Phylogenomics facilitates stable scorpion systematics: Reassessing the relationships of Vaejovidae and a new higher-level classification of Scorpiones (Arachnida)

Abstract

The Neartic family Vaejovidae (Scorpiones: Chactoidea) has long been treated as a diverse and systematically cohesive group of scorpions, but its monophyly and relationship to other scorpion families have historically been questioned. Morphological data have supported its monophyly and a variety of phylogenetic placements within the superfamily Chactoidea. Recent phylogenomic analyses have instead recovered vaejovids as polyphyletic (albeit with minimal taxonomic sampling) and Chactoidea as paraphyletic. Here, we reexamined the monophyly and phylogenetic placement of the family Vaejovidae, sampling 17 new vaejovid libraries using high throughput transcriptomic sequencing. Our phylogenomic analyses revealed a previous misplacement of Smeringurus mesaensis. Regardless, we recovered Vaejovidae as diphyletic due to the placement of the enigmatic genus Uroctonus. The remaining vaejovids formed a clade that was strongly supported as the sister group of the superfamily Scorpionoidea, a placement insensitive to matrix completeness or concatenation vs. species tree approaches to inferring the tree topology. Chactoidea was invariably recovered as a paraphyletic group due to the nested placement of Scorpionoidea. As first steps to resolving the paraphyly of Chactoidea, we take the following systematic actions: (1) we establish the superfamily Superstitionoidea (new superfamily) to accommodate Superstitioniidae; (2) we restore Vaejovoidea (status revalidated) as a valid superfamily that excludes Uroctonus; and (3) we treat the families Caraboctonidae, Troglotayosicidae, and the subfamily Uroctoninae as incertae sedis with respect to superfamilial placement. Our systematic actions thus establish the monophyly of the presently redefined Chactoidea and Vaejovoidea.