Head morphology of the smallest beetles (Coleoptera: Ptiliidae) and the evolution of sporophagy within Staphyliniformia

Abstract

Ptiliidae include the smallest known beetles. External and internal head structures of species with different body sizes and feeding preferences were examined and described in detail. Saprophagous and sporophagous species are compared. The observed features are evaluated with respect to their phylogenetic and functional significance, and their correlation with extreme size reduction. A putative autapomorphy of Staphyliniformia is an unusual extrinsic maxillary muscle, which among ptiliids is only present in the saprophagous species. Synapomorphies of Ptiliidae and their sister group Hydraenidae are a lateral mandibular process forming a unique locking device with a lateral groove of the labrum, and mandibles divided into a main body and a mesal molar part, both connected by a membrane. Extreme body size reduction is a presumptive autapomorphy of Ptiliidae that probably resulted in the following derived features: the loss of cephalic sutures and ridges, a simplified tentorium, and a brain modified in shape and very large in relation to the head size. The ptiliid species with saprophagous and sporophagous feeding habits show only subtle differences in their cephalic structures, notably in details of the epipharynx and galeae and in the configuration of maxillary muscles. Two alternative scenarios are suggested for the evolution of feeding habits, based on the morphological results and presently available information on phylogenetic relationships. One option is to assign saprophagy to the groundplan of the family, with two switches to sporophagy; first in the basal Nossidium and then a second time in the extremely small Nanosellini, which are characterized by feeding habits that we address as microsporophagy. An alternative scenario is that feeding on spores is ancestral for Ptiliidae, with reversals to saprophagy in several branches of the family, and a specialization on very small spores in the strongly miniaturized nanoselline species. A well-founded species level phylogeny of Ptiliidae with a dense taxon sampling will help to clarify this issue.