). Caucasian Entomological Bulletin, 7, 153–162. [in Russian]

How to Cite

ARZANOV, Y. G., & GREBENNIKOV, V. V. (2017). Cleonini (Coleoptera: Curculionidae: Lixinae) are monophyletic and flightless: tribe overview, rampant adult homoplasy and illustrated global diversity. Zootaxa, 4329(1), 1–63. https://doi.org/10.11646/zootaxa.4329.1.1