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{{short description|Superfamily of beetles}} {{About|the beetles}} {{Automatic taxobox | fossil_range = {{fossilrange|166|0|[[Middle Jurassic]] - [[Recent]]|earliest=Ladinian}} | name = Weevils | image = Weevil September 2008-1.jpg | image_caption = ''[[Lixus pulverulentus]]'' | taxon = Curculionoidea | display_parents = 3 | authority = [[Pierre André Latreille|Latreille]], 1802 | subdivision_ranks = [[Family (biology)|Families]] | subdivision = * [[Anthribidae]] * [[Attelabidae]] * [[Belidae]] * [[Brentidae]] * [[Caridae]] * [[Cimberididae]] * [[Curculionidae]] * {{extinct}}[[Mesophyletidae]] * [[Nemonychidae]] * {{extinct}}?[[Obrieniidae]] }} '''Weevils''' are [[beetle]]s belonging to the [[Taxonomic rank|superfamily]] '''Curculionoidea''', known for their elongated snouts. They are usually small – less than {{convert|6|mm|abbr=on|frac=4}} in length – and [[Herbivore|herbivorous]]. Approximately 97,000 species of weevils are known. They belong to several families, with most of them in the family [[Curculionidae]] (the [[true weevil]]s). It also includes [[Bark beetle|bark beetles]], which while morphologically dissimilar to other weevils in lacking the distinctive snout, is a subfamily of Curculionidae. Some other beetles, although not closely related, bear the name "weevil", such as the [[leaf beetle]] subfamily [[bean weevil|Bruchinae]], known as "bean weevils", or the [[Drugstore beetle|biscuit weevil]] (''Stegobium paniceum''), which belongs to the family [[Ptinidae]]. Many weevils are considered pests because of their ability to damage and kill crops. The grain or [[wheat weevil]] (''Sitophilus granarius'') damages stored [[cereal|grain]], as does the [[maize weevil]] (''Sitophilus zeamais''), among others. The [[boll weevil]] (''Anthonomus grandis'') attacks [[cotton]] crops; it lays its eggs inside cotton bolls and the larvae eat their way out. Other weevils are used for biological control of invasive plants. A weevil's [[Rostrum (anatomy)|rostrum]], or elongated snout, hosts chewing [[Arthropod mouthparts|mouthparts]] instead of the piercing mouthparts that [[proboscis]]-possessing insects are known for. The mouthparts are often used to excavate tunnels into grains.<ref>{{Cite journal|last=Moon|first=Myung-Jin|date=2015|title=Microstructure of mandibulate mouthparts in the greater rice weevil, Sitophilus zeamais (Coleoptera: Curculionidae)|journal=Entomological Research|language=en|volume=45|issue=1|pages=9–15|doi=10.1111/1748-5967.12086|s2cid=82634306|issn=1748-5967}}</ref> In more derived weevils, the rostrum has a groove in which the weevil can fold the first segment of its antennae. Most weevils have the ability to fly (including pest species such as the [[rice weevil]]),<ref>{{cite web|url=https://www.thespruce.com/what-is-a-weevil-2656439|title=What Is a Weevil and How Did That Bug Get in My Food?}}</ref><ref>{{cite web|url=http://ento.psu.edu/extension/factsheets/weevils-on-stored-grain|title=Weevils on Stored Grain (Department of Entomology)|website=Department of Entomology (Penn State University)}}</ref> though a significant number are flightless, such as the genus ''[[Otiorhynchus]]'', and others can jump. One species of weevil, ''[[Austroplatypus incompertus]]'', exhibits [[eusociality]], one of the few insects outside the [[Hymenoptera]] and the [[Termite|Isoptera]] to do so. [[File:Weevil3.jpg|thumb|''[[Curculio nucum]]'']] ==Taxonomy and phylogeny== Because so many species exist in such diversity, the higher classification of weevils is in a state of flux. They are generally divided into two major divisions, the Orthoceri or primitive weevils, and the Gonatoceri or true weevils ([[Curculionidae]]). [[Elwood Zimmerman|E. C. Zimmerman]] proposed a third division, the Heteromorphi, for several intermediate forms.<ref>{{cite book |author=E. C. Zimmerman |year=1994 |title=Australian weevils (Coleoptera: Curculionidae). Volume 1. Orthoceri: Anthribidae to Attelabidae: the primitive weevils |publisher=[[Commonwealth Scientific and Industrial Research Organisation|CSIRO]] |location=[[East Melbourne, Victoria|East Melbourne]] |pages=741 pp}}</ref> Primitive weevils are distinguished by having straight antennae, while true weevils have elbowed (geniculate) antennae. The elbow occurs at the end of the [[antenna (biology)#Structure|scape]] (first antennal segment) in true weevils, and the scape is usually much longer than the other antennal segments. Some exceptions occur, such as [[Nanophyini]], primitive weevils with long scapes and geniculate antennae, while among the true weevils, [[Gonipterinae]] and ''[[Ramphus]]'' have short scapes and little or no "elbow". A 1995 classification system to family level was provided by Kuschel,<ref>{{cite journal |author=G. Kuschel |year=1995 |title=A phylogenetic classification of Curculionoidea to families and subfamilies |journal=[[Memoirs of the Entomological Society of Washington]] |volume=14 |pages=5–33}}</ref> with updates from Marvaldi ''et al.'' in 2002,<ref name=Marvaldi2002>{{cite journal |last1=Marvaldi |first1=Adriana E. |last2=Sequeira |first2=Andrea S. |last3=O'Brien |first3=Charles W. |last4=Farrell |first4=Brian D. |year=2002 |title=Molecular and morphological phylogenetics of weevils (Coleoptera, Curculionidae): do niche shifts accompany diversification? |journal=[[Systematic Biology]] |volume=51 |issue=5 |pages=761–785 |doi=10.1080/10635150290102465 |pmid=12396590|doi-access=free }}</ref> and was achieved using [[phylogenetic]] analyses. The accepted families were the primitive weevils, [[Anthribidae]], [[Attelabidae]], [[Belidae]], [[Brentidae]], [[Caridae]], and [[Nemonychidae]], and the true weevils [[Curculionidae]]. Most other weevil families were demoted to subfamilies or tribes. Further work resulted in the elevation of [[Cimberididae]] to family from placement as a subfamily of Nemonychidae in 2017<ref name=Shin2018>{{cite journal |last1=Shin |first1=Seunggwan |last2=Clarke |first2=Dave J. |last3=Lemmon |first3=Alan R. |last4=Lemmon |first4=Emily Moriarty |last5=Aitken |first5=Alexander L. |last6=Haddad |first6=Stephanie |last7=Farrell |first7=Brian D. |last8=Marvaldi |first8=Adriana E. |last9=Oberprieler |first9=Rolf G. |last10=McKenna |first10=Duane D. |title=Phylogenomic data yield new and robust insights into the phylogeny and evolution of weevils |journal=Molecular Biology and Evolution |year=2018 |volume=35 |issue=4 |pages=823–836 |doi=10.1093/molbev/msx324|pmid=29294021 |doi-access=free |hdl=11336/57287 |hdl-access=free }}</ref> and the recognition of the [[Cretaceous]] age family [[Mesophyletidae]] in 2018 from [[Burmese amber]].<ref name=ClarkeetalDiversity>{{cite journal |author1=Dave J. Clarke |author2=Ajay Limaye |author3=Duane D. McKenna |author4=Rolf G. Oberprieler |year=2018 |title=The weevil fauna preserved in Burmese amber—snapshot of a unique, extinct lineage (Coleoptera: Curculionoidea) |journal=Diversity |volume=11 |issue=1 |pages=Article 1 |doi=10.3390/d11010001 |doi-access=free }}</ref> The oldest weevils date to the Middle-Late Jurassic boundary, found in the [[Karabastau Formation]] of Kazakhstan, the [[Ulan Malgait Formation|Shar-Teg]] locality of Mongolia, the [[Haifanggou Formation|Daohugou]] locality in Inner Mongolia, China, and the [[Talbragar fossil site|Talbragar site]] in Australia.<ref>{{Cite journal |last=Legalov |first=A. A. |date=April 2022 |title=Review of the Jurassic Weevils of the Genus Belonotaris Arnoldi (Coleoptera: Nemonychidae) with a Straight Rostrum |url=http://dx.doi.org/10.1134/s0031030122020071 |journal=Paleontological Journal |volume=56 |issue=2 |pages=199–207 |doi=10.1134/s0031030122020071 |s2cid=248303405 |issn=0031-0301|url-access=subscription }}</ref> The extinct family [[Obrieniidae]], with species dating from the [[Ladinian]] stage of the [[Triassic]] through to tentatively the [[Oxfordian (stage)|Oxfordian]], have sometimes been considered weevils. Genera of the family have only been found in three formations in [[Kazakhstan]], with most named in 1993.<ref name="Legalovlist2010">{{cite journal |last1=Legalov |first1=A. A. |year=2010 |title=Checklist of Mesozoic Curculionoidea (Coleoptera) with description of new taxa |journal=Baltic Journal of Coleopterology |volume=10 |issue=1 |pages=71–101}}</ref> However, their phylogenetic position is contested, with others considering it part of [[Archostemata]].<ref>{{Cite journal |last=Legalov|first=A. |date=2020 |title=Fossil History of Curculionoidea (Coleoptera) from the Paleogene |journal=Geosciences |volume=10 |issue=9 |pages=358 |doi=10.3390/geosciences10090358|bibcode=2020Geosc..10..358L |doi-access=free |language=en}}</ref> The interfamilial relationships of Curculionoidea have been generally well resolved.<ref name=Marvaldi2002/><ref name=Shin2018/><ref>{{cite journal |last1=Marvaldi |first1=A. E. |last2=Morrone |first2=J. J. |year=2000 |title=Phylogenetic systematics of weevils (Coleoptera: Curculionoidea): A reappraisal based on larval and adult morphology |journal=Insect Systematics & Evolution |volume=31 |pages=43–58 |doi=10.1163/187631200X00309 |hdl=11336/152180 |hdl-access=free }}</ref><ref>{{cite journal |last1=Marvaldi |first1=Adriana E. |last2=Duckett |first2=Catherine N. |last3=Kjer |first3=Karl M. |last4=Gillespie |first4=Joseph J. |year=2009 |title=Structural alignment of 18S and 28S rDNA sequences provides insights into phylogeny of Phytophaga (Coleoptera: Curculionoidea and Chrysomeloidea) |journal=[[Zoologica Scripta]] |volume=38 |pages=63–77 |doi=10.1111/j.1463-6409.2008.00360.x |s2cid=84345520 |hdl=11336/92765 |hdl-access=free }}</ref><ref name=Li2023>{{cite journal |last1=Li |first1=Yan-Da |last2=Engel |first2=Michael S. |last3=Tihelka |first3=Erik |last4=Cai |first4=Chenyang |year=2023 |title=Phylogenomics of weevils revisited: data curation and modelling compositional heterogeneity |journal=Biology Letters |volume=19 |issue=9 |page=20230307 |doi=10.1098/rsbl.2023.0307 |pmid=37727076 |pmc=10509570 |doi-access=free }}</ref> The [[phylogeny]] by Li et al. (2023) based on phylogenomic data is suggested below:<ref name=Li2023/> {{clade|style=font-size:100%;line-height:100% |1={{clade |1=[[Cimberididae]] |2={{clade |1={{clade |1=[[Nemonychidae]] |2=[[Anthribidae]] }} |2={{clade |1=[[Belidae]] |2={{clade |1=[[Attelabidae]] |2={{clade |1=[[Caridae]] |2={{clade |1=[[Brentidae]] |2=[[Curculionidae]] }} }} }} }} }} }} }} ==Families== * [[Anthribidae]]—fungus weevils * [[Attelabidae]]—leaf rolling weevils * [[Belidae]]—primitive weevils * [[Brentidae]]—straight snout weevils * [[Caridae]] * [[Cimberididae]] * [[Curculionidae]]—true weevils * {{extinct}}[[Mesophyletidae]] * [[Nemonychidae]]—pine flower weevils * {{extinct}}?[[Obrieniidae]] == Sexual dimorphism == ''[[Rhopalapion longirostre]]'' exhibits an extreme case of [[sexual dimorphism]]. The female [[rostrum (anatomy)|rostrum]] is twice as long and its surface is smoother than in the male. The female bores egg channels into the buds of ''[[Alcea rosea]]''. Thus, the dimorphism is not attributed to sexual selection. It is a response to ecological demands of egg deposition.<ref>{{cite journal |author=G. Wilhelm|year=2011 |title=Sexual dimorphism in head structures of the weevil ''Rhopalapion longirostre'': a response to ecological demands of egg deposition |journal=[[Biological Journal of the Linnean Society]] |volume=104 |pages=642–660 |doi=10.1111/j.1095-8312.2011.01751.x|display-authors=etal|doi-access=free }}</ref> Another example of extreme dimorphism in weevils is that of the [[New Zealand giraffe weevil]]. Males measure up to {{convert|90|mm|in|abbr=on}} and females {{convert|50|mm|in|abbr=on}}, although there is an extreme range of body sizes in both sexes. ==References== {{Reflist}} ==Further reading== * Bright, Donald E.; Bouchard, Patrice (2008). [https://books.google.com/books?id=_IFYt8o7giYC ''Coleoptera, Curculionidae, Entiminae: Weevils of Canada and Alaska Volume 2'']. Insects and Arachnids of Canada Series, Part 25. Ottawa: NRC Research Press. {{ISBN|978-0-660-19400-4}}. == External links == * {{Commons category-inline|Curculionoidea}} {{Coleoptera|4}} {{Taxonbar|from=Q383963}} {{Authority control}} {{DEFAULTSORT:Weevil}} [[Category:Weevils| ]] <!-- [[Category:Polyphaga]] category above is already included here --> [[de:Rüsselkäfer]]
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