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Type: Article
Published: 2021-08-25
Page range: 201-220
Abstract views: 633
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The first molecular phylogeny of the weevil subfamily Lixinae (Coleoptera: Curculionidae) casts doubts on the monophyly of its tribes

National Nature Park “Priazovskyi”, Melitopol, Ukraine.
Ecology Group, Department of Biology, National University of Mongolia, Ikh Surguuliin Gudamj 1, Ulaanbaatar 14201, Mongolia.
Ecology Group, Department of Biology, National University of Mongolia, Ikh Surguuliin Gudamj 1, Ulaanbaatar 14201, Mongolia. &
Institute of General and Experimental Biology, Siberian Branch, Russian Academy of Sciences, Ulan-Ude, Russia.
Institute of Zoology, Al-Farabi, 93, Almaty 050060 Kazakhstan.
3Institute of General and Experimental Biology, Siberian Branch, Russian Academy of Sciences, Ulan-Ude, Russia.
Severtsov Institute of Ecology and Evolution, Russian Academy of Sciences, Leninskiy Prospect 33, Moscow, Russia.
Canadian Food Inspection Agency, 960 Carling Avenue, Ottawa, ON, K1A 0C6, Canada.
Coleoptera Curculionidae


This paper targets the nearly cosmopolitan weevil subfamily Lixinae. A phylogenetic analysis of one mitochondrial and two nuclear DNA fragments obtained from 87 Lixinae and 49 outgroup terminals strongly supports the monophyly of the subfamily. The molytine tribe Mecysolobini is the likeliest sister group of Lixinae; adults of both taxa share the likely synapomorphic condition of the greatly shortened labial palpi. Phylogenetic relationships within the subfamily are poorly resolved. None of three Lixinae tribes is recovered as monophyletic. The subfamily’s oldest dichotomy is formed by a clade of two genera Rhabdorrhynchus plus Pachycerus sister to the rest of the subfamily. The genera Chromonotus, Larinus, Lixus, and Stephanocleonus are recovered as non-monophyletic. The genera Asproparthenis, Chromonotus, and Maximus form a strongly supported clade. The genus Eumecops is the likeliest sister to the clade formed by the genera Stephanocleonus plus Coniocleonus. The cleonine genus Scaphomorphus is a sister to a subset of externally most similar species of the genus Lixus; the same clade likely includes the genus Lixoglyptus not represented in the analysis. As an aside we provide a short summary of active flight in adult Cleonini.



  1. Alonso-Zarazaga, M.A., Barrios, H., Borovec, R., Bouchard, P., Caldara, R., Colonnelli, E., Gültekin, L., Hlaváč, P., Korotyaev, B.A., Lyal, C.H.C., Machado, A., Meregalli, M., Pierotti, H., Ren, L., Sánchez-Ruiz, M., Sforzi, A., Silfverberg, H., Skuhrovec, J., Trýzna, M., Velázquez de Castro, A.J. & Yunakov, N.N. (2017) Cooperative Catalogue of Palaearctic Coleoptera Curculionoidea. Sociedad Entomológica Aragonesa, Monografias electrónicas SEA, 8, 1–729. Available from: (accessed 15 October 2019)
    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.
    Anderson, R.S. (1988) Systematics, phylogeny and biogeography of New World weevils traditionally of the tribe Cleonini (Coleoptera: Curculionidae; Cleoninae). Quaestiones Entomologicae, 23 (1987), 431–709.
    Anderson, R.S. (2018) Purealus beckelorum, a new genus and species of cleonine weevil from western Texas and eastern New Mexico (Coleoptera, Curculionidae, Lixinae, Cleonini). ZooKeys, 785, 1–10.
    Arzanov, Yu. G. (2009) 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. & 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.
    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]
    Auersch, O. (1961) Zur Kenntnis des Rübenderbrusslers (Bothynoderes punctiventris Germ.). Teil 2. Zeitschrift für Angewandte Entomologie, 49, 50–77.
    Brunner, Yu. N. (1959) Samarkandskyi sveklovichnyi dolgonosik. In: Savchenko, E.N. (Ed.), Sveklovodstvo. Vol. 3. Gosselkhozizdat USSR, Kiev, pp. 135–136. [in Russian]
    Chamorro, M.L., de Medeiros, B.A.S. & Farrell, B.D. (2021) First phylogenetic analysis of Dryophthorinae (Coleoptera, Curculionidae) based on structural alignment of ribosomal DNA reveals Cenozoic diversification. Ecology & Evolution, 11, 1984–1998.
    Colonnelli, E. (2003) A revised checklist of Italian Curculionoidea (Coleoptera). Zootaxa, 337, 10, 1–142.
    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.
    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.
    Grebennikov, V.V. & Anderson, R.S. (2021) Late Miocene eastwards transatlantic dispersal of flightless anchonine weevils (Coleoptera: Curculionidae: Molytinae). Zootaxa, 4952 (1), 55–70.
    Gunter, N.L., Oberprieler, R.G. & Cameron, S.L. (2016) Molecular phylogenetics of Australian weevils (Coleoptera: Curculionoidea): exploring relationships in a hyperdiverse lineage through comparison of independent analyses. Austral Entomology, 55, 217–233.
    Gültekin, L. (2010) Taxonomic remarks on some genera of Lixini Schoenherr, 1823 (Coleoptera: Curculionidae). Zootaxa, 2411 (1), 1–21.
    Gültekin, L. (2012) Allolarinus, a new genus and species of Lixini (Coleoptera: Curculionidae: Lixinae) from Congo. Annales de la Société Entomologique de France, 48, 67–65.
    Gültekin, L. (2013a) Afrolarinus, a new genus of Lixini (Coleoptera: Curculionidae: Lixinae) from Afrotropical region with taxonomic revision. Deutsche Entomologische Zeitschrift, 60, 251–260.
    Gültekin, L. (2013b) A new weevil genus Nefis gen. nov. (Coleoptera: Curculionidae: Lixinae): systematics and taxonomic revision. Journal of Insect Biodiversity, 1, 1–51.
    Gültekin, L. (2016) A remarkable new genus and species of Lixini (Coleoptera: Curculionidae: Lixinae) from Madagascar. Journal of Insect Biodiversity, 4, 1–10.
    Gültekin, L., Diotti, L. & Caldara, R. (2019) On the genus Rhinocyllus Germar (Coleoptera: Curculionidae: Lixinae) with description of a new subgenus and a new species from Italy. Journal of Insect Biodiversity, 11, 10–23.
    Gültekin, L. & Lyal, C.H.C. (2014) A taxonomic review on the genus Larinodontes Faust, 1898 (Coleoptera: Curculionidae) from the Oriental Region with a description of a new species from India. Journal of Insect Biodiversity, 2, 1–13.
    Haran, J., Timmermans, M.J.T.N. & Vogler, A.P. (2013) Mitogenome sequences stabilize the phylogenetics of weevils (Curculionoidea) and establish the monophyly of larval ectophagy. Molecular Phylogenetics and Evolution, 6, 156–166.
    Hebert, P.D.N., Ratnasingham, S. & DeWaard, J.R. (2003a) Barcoding animal life: cytochrome c oxidase subunit 1 divergences among closely related species. Proceedings of the Royal Society B: Biological Sciences, 270, 96–99.
    Hebert, P.D.N., Cywinska, A., Ball, S.L. & DeWaard, J.R. (2003b) Biological identifications through DNA barcodes. Proceedings of the Royal Society B: Biological Sciences, 270, 313–321.
    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.
    Jordal, B.H., Sequeira, A.S. & Cognato, A.I. (2011) The age and phylogeny of wood boring weevils and the origin of subsociality. Molecular Phylogenetics and Evolution, 59, 709–724.
    Katoh, K., Rozewicki, J. & Yamada, K.D. (2017) MAFFT online service: multiple sequence alignment, interactive sequence choice and visualization. Briefings in Bioinformatics, bbx, 108, 1–7.
    Kokovikhin, А.I. (1966) K biologii Chromoderus fasciatus Müll. (Coleoptera, Curculionidae ) v stepnoy zone Omskoi oblasti. In: Cherepanov, A.I. (Ed.), Fauna i Ekologija Chlenistonogikh Sibiri. Nauka—Sibirskoe otdelenie, Novosibirsk, рp. 29–32. [in Russian]
    Kuraku, S., Zmasek, C.M., Nishimura, O. & Katoh, K. (2013) aLeaves facilitates on-demand exploration of metazoan gene family trees on MAFFT sequence alignment server with enhanced interactivity. Nucleic Acids Research, 41, W22–W28.
    Kuschel, G. (1987) The subfamily Molytinae (Coleoptera: Curculionidae): general notes and descriptions of new taxa from New Zealand and Chile. New Zealand Entomologist, 9, 11–29.
    Letsch, H., Balke, M., Toussaint, E.F.A. & Riedel, A. (2020) Historical biogeography of the hyperdiverse hidden snout weevils (Coleoptera, Curculionidae, Cryptorhynchinae). Systematic Entomology, 45, 312–326.
    Lyal, C.H.C. (2014) 3.7.7 Molytinae Schoenherr, 1823. In: Leschen, R.A.B. & Beutel, R.G. (Eds.), Handbook of Zoology, Arthropoda: Insecta: Coleoptera. Vol. 3. Morphology and Systematics (Phytophaga). Walter de Gruyter, Berlin, pp. 529–570.
    Maddison, W.P. & Maddison, D.R. (2019) Mesquite: a modular system for evolutionary analysis. Version 3.5. Program and documentation. Available from: (accessed 15 March 2019)
    Marvaldi, A.E., Sequeira, A.S., O’Brien, C.W. & Farrell, B.D. (2002) Molecular and morphological phylogenetics of weevils (Coleoptera, Curculionoidea): do niche shifts accompany diversification? Systematic Biology, 51, 761–785.
    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 USA, 106, 7083–7088.
    Meregalli, M. (2005) Eurycleonus talamellii n. sp. of Cleonine weevil from the Moroccan desert (Coleoptera: Curculionidae: Lixinae: Cleonini). Zootaxa, 1053 (1), 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.
    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. (2017a) World catalogue of the Curculionidae: Lixinae: Cleonini. Available from: (accessed 19 July 2017).
    Meregalli, M. (2017b) Taxonomic and nomenclatural acts formalizing the classification provided in the world catalogue of the Cleonini (Curculionidae: Lixinae). Zootaxa, 4323 (3), 445–450.
    Miller, M., Pfeiffer, W. & Schwartz, T. (2010) Creating the CIPRES Science Gateway for inference of large phylogenetic trees. Proceedings of the Gateway Computing Environments Workshop (GCE), New Orleans, Louisiana, 2010, pp. 1–8.
    Mugu, S., Pistone, P. & Jordal, B.H. (2018) New molecular markers resolve the phylogenetic position of the enigmatic wood-boring weevils Platypodinae (Coleoptera: Curculionidae). Arthropod Systematics and Phylogeny, 76, 45–58.
    Orlachiova, K.A. (1956) Faktory, opredelyayushchie perelioty zhukov sveklovichnogo dolgonosika. In: Vasiliev, V.P. (Ed.), Sveklovichnyi Dolgonosik i Borba s Nim. Sbornik Nauchnykh Trudov Instituta Entomologii i Fitopatologii AN USSR. Vol. 7. Akademia Nauk USSR, Kiev, pp. 158–194 [in Russian].
    Pistone, D., Gohli, J. & Jordal, B.H. (2018) Molecular phylogeny of bark and ambrosia beetles (Curculionidae: Scolytinae) based on 18 molecular markers. Systematic Entomology, 43, 387–406.
    Rambaut, A. (2019) FigTree. Version 1.4.0. Program and documentation. Available from: (accessed 15 March 2019)
    Ratnasingham, S. & Hebert, P.D.N. (2007) BOLD: The Barcode of Life Data System ( Molecular Ecology Notes, 7, 355–364.
    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.
    Shin, S., Clarke, D.J., Lemmon, A.R., Lemmon, E.M., Aitken, A.L., Haddad, S., Farrell, B.D., Marvaldi, A.E., Oberprieler, R.G. & McKenna, D.D. (2017) Phylogenomic data yield new and robust insights into the phylogeny and evolution of weevils. Molecular Biology and Evolution, 35, 823–836.
    Skuhrovec, J., Volovnik, S., Gosik, R., Stejskal, R. & Trnka, F. (2019) Cleonis pigra (Scopoli, 1763) (Coleoptera: Curculionidae: Lixinae): morphological redescription of the immature stages, keys, tribal comparisons and biology. Insects, 10 (325), 1­–25.
    Skuhrovec, J., Caldara, R., Gosik, R., Trnka, F. & Stejskal, R. (2021) On the affinities and systematic position of Lachnaeus Schoenherr and Rhinocyllus Germar in the tribe Lixini (Coleoptera: Curculionidae: Lixinae) based on the morphological characters of the immature stages. Insects, 12 (6), 489, 1–25.
    Stamatakis, A. (2014) RAxML version 8: a tool for phylogenetic analysis and post-analysis of large phylogenies. Bioinformatics, 30, 1312–1313.
    Stamatakis, A., Hoover, P. & Rougemont, J. (2008) A rapid bootstrap algorithm for the RAxML web servers. Systematic Biology, 57, 758–771.
    Ter-Minassian, M.E. (1967) Zhuki-dolgonosiki podsemejstva Cleoninae fauny SSSR. Tsvetozh­ily i stebleedy (triba Lixini). Nauka, Leningrad, 140 + 1 pp. [1978 English translation: Weevils of the Subfamily Cleoninae in the Fauna of the USSR. Tribe Lixini. USDA Agricultural Research Service, Washington, D.C. and Amerind Publishing Co. Pvt. Ltd., New Delhi, 166 pp.]
    Tóth, M., Ujváry, I., Sivcev, I., Imrei, Z., Szarukán, I., Farkas, O., Gömöry, Á., Gács-Baitz, E. & Francke, W. (2006) An aggregation attractant for the sugar-beet weevil, Bothynoderes punctiventris. Entomologia Experimentalis et Applicata, 122, 125–132
    Volovnik, S.V. (2010) Weevils Lixinae (Coleoptera, Curculionidae) as gall formers. Entomological Review, 90, 585–590.
    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.
    Zherikhin, V.V. & Egorov, A.B. (1991) Zhuki-Dolgonosiki (Coleoptera, Curculionidae) Dal’nego Vostoka SSSR (Obzor Podsemestv s Opisaniem Novykh Taksonov). Akademiya Nauk SSSR, Vladivostok, 164 pp. [in Russian, 1990]