Click Beetle Mitogenomics with the Definition of a New Subfamily
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insects Article Click Beetle Mitogenomics with the Definition of a New Subfamily Hapatesinae from Australasia (Coleoptera: Elateridae) Dominik Kusy † , Michal Motyka † and Ladislav Bocak * Laboratory of Biodiversity and Molecular Evolution, CATRIN-CRH, Palacky University, 17. listopadu 50, 771 46 Olomouc, Czech Republic; [email protected] (D.K.); [email protected] (M.M.) * Correspondence: [email protected] † These authors contributed equally to this work. Simple Summary: The classification of click beetles is revisited with newly sequenced mitochondrial genomes of eight species belonging to seven different subfamilies. The genus Hapatesus Candèze, 1863, is herein excluded from Dentrometrinae and designated as a type genus of Hapatesinae, a new subfamily. Phylogenetic analyses suggest that Eudicronychinae is a terminal lineage in Elaterinae. Consequently, we propose Eudicronychini, a new status. The deep mtDNA-based split between Elaterinae and the clade of other click beetle subfamilies agrees with the results of phylogenomic analyses and mitochondrial genomes provide a sufficient signal for inference of shallow splits. Abstract: Elateridae is a taxon with very unstable classification and a number of conflicting phy- logenetic hypotheses have been based on morphology and molecular data. We assembled eight complete mitogenomes for seven elaterid subfamilies and merged these taxa with an additional 22 elaterids and an outgroup. The structure of the newly produced mitogenomes showed a very similar arrangement with regard to all earlier published mitogenomes for the Elateridae. The maximum likelihood and Bayesian analyses indicated that Hapatesus Candèze, 1863, is a sister of Parablacinae Citation: Kusy, D.; Motyka, M.; and Pityobiinae. Therefore, Hapatesinae, a new subfamily, is proposed for the Australian genera Bocak, L. Click Beetle Mitogenomics Hapatesus (21 spp.) and Toorongus Neboiss, 1957 (4 spp.). Parablacinae, Pityobiinae, and Hapatesinae with the Definition of a New have a putative Gondwanan origin as the constituent genera are known from the Australian region Subfamily Hapatesinae from (9 genera) and Neotropical region (Tibionema Solier, 1851), and only Pityobius LeConte, 1853, occurs Australasia (Coleoptera: Elateridae). in the Nearctic region. Another putative Gondwanan lineage, the Afrotropical Morostomatinae, Insects 2021, 12, 17. https://doi.org/ forms either a serial paraphylum with the clade of Parablacinae, Pityobiinae, and Hapatesinae or 10.3390/insects12010017 is rooted in a more terminal position, but always as an independent lineage. An Eudicronychinae lineage was either recovered as a sister to Melanotini or as a deep split inside Elaterinae and we Received: 8 December 2020 herein transfer the group to Elaterinae as Eudicronychini, a new status. The mitochondrial genomes Accepted: 23 December 2020 Published: 29 December 2020 provide a sufficient signal for the placement of most lineages, but the deep bipartitions need to be Revised: 6 January 2021 compared with phylogenomic analyses. Publisher’s Note: MDPI stays neu- Keywords: taxonomy; mitochondrial genomes; phylogeny; Gondwana; Australian region; tral with regard to jurisdictional claims new subfamily; new status in published maps and institutional affiliations. 1. Introduction Copyright: © 2020 by the authors. Li- Click beetles, Elateridae, are a major elateroid family with ~10,000 recognized species censee MDPI, Basel, Switzerland. This from all zoogeographic regions. The group is well-known even to non-specialists due article is an open access article distributed to its common occurrence in all ecosystems and characteristic clicking mechanism [1] under the terms and conditions of the (Figure1A–D). The clicking elateroids are morphologically very uniform and they resemble Creative Commons Attribution (CC BY) the general appearance of false click beetles (Eucnemidae; Figure1E) and small false click license (https://creativecommons.org/ beetles (Throscidae) which are distantly related [2]. The characteristic slender body shape, licenses/by/4.0/). rectangular to acutely projected posterior angles of the pronotum, and short slender legs Insects 2021, 12, 17. https://doi.org/10.3390/insects12010017 https://www.mdpi.com/journal/insects Insects 2021, 12, x FOR PEER REVIEW 2 of 17 Insects 2021, 12, 17 2 of 17 beetles (Throscidae) which are distantly related [2]. The characteristic slender body shape, rectangular to acutely projected posterior angles of the pronotum, and short slender legs andand antennae,antennae, bothboth often partly sunk sunk in in grooves grooves in in a a resting resting position, position, are are characteristic characteristic of ofmost most elaterids elaterids (Figure (Figure 1A–D).1A–D). A Adependence dependence on onclicking clicking as an as anantipredator antipredator strategy strategy pos- possiblysibly constrains constrains the themorphological morphological evolut evolutionion of the of thegroup group because because the themechanism mechanism de- dependspends on on precise precise interaction interaction between between pro- pro- and and mesothoracic mesothoracic structures structures that thatkeep keep the body the bodyin a brace in a brace position position until untiltheir theirrelease release [3,4].[ 3Some,4]. Some forms forms similar similar to extant to extant taxa taxahave have been beenreported reported already already from from the Late the LateTriassic Triassic and andEarly Early Jurassic, Jurassic, the theearly-Cretaceous early-Cretaceous Jinju Jinju for- formation,mation, and and Burmese, Burmese, Dominican, Dominican, and and Baltic Baltic amber amber inclusions inclusions [5–8]. [5–8 ]. FigureFigure 1.1.Clicking Clicking elateroids elateroids in in nature. nature. (A –(AD–)D click) click beetles, beetles, Elateridae; Elateridae; (E) false (E) false click click beetle, beetle, Eucnemidae. Eucnemidae. Despite high morphological uniformity (Figure1), the earlier students of click beetles describedDespite a number high morphological of family group uniformity taxa, i.e., (Figure subfamilies, 1), the tribes, earlier and students subtribes of click (altogether beetles >130described taxa; a [ 9number]), but theirof family relationships group taxa, have i.e., subfamilies, remained contentious, tribes, and subtribes reliable diagnostic(altogether characters>130 taxa; [9]), are absent,but their constituent relationships genera have areremained often transferredcontentious, between reliable diagnostic subfamilies char- or tribes,acters andare absent, the ranks constituent of individual genera taxa are are often unstable transferred [1,10]. between Not only subfamilies morphology-based or tribes, hypothesesand the ranks on relationships,of individual taxa but alsoare unstable Sanger molecular [1,10]. Not data only have morphology-based provided weak supporthypoth- foreses the on click relationships, beetle classification, but also Sanger and additionally,molecular data recovered have provided topologies weak were support in conflict for the withclickprevious beetle classification, definitions ofand separate additionally, “cantharoid” recovered and topologies “elateroid” were groups in conflict of families with [ pre-11]. Thevious morphological definitions of delimitation separate “cantharoid” of click beetles and has “elateroid” been recently groups seriously of families questioned [11]. The by themorphological inclusion of somedelimitation soft-bodied, of click morphologically beetles has been highly recently divergent seriously “cantharoid” questioned lineages by the intoinclusion Elateridae. of some Already, soft-bodied, Bocakova morphologica et al. [2] showedlly highly that divergentDrilus Olivier, “cantharoid” 1790, believed lineages atinto the Elateridae. time to be Already, a genus ofBocakova the family et al. Drilidae [2] showedsensu Crowsonthat Drilus [12 Olivier,], might 1790, be a believed modified at clickthe time beetle to andbe a itsgenus position of the in family the Agrypninae Drilidae sensu was Crowson confirmed [12], with might further be mito-ribosomala modified click databeetle [13 and,14 ].its Later, position Plastoceridae in the Agrypninae were transferred was confirmed to Elateridae with further and mito-ribosomal given the rank ofdata a subfamily[13,14]. Later, [15]. Plastoceridae Due to limited were statistical transferred support to ofElateridae mito-ribosomal and given phylogenies the rank forof a deep sub- relationshipsfamily [15]. Due [2,13 to–16 limited], the resultsstatistical were support considered of mito-ribosomal by most students phylogenies of click beetles for deep with re- reluctancelationships [ 17[2,13–16],,18], see the [19 ]results for further were information.considered by Therefore, most students the Sanger of click data beetles for click with beetlesreluctance have [17,18], not recently see [19] been for intensively further information. produced, and Therefore, only a few the sequences Sanger data for elateridfor click taxabeetles have have been not reported recently inbeen the intensively last years toproduced, identify and relationships only a few of sequences some species-poor for elaterid lineages,taxa have e.g., been Pityobiinae reported in and the Parablacinae last years to [ 20identify]. Similarly, relationships a limited of number some species-poor of protein- codinglineages, nuclear e.g., Pityobiinae genes did and not performParablacinae well, [20]. and Similarly, click beetles a limited were notnumber identified of protein- as a monophylumcoding nuclear when genes