A Combined Molecular and Morphological Approach to Explore the Higher Phylogeny of Entimine Weevils (Coleoptera: Curculionidae), with Special Reference to South American Taxa

Abstract

The Entiminae are broad-nosed weevils constituting the most diverse subfamily of Curculionidae, with over 50 tribes. We performed Bayesian and Maximum Parsimony combined phylogenetic analyses with the main objective of testing higher-level relationships and the naturalness of the major Neotropical and Southern South American (Patagonia and Andes) tribes, including some members from other regions. We compiled a data matrix of 67 terminal units with 63 Entiminae species, as well as four outgroup taxa from Cyclominae, by 3522 molecular (from nuclear 18S rDNA and 28S rDNA, and mitochondrial 16S rDNA and COI gene sequences) and 70 morphological characters. The resulting trees recover a clade Entiminae with a monophyletic Cylydrorhinini and Premnotrypes branching off early. The tree resulting from parsimony analysis shows a clade of Leptopiini from the Australian region and another clade including taxa mainly distributed in the Palaearctic and Neotropical regions, but in the Bayesian tree the South American and Australian Leptopiini are grouped together. The mainly Palaearctic Entiminae (e.g., Brachyderini, Laparocerini, Otiorhynchini, Peritelini, Polydrusini, Phyllobiini and Sciaphylini) form a subclade separated from Southern Hemisphere taxa. Among the latter, the well-supported Naupactini are the sister group of the South American Tanymecini, excluding Platyaspistes, herein transferred to Leptopiini (new placement). Another well-justified clade is Eustylini–Geonemini, which also includes the enigmatic Galapagonotus, and the genus Artipus, thus corroborating its recent exclusion from Naupactini.

Systematic revision of the genusIsodacrysSharp, 1911 (Coleoptera: Curculionidae: Entiminae: Tanymecini)

“…Howden (1959) first used the term nasal plate for Tanymecini as “the sclerite that is within the apical emargination or extend forward from the apex of the beak”, considering it as a synonym of the epistome. In the literature, the terms nasal plate and epistome have been used to refer to the same structure or different parts of the apical region of the rostrum ( Pierce, 1913 ; Blatchley & Leng, 1916 ; Howden, 1959 ; Anderson, 2002 ; Franz, 2012 ; Oberprieler, Anderson & Marvaldi, 2014 , Jansen & Franz, 2015 , 2018 ; Marvaldi et al, 2018 ; Girón & Howden, 2019 ). In the present study, the apical emargination refers to the posterior margin of the nasal plate.…”

“…Non-additive. See character 3 in Marvaldi et al (2018) . Character state 1 synapomorphy for the Isodrusus debilis - I. guajavus clade; character state 2 synapomorphy for the Pandeleteius hilaris - P. inflatus clade (l: 2, ci: 1, ri: 1).…”

“…Tribal concepts follow Alonso-Zarazaga & Lyal (1999) , except for the genus Platyaspistes Schönherr, 1840 which is considered to belong to Leptopiini Oke, 1951 instead of Tropiphorini Marseul, 1863 (see Marvaldi et al, 2018 ). An exemplar approach based on morphology was followed ( Prendini, 2001 ).…”

“…The outgroup comprised 21 species representing eight tanymecine genera ( Table 1 ), and one member of the tribes Leptopiini ( Platyaspistes prasinus [Erichson, 1834], recently transferred from Piazomiina Reitter (1913) by Marvaldi et al (2018) and Naupactini Gistel (1856) ( Megalostylus albicans [Lacordaire]), all distributed in the Americas. Specimens of the type species of each outgroup genus were included when available.…”

“…Phylogenetic analyses were performed under maximum parsimony as optimality criterion in TNT v1.5 ( Goloboff & Catalano, 2016 ) using heuristic traditional search under equal weights. Cladograms obtained were rooted with Megalostylus albicans Champion, 1911 (Entiminae: Naupactini) based on recent analyses of Entiminae weevils depicting the phylogenetic closeness of Naupactini and Tanymecini ( Marvaldi et al, 2018 ; contra Gillet et al, 2018 ). Most parsimonious trees (MPT’s) were consistently obtained through several rounds of analyses modifying the number of replicates and trees retained per replicate.…”