Reevaluation ofBlapimorpha and Opatrinae: addressing a major phylogeny‐classification gap in darkling beetles (Coleoptera: Tenebrionidae: Blaptinae)

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.

On the type species of the genus Dendarus Dejean, 1821 (Coleoptera: Tenebrionidae)

“…They used "Dendarus tristis Laporte, 1840" as a valid name (with Pandarus coarcticollis Mulsant, 1854 as a synonym) and considered it to be the type of Dendarus. Many subsequent authors followed this concept (Gebien 1938, Español 1961, Medvedev 1968, Chatzimanolis & Löbl 2003, Cherney 2005, Iwan et al 2020. However, as mentioned before "D. tristis Laporte" was not originally included in Dendarus by Dejean (1821) and therefore cannot be considered as a type species of that genus.…”

“…Several authors considered Pandarus libanicus Desbrochers des Loges, 1881 as a junior synonym of D. lugens Mulsant & Rey, 1854(e.g., Gebien 1938, Löbl et al 2008, Iwan et al 2020. However, as noted by Koch (1948) Pandarus libanicus is in fact a synonym of Dendarus calcaratus Baudi di Selve, 1881.…”

“…Dendarus Dejean, 1821 (Coleoptera: Tenebrionidae) is a well-known genus of flightless darkling beetles inhabiting the Western Palearctic Region (Trichas et al 2020). The genus is represented by 89 species and subspecies classified in six subgenera (Iwan et al 2020, Bouchard et al 2021, and is currently placed within the tribe Dendarini Mulsant & Rey, 1854 of the subfamily Blaptinae Leach, 1815 (Kamiński et al 2020). Throughout history, several contributors questioned the validity of the type species fixation for Dendarus (see Löbl et al 2008, Iwan et al 2020).…”

“…The genus is represented by 89 species and subspecies classified in six subgenera (Iwan et al 2020, Bouchard et al 2021, and is currently placed within the tribe Dendarini Mulsant & Rey, 1854 of the subfamily Blaptinae Leach, 1815 (Kamiński et al 2020). Throughout history, several contributors questioned the validity of the type species fixation for Dendarus (see Löbl et al 2008, Iwan et al 2020). However, despite intensive taxonomic work on the genus this problem has not yet been solved.…”

Distribution and environmental determinants of darkling beetles assemblages (Coleoptera: Tenebrionidae) in Península Valdés (Argentinean Patagonia)

“…reviews and keys by Kulzer (1955Kulzer ( , 1963, Flores (1999Flores ( , 2004, Flores et al (2011) and comparisons with material deposited in the IADIZA collection. We followed the classification proposed by Matthews et al (2010) and Kamiński et al (2021) to assign species and genera into subfamilies and tribes. Following the recent taxonomic changes suggested by , Nyctelia nodosa (Germar, 1823) mentioned in previous works (Cheli et al 2010, 2013, Carrara et al 2011b (Cheli et al 2010, 2013, Flores et al 2011, Carrara et al 2011b, Martínez et al 2018) and the collecting campaign, a matrix of species incidence by environmental unit was constructed (Table II) to perform all statistical analyses.…”