Phylogenetic relationships of “Tenebrionidae clade”
In the “Tenebrionidae clade”, the family Lagriidae and Tenebrionidae are monophyletic, and both of them are sister groups each other in the present study. The classification of Lagriidae has been debated over the years, and some scholars argue that Lagriidae should be classified as a subfamily within Tenebrionidae. However, the beetles of Lagriidae are leafivorous like beetles of Chrysomelidae, and morphological characteristics of Lagriidae adapt much more for free-moving and leaf-feeding than the beetles from other subfamilies in Tenebrionidae. The monophyly of Lagriidae was proposed in the earlier morphology study based on the characteristics of abdominal defense glands, female reproductive tract, mouthparts morphology and structure of the wings in Lagriid and Tenebrionoid (Doyen & Tschinkel, 1982). The monophyly was also proposed in earlier phylogenetic inference based on mtgenomes and nuclear genes (Gunter et al. , 2014; Kergoat et al. , 2014a). The mtgenome-based phylogeny of 36 species in Tenebrionidae suggested the monophyly of Lagriidae and Tenebrionidae, and their sister relationships based on PCG123 datasets using BI and ML (Wu et al. , 2021), which is consistent with the present study. The monophyly of Tenebrionidae was also proposed in some earlier phylogenetic studies based on mitochondrial and nuclear genes (Hunt et al ., 2007; Timmermans et al. , 2015; Mckennaet al. , 2019).
The present study suggests the phylogenetic relationships of Adeliinae + (Lagriinae + Statininae) in the family Lagriidae, and the monophyly of Lagriinae and Statininae. The monophyly of Lagriinae was proposed in earlier phylogenetic studies based on the morphology characteristics (Doyen, 1989), and also based on mitochondrial and nuclear genes (Gunteret al ., 2014; Wu et al ., 2021). The present study supports the monophyly of the subfamily Statiriinae for the first time. The Adeliinae is at the base of the family Lagriidae in this study, which is consistent with the earlier mtgenome-based study (Wu et al ., 2021). The monophyly of Adeliinae in Lagriidae is not yet determined due to only one species to be included.
In the family Tenebrionidae, the present study supports the monophyly of the subfamily Pimeliinae, Stenochiinae and Alleculinae, whereas Tenebrioninae and Diaperinae are recognized as a polyphyly. The subfamily Pimeliinae was also proposed to be monophyletic in the earlier studies based on nuclear genes and mitochondrial genes (Gunter et al. , 2014; Wu et al ., 2021). The present study supports the monophyly of the subfamilies Stenochiinae and Alleculinae in Tenebrionidae, which was also proposed in earlier phylogenetic studies based on mitochondrial and nuclear genes (Kergoat et al. , 2014a; Wu et al ., 2021). The subfamily Tenebrioninae and Diaperinae in these molecular-based studies are found to be polyphyletic, which is need to be elucidated with more species involved.