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Lachesillidae is one of the largest families of bark lice and includes more than 420 described species, in 26 genera and three subfamilies. This family belongs in the suborder Psocomorpha, infraorder Homilopsocidea. The classification of Lachesillidae is based on male and female genital morphologies, but questions remain regarding the monophyly of the family and some of its genera. Here, we used whole genome and transcriptome data to generate a 2060 orthologous gene data matrix of 2,438,763 aligned bp and used these data to reconstruct the phylogenetic relationships of species of Lachesillidae and relatives. Taxon sampling included 24 species from Lachesillidae and 23 additional species belonging to related families from the infraorders Homilopsocidea and Caeciliusetae. Phylogenetic relationships reconstructed with maximum likelihood and coalescent‐based analyses indicated paraphyly of Lachesillidae, and monophyly of the tribe Graphocaeciliini and the genusLachesillawere also never recovered. Instability was observed in the position ofEolachesilla chilensis, which was recovered either as sister to Elipsocidae or to Mesopsocidae species, so we cannot conclusively determine the position of this genus within the Homilopsocidea. Given our results, a reclassification is necessary, but more taxon sampling of other species in Mesopsocidae and Peripsocidae would be useful to add to a tree in future before proposing a new classification.

Psocodea (booklice and parasitic lice) is an order of insects containing species with extensive mitochondrial genome rearrangements, particularly within the suborder Troctomorpha, in which some species possess an extremely fragmented mitochondrial genome with several small minichromosomes. In the remaining suborders of Psocodea, there are groups with the ancestral pancrustacean arrangement, quite extensive rearrangements (e.g. Trogiomorpha), or in which the small number of species analysed to date have rearrangements of only a few protein‐coding genes and/or tRNAs (e.g. Psocomorpha). Despite the apparent high rate of rearrangements in the order as a whole, a small number of complete mitochondrial genomes are available, especially for suborder Psocomorpha, the largest free‐living suborder. To understand the evolution of the gene arrangement of the mitochondrial genome within Psocomorpha and its phylogenetic implications, we assembled and analysed the mitochondrial genomes of 33 species of bark lice belonging to nine families in two infraorders. Within the infraorder Homilopsocidea, four families were analysed, mainly from Lachesillidae (which included 22 species of this family). Within the infraorder Caeciliusetae, seven species representing five families were analysed. Mitochondrial gene rearrangements were identified in seven of the nine families. Some of these rearrangements were unique to a single species, while some contained phylogenetic signal, being shared by related species. These rearrangements typically corresponded to transpositions and inversions of tRNAs, possibly caused by tandem duplication–random loss (TDRL) and/or recombination events. Phylogenetic analyses of mitochondrial gene sequences provided phylogenetic resolution for several branches of the tree, including monophyly of Lachesillinae. The genusHemicaeciliusEnderlein was found to be embedded within the genusLachesillaWestwood, rending the latter paraphyletic. Monophyly was also never recovered for Lachesillidae and Elipsocidae as currently defined. However, instability was observed for some higher level relationships within Psocomorpha, including the relationships among the major clades of Lachesillidae.