The first molecular phylogeny of the tribe Sepidiini is inferred from analyses of DNA sequence data from the following five loci (CAD, wg, COI, COII, 28S rRNA). Bayesian and maximum likelihood analyses were performed on a dataset containing 41 taxa, of which a majority represent Molurina (27) and Phanerotomeina (6). The resulting topologies were used to discuss phylogenetic placement and diagnostic characters of all of the genera representing Molurina. Within the subtribe, the results revealed paraphyly of the genus Psammodes. The P. vialis species-group, currently classified within Psammodes, was recovered as sister to all other Molurina genera. Based on this topology and morphological investigations, a new genus named Toktokkus gen. nov. is established. Within Phanerotomeina, Ocnodes is paraphyletic with regard to Tarsocnodes. In order to restore the monophyly of Ocnodes, the subgenus Chiliarchum stat. nov. is elevated to generic level. Finally, as the homology of female terminalia structures has never been fully assessed for Sepidiini, a comparative study of ovipositor morphology was conducted. As a result, this paper presents the first fully annotated ovipositors for tok-tokkie beetles.
Sepidiini is a speciose tribe of desert-inhabiting darkling beetles, which contains a number of poorly defined taxonomic groups and is in need of revision at all taxonomic levels. In this study, two previously unrecognized lineages were discovered, based on morphological traits, among the extremely speciose genera Psammodes Kirby, 1819 (164 species and subspecies) and Ocnodes Fåhraeus, 1870 (144 species and subspecies), namely the Psammodes spinosus species-group and Ocnodes humeralis species-group. In order to test their phylogenetic placement, a phylogeny of the tribe was reconstructed based on analyses of DNA sequences from six nonoverlapping genetic loci (CAD, wg, COI JP, COI BC, COII, and 28S) using Bayesian and maximum likelihood inference methods. The aforementioned, morphologically defined, species-groups were recovered as distinct and well-supported lineages within Molurina + Phanerotomeina and are interpreted as independent genera, respectively, Tibiocnodes Gearner & Kamiński gen. nov. and Tuberocnodes Gearner & Kamiński gen. nov. A new species, Tuberocnodes synhimboides Gearner & Kamiński sp. nov., is also described. Furthermore, as the recovered phylogenetic placement of Tibiocnodes and Tuberocnodes undermines the monophyly of Molurina and Phanerotomeina, an analysis of the available diagnostic characters for those subtribes is also performed. As a consequence, Phanerotomeina is considered as a synonym of the newly redefined Molurina sens. nov. Finally, spectrograms of vibrations produced by substrate tapping of two Molurina species, Toktokkus vialis (Burchell, 1822) and T. synhimboides, are presented.
Tok-tokkies are one of the most iconic lineages within Tenebrionidae. In addition to containing some of the largest darkling beetles, this tribe is recognized for its remarkable form of sexual communication known as substrate tapping. Nevertheless, the phylogenetic relationships within the group remain poorly understood. This study investigates the usefulness of female terminalia morphology for delimiting Sepidiini and reconstructing relationships among it. Data on the structure of the ovipositors, genital tubes and spicula ventrali have been generated for >200 species representing 28 Pimeliinae tribes. This dataset was used in a comparative analysis at the subfamilial level, which resulted in recognition of several unique features of tok-tokkie terminalia. Additionally, new features linking phenotypically challenging tribes also were recovered (Cryptochilini + Idisiini + Pimeliini). Secondly, 23 characters linked to the structure of female terminalia were defined for tok-tok beetles. Cladistic analysis demonstrates the nonmonophyletic nature of most of the recognized subtribes. The morphological dataset was analysed separately and in combination with available molecular data (CAD, Wg, cox1, cox2, 28S). All obtained topologies were largely congruent, supporting the following changes: Palpomodina Kami nski & Gearner subtr.n. is erected to accommodate the genera Namibomodes and Palpomodes; Argenticrinis and Bombocnodulus are transferred from Hypomelina to Molurina; 153 species and subspecies previously classified within Psammodes are distributed over three separate genera (Mariazofia Kami nski nom.n., Piesomera stat.r., Psammodes sens.n.). Psammodes sklodowskae Kami nski & Gearner sp.n. is described. Preliminary investigation of the ovipositor of Mariazofia basuto (Koch) comb.n. was carried out with the application of microcomputed tomography, illuminating the muscular system as a reliable reference point for recognizing homologous elements in highly modified ovipositors.
The darkling beetle tribe Adesmiini (Tenebrionidae: Pimeliinae) is a prominent part of African and western Palearctic desert faunas, with most species being day-active fast-running detritivores. Taxonomic diversity within the tribe is highest in the southern Afrotropical realm (where all genera are present); only 1 genus, the species-rich Adesmia, occurs north of the Sahara. Despite notable species, such as the fog-basking beetle Onymacris unguicularis (a focal taxon in desert ecological research), Adesmiini has undergone few modern taxonomic or phylogenetic studies. Hence, generic concepts and pronounced diurnal activity, rare in the primarily nocturnal family Tenebrionidae, remain poorly explored. To investigate evolutionary relationships and diurnal origins within the tribe, we generated a genomic dataset of 529 protein-coding genes across 43 species spanning 10 of 11 Adesmiini genera. Our resulting phylogeny for the tribe rejects the monophyly of 5 currently recognized Adesmiini genera (i.e., Adesmia, Metriopus, Onymacris, Physadesmia, and Stenocara). Ancestral state reconstruction of diurnal activity using eye shape as a proxy supports the hypothesis that Adesmiini were primitively diurnal, followed by at least 4 shifts to nocturnal or crepuscular activity.
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