Monday, October 10, 2022

Taxonomy and Phylogeny of Weevils

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). E. C. Zimmerman proposed a third division, the Heteromorphi, for several intermediate forms. Primitive weevils are distinguished by having straight antennae, while true weevils have elbowed (geniculate) antennae. The elbow occurs at the end of the 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, with updates from Marvaldi et al. in 2002, 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 and the recognition of the Cretaceous age family Mesophyletidae in 2018 from Burmese amber. The oldest known weevils are from the Middle-Late Jurassic Karabastau Formation of Kazakhstan, with species assigned to the extant families Nemonychidae, Anthribidae and Ithyceridae. The extinct family Obrieniidae, with species dating from the Ladinian stage of the Triassic through to tentatively the Oxfordian, have sometimes been considered weevils. Genera of the family have been only found in three formations in Kazakhstan, with most named in 1993. However, their phylogenetic position is contested, with others considering it part of Archostemata.

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