No abstract

The taxonomic concepts of Blapimorpha and Opatrinae (informal and traditional, morphology‐based groupings among darkling beetles) are tested using molecular phylogenetics and a reassessment of larval and adult morphology to address a major phylogeny‐classification gap in Tenebrionidae. Instead of a holistic approach (family‐level phylogeny), this study uses a bottom‐up strategy (tribal grouping) in order to define larger, monophyletic lineages within Tenebrioninae. Sampling included representatives of 27 tenebrionid tribes: Alleculini, Amarygmini, Amphidorini, Blaptini, Bolitophagini, Branchini, Cerenopini, Coniontini, Caenocrypticini, Dendarini, Eulabini, Helopini, Lagriini, Melanimini, Opatrini, Pedinini, Phaleriini, Physogasterini, Platynotini, Platyscelidini, Praociini, Scaurini, Scotobiini, Tenebrionini, Trachyscelini, Triboliini and Ulomini. Molecular analyses were based on DNA sequence data from four non‐overlapping gene regions: carbamoyl‐phosphate synthetase domain of rudimentary (CAD) (723 bp), wingless (wg) (438 bp) and nuclear ribosomal 28S (1101 bp) and mitochondrial ribosomal 12S (363 bp). Additionally, 15 larval and imaginal characters were scored and subjected to an ancestral state reconstruction analysis. Results revealed that Amphidorini, Blaptini, Dendarini, Pedinini, Platynotini, Platyscelidini and Opatrini form a clade which can be defined by the following morphological features: adults—antennae lacking compound/stellate sensoria; procoxal cavities externally and internally closed, intersternal membrane of abdominal ventrites 3–5 visible; paired abdominal defensive glands present, elongate, not annulated; larvae—prolegs enlarged (adapted for digging); ninth tergite lacking urogomphi. To accommodate this monophyletic grouping (281 genera and ∼4000 species), the subfamily Blaptinae sens. nov. is resurrected. Prior to these results, all of the tribes within Blaptinae were classified within the polyphyletic subfamily Tenebrioninae. The non‐monophyletic nature of Terebrioninae has already been postulated by previous authors, yet no taxonomic decisions were made to fix its status. The reinstatement of Blaptinae, which groups ∼50% of the former Tenebrioninae, helps to clarify phylogenetic relations among the whole family and is the first step towards a complete higher‐level revision of Tenebrionidae. The Central Asian tribe Dissonomini (two genera, ∼30 species) was not included in Blaptinae due to a lack of representatives in the performed phylogenetic analyses; however, based on morphological features, the tribe is listed as a potential addition to the subfamily.

This paper summarizes currently available morphological data on larval stages of representatives of the 'Opatrinoid' clade (Tenebrionidae: Tenebrioninae). Literature research revealed that larval morphology of approximately 6% of described species representing this lineage is currently known (139 out of ~ 2325 spp.). Larvae of the five following species are described and illustrated: Zadenos mulsanti (Dendarini: Melambiina; South Africa), Blapstinus histricus, Blapstinus longulus, Trichoton sordidum (Opatrini: Blapstinina; North America), and Eurynotus rudebecki (Platynotini: Eurynotina; South Africa). The majority of studied larvae were associated with adults using molecular tools, resulting in an updated phylogeny of the 'Opatrinoid' clade. This revised phylogeny provides an evolutionary context for discussion of larval morphology. Based on the morphological and molecular evidence, the following synonym is proposed within Blapstinina: Trichoton Hope, 1841 (= Bycrea Pascoe, 1868 syn. nov.). Based on this decision, a new combination is introduced: Trichoton villosum Pascoe, 1868 comb. nov. The economic importance of the 'Opatrinoid' clade larvae is also briefly discussed, as well as potential future avenues of research.

The monophyly of the North and South American endemic subtribe Blapstinina (Tenebrionidae: Opatrini) is tested through phylogenetic analyses using five molecular markers [nuclear ribosomal 28S (28S), cytochrome oxidase subunit II (COII), arginine kinase (ArgK), carbamoyl‐phosphate synthetase domain of rudimentary (CAD), wingless (wg)]. Representatives of several opatrinoid subtribes were taken into consideration, including other geographically overlapping endemic genera, namely Ammodonus, Ephalus and Pseudephalus (all previously considered representatives of Ammobiina). A comparative study of morphology was performed to assess resulting phylogenetic hypotheses. Analyses support the monophyly of Blapstinina; however, they also strongly indicate that Ammodonus should be included within the subtribe. Mecysmus is nested within Blapstinus and therefore, a new synonymy, Blapstinus (= Mecysmus syn.n.), and the following combinations are introduced: Blapstinus advena comb.n., B. angustus comb.r., B. laticollis comb.n., B. parvulus comb.n., B. tenuis comb.n. Morphological analysis showed a close affiliation between Ephalus and Pseudephalus. Based on these results, Pseudephalus is synonymized with Ephalus [Ephalus (= Pseudephalus syn.n.)], and the following combination is introduced: Ephalus brevicornis comb.n. Recovered topologies also strongly support transferring Ephalus stat.n. into Opatrina, making the distribution of Opatrina amphi‐Atlantic.