Untangling the evolution of soldier beetles (Coleoptera: Cantharidae) and the evaluation of the morphological phylogenetic signal in a soft-bodied elateroid lineage.

This study addresses the long-standing uncertainty about the internal classification of soldier beetles (Elateroidea: Cantharidae). Four datasets were compiled and analysed: 66 genes for 14 terminals, 15 mtDNA genes for 79 terminals, one mtDNA and two rRNA genes for 217 terminals, and barcodes for 576 terminals. Based on congruent topologies, Chauliognathinae is proposed as a sister to the remaining Cantharidae, followed by the redefined Malthininae (including Tytthonyxini), the paraphyletic "dysmorphocerine" lineages (Dysmorphocerinae sensu stricto and Heteromastiginae subfam. nov.), and Silinae + Cantharinae as a terminal clade. The present phylogeny supersedes earlier morphology and short-fragment molecular hypotheses that have not converged on a consensus. Few morphological characters corroborate the DNA-based relationships (see the adults and larval keys). However, morphology-based hypotheses have relied on a few informative characters, and no evidence strongly rejects the preferred molecular topology. The interpretation of morphological characters and uncertain polarity resulting from the high phenotypic disparity of Elateroidea are discussed in detail. The dated phylogeny hypothesizes the earliest split within the Cantharidae in the Berriasian stage (Early Cretaceous, ~141 Myr) and the diversification of most extant subfamilies and tribes already in the Late Cretaceous. The most diverse subfamily, Cantharinae, represents a delayed radiation that started during the Eocene climatic optimum, 55.5 Myr. The late origin of Cantharinae questions the classification of Cretaceous Cantharidae as members of Cantharinae. Instead, the results suggest their deeper rooting after separating from dysmorphocerine lineages and before the node between Cantharinae and Silinae.

Beetle bioluminescence outshines extant aerial predators.

We understand very little about the timing and origins of bioluminescence, particularly as a predator avoidance strategy. Understanding the timing of its origins, however, can help elucidate the evolution of this ecologically important signal. Using fireflies, a prevalent bioluminescent group where bioluminescence primarily functions as aposematic and sexual signals, we explore the origins of this signal in the context of their potential predators. Divergence time estimations were performed using genomic-scale datasets providing a robust estimate for the origin of firefly bioluminescence as both a terrestrial and as an aerial signal. Our results recover the origin of terrestrial beetle bioluminescence at 141.17 (122.63-161.17) Ma and firefly aerial bioluminescence at 133.18 (117.86-152.47) Ma using a large dataset focused on Lampyridae; and terrestrial bioluminescence at 148.03 (130.12-166.80) Ma, with the age of aerial bioluminescence at 104.97 (99.00-120.90) Ma using a complementary Elateroidea dataset. These ages pre-date the origins of all known extant aerial predators (i.e. bats and birds) and support much older terrestrial predators (assassin bugs, frogs, ground beetles, lizards, snakes, hunting spiders and harvestmen) as the drivers of terrestrial bioluminescence in beetles. These ages also support the hypothesis that sexual signalling was probably the original function of this signal in aerial fireflies.

Persistence of multiple patterns and intraspecific polymorphism in multi-species Müllerian communities of net-winged beetles.

BACKGROUND: In contrast to traditional models of purifying selection and a single aposematic signal in Müllerian complexes, some communities of unprofitable prey contain members with multiple aposematic patterns. Processes responsible for diversity in aposematic signaling are poorly understood and large multi-species communities are seldom considered. RESULTS: We analyzed the phylogeny and aposematic patterns of closely related Eniclases net-winged beetles in New Guinea using mtDNA and nextRAD data. We suggest three clades of closely related and incompletely reproductively isolated lineages, detail the extent of polymorphism among Eniclases, and categorize their low-contrast aposematic patterns. The warning signal of Eniclases consists of body shape and color, with ambiguous color perception under some circumstances, i.e., when resting on the undersides of leaves. Field observations suggest that perception of the aposematic signal is affected by beetle behavior and environmental conditions. Local communities containing Eniclases consisted of 7-85 metriorrhynchine species assigned to 3-10 colour patterns. CONCLUSION: As a result, we suggest that under certain light conditions the aposematic colour signal is less apparent than the body shape in net-winged beetle communities. We document variable environmental factors in our study area and highly diverse multi-species communities of other net-winged beetles. Which implies dynamically changing community structure in space and time. Variable environmental conditions and diverse community composition are suggested to be favorable for the persistence of multiple aposematic patterns, imperfect mimics, and intraspecific polymorphism. Further research should identify the relative effect of these factors on purifying selection and the alleles which are responsible for phenotypic differences.

The contribution of mitochondrial metagenomics to large-scale data mining and phylogenetic analysis of Coleoptera.

A phylogenetic tree at the species level is still far off for highly diverse insect orders, including the Coleoptera, but the taxonomic breadth of public sequence databases is growing. In addition, new types of data may contribute to increasing taxon coverage, such as metagenomic shotgun sequencing for assembly of mitogenomes from bulk specimen samples. The current study explores the application of these techniques for large-scale efforts to build the tree of Coleoptera. We used shotgun data from 17 different ecological and taxonomic datasets (5 unpublished) to assemble a total of 1942 mitogenome contigs of >3000 bp. These sequences were combined into a single dataset together with all mitochondrial data available at GenBank, in addition to nuclear markers widely used in molecular phylogenetics. The resulting matrix of nearly 16,000 species with two or more loci produced trees (RAxML) showing overall congruence with the Linnaean taxonomy at hierarchical levels from suborders to genera. We tested the role of full-length mitogenomes in stabilizing the tree from GenBank data, as mitogenomes might link terminals with non-overlapping gene representation. However, the mitogenome data were only partly useful in this respect, presumably because of the purely automated approach to assembly and gene delimitation, but improvements in future may be possible by using multiple assemblers and manual curation. In conclusion, the combination of data mining and metagenomic sequencing of bulk samples provided the largest phylogenetic tree of Coleoptera to date, which represents a summary of existing phylogenetic knowledge and a defensible tree of great utility, in particular for studies at the intra-familial level, despite some shortcomings for resolving basal nodes.

Genome sequencing of Rhinorhipus Lawrence exposes an early branch of the Coleoptera.

BACKGROUND: Rhinorhipidae Lawrence, 1988 is an enigmatic beetle family represented by a single species, Rhinorhipus tamborinensis Lawrence, 1988, from Australia, with poorly established affinities near the superfamily Elateroidea (click beetles, soldier beetles and fireflies) or the more inclusive series (infraorder) Elateriformia. Its evolutionary position may inform the basal relationships of the suborder Polyphaga, the largest clade of Coleoptera. RESULTS: We analyzed four densely sampled DNA datasets of major coleopteran lineages for mitogenomes, rRNA genes and single copy nuclear genes. Additionally, genome sequencing was used for incorporation of R. tamborinensis into a set of 4220 orthologs for 24 terminals representing 12 polyphagan superfamilies. Topologies differed to various degrees, but all consistently refute the proposed placement of Rhinorhipidae in Elateroidea and instead indicate either sister relationships with other Elateriformia, frequently together with Nosodendridae, another divergent small family hitherto placed in Derodontoidea, or in an isolated position among the deepest lineages of Polyphaga. The phylogenomic analyses recovered Rhinorhipus in a sister position to all other Elateriformia composed of five superfamilies. Therefore, we erect the new superfamily Rhinorhipoidea Lawrence, 1988, stat. Nov., with the type-family Rhinorhipidae. The origins of the Rhinorhipidae were dated to the Upper Triassic/Lower Jurassic at the very early phase of polyphagan diversification. CONCLUSIONS: Thus, Rhinorhipidae adds another example to several recently recognized ancient relict lineages which are interspersed within contemporaneous hugely species-rich lineages of Coleoptera.

The comprehensive phylogeny of the superfamily Elateroidea (Coleoptera: Elateriformia).

Elateriformia consists of Dascilloidea, Buprestoidea (jewel beetles), Byrrhoidea and Elateroidea (click beetles, fireflies and relatives). Numerous elateroid lineages contain taxa with modified metamorphosis resulting in sexual maturity while retaining larval characters. Additionally, they evolved unique defensive strategies including clicking mechanism, aposematic coloration and bioluminescence. To investigate the phylogenetic position of Elateroidea within Coleoptera, we merged 1048 newly produced 18S rRNA, 28S rRNA, rrnL mtDNA, and cox1 mtDNA sequences for ∼300 elateriform taxa with data from GenBank. The 975-taxa dataset aligned in BlastAlign was analyzed under maximum likelihood criterion. The results agreed in most aspects with the current morphology-based classification and results of molecular studies. Elateriformia were monophyletic and Elateroidea were sister to Byrrhoidea. Further, we analyzed all-data (513 elateriform taxa) and pruned matrix (417 elateriform taxa, all fragments present) using parsimony and maximum likelihood methods to reveal the phylogenetic relationships among elateroid lineages and examine the evolution of soft-bodiedness, neoteny and bioluminescence. We confirmed the monophyly of Elateroidea sensu lato and most of the families, with Telegeusidae inferred in most trees within paraphyletic Omethidae. The clade Artematopodidae+Telegeusidae+Omethidae was a sister to remaining elateroids. All topologies reject the relationships of hard-bodied Elateridae, Eucnemidae, Throscidae and Cerophytidae, formerly supposed to be a monophylum. Eucnemidae and Throscidae formed independent lineages and the position of Cerophytidae was variable - either a sister to Throscidae, or an independent lineage. The Lampyridae+Cantharidae clade was in most trees sister to Phengodidae+Rhagophthalmidae+Omalisidae+Elateridae. Molecular phylogeny of Elateroidea confirmed the multiple origins of soft-bodied, neotenic and light emiting lineages. On the basis of our molecular phylogeny, we place former Telegeusidae as a subfamily in Omethidae.

Towards a phylogeny of the Tenebrionoidea (Coleoptera).

The phylogenetic relationships of the beetle superfamily Tenebrionoidea are investigated using the most comprehensive genetic data set compiled to date. With ∼34,000 described species in approximately 1250 genera and 28 families, Tenebrionoidea represent one of the most diverse and species-rich superfamilies of beetles. The interfamilial relationships of the Tenebrionoidea are poorly known; previous morphological and molecular phylogenies recovered few well-supported and often conflicting relationships between families. Here we present a molecular phylogeny of Tenebrionoidea based on genes commonly used to resolve family and superfamily-level phylogenies of beetles (18S, 28S, 16S, 12S, tRNA Val and COI). The alignment spanned over 6.5KB of DNA sequence and over 300 tenebrionoid genera from 24 of the 28 families were sampled. Maximum Likelihood and Bayesian analysis could not resolve deeper level divergences within the superfamily and very few relationships between families were supported. Increasing gene coverage in the alignment by removing taxa with missing data did not improve clade support but when rogue taxa were removed increased resolution was recovered. Investigation of signal strength suggested conflicting phylogenetic signal was present in the standard genes used for beetle phylogenetics, even when rogue taxa were removed. Our study of Tenebrionoidea highlights that even with relatively comprehensive taxon sampling within a lineage, this standard set of genes is unable to resolve relationships within this superfamily.

A review of Drilini (Coleoptera: Elateridae: Agrypninae) of the Northern Levant, with description of a new species from Syria and a key to Levantine species.

The species of the elaterid soft-bodied tribe Drilini from the Levant are reviewed. All known species are redescribed and Drilus nemethi sp. nov. is described. Drilus adustus (Chevrolat, 1854), comb. nov. and D. akbesianus (Fairmaire, 1895), comb. nov. are transferred from Malacogaster Bassi, 1834 to Drilus Olivier, 1790. Drilus posticus Schaufuss, 1867, syn. nov. is a junior subjective synonym of Drilus adustus (Chevrolat, 1854). All Levantine species are keyed and diagnostic characters are illustrated. Their distribution, morphological diversity and the statuses of type specimens of Drilus reitteri Bourgeois, 1908 and Cydistus reitteri Bourgeois, 1885 are discussed.

A new species of Dexoris (Coleoptera: Lycidae) and parallel evolution of brachyptery in the soft-bodied elateroid beetles.

Dexoris chome sp. nov. is described from South Pare mountains, Tanzania, based on a male specimen. This is the only Dexoris with shortened elytra, rudimentary hind wings and large, larviform abdomen. Unlike males of other species in the genus, D. chome sp. nov. has a modified number of palpomeres and remarkably short, robust legs and antennae. Such modifications are similar to those in the neotenic female of Omalisus fontisbellaquei Fourcroy, 1785 (Omalisidae), suggesting analogous morphological changes in unrelated lineages supposedly caused by similar modifications of their metamorphosis. The distribution of all 11 known species of African Dexoris closely overlap with the location of the hypothesized centres for evolution of new species in the Afrotropical region.

Paussus (Edaphopaussus) asseyi sp. nov. (Coleoptera: Carabidae: Paussinae) from the Nguru Mountains of the Eastern African Arc in Tanzania.

A new species of Paussus L., 1775 (Coleoptera: Carabidae) is described from the Nguru Mountains of the Eastern African Arc in Tanzania. Paussus asseyi sp. nov. is diagnosed by the shape of the antennal club, which bears two sharp edges at the outer margin, the blunt spines on tibiae, and the characteristic colouration that allows this species to be distinguished from all other species in Paussus (Series I) defined in the latest molecular phylogeny. Other characters, i.e., the shape of the pronotum and absent pronotal setae, place the new species in the subgenus Edaphopaussus Kolbe, 1920. The colouration resembles P. dichrous Janssens, 1950 from Democratic Republic Congo and Angola and some Asian species, yet all are morphologically different from the new species. The robust head, small eyes, and clypeus shape resemble distantly related P. (Hylotorus) cephalotes Raffray, 1886. The individual mountain massifs of Eastern Tanzania represent rainforest islands set in much drier lowlands with savannah ecosystems. The new species contributes to our understanding of the isolated mountain ranges' relatively poorly investigated beetle fauna.

New Species of Paussus, Subgenus Scaphipaussus (Coleoptera: Carabidae: Paussinae), from Southeast Asia Reveal Ambiguities in Species Group Limits and High Species Diversity in the Oriental Region.

Paussus, commonly known as ant nest beetles, is the most diverse genus of Paussinae (Coleoptera: Carabidae) with a very complex taxonomic history. Biodiversity research in Southeast and South Asia yields new species that can contribute to a better understanding of the morphological disparity and species-group or subgenus delimitation. Here, we describe nine new species from Southeast Asia and China: Paussus (Scaphipaussus) fencli sp. nov. (China), P. (S.) mawdsleyi sp. nov. (Borneo), P. (S.) bakeri sp. nov. (Philippines), P. (S.) jendeki sp. nov. (Laos), P. (S.) saueri sp. nov. (India), P. (S.) annamensis sp. nov. (Vietnam), P. (S.) phoupanensis sp. nov. (Laos, Vietnam), P. (S.) bilyi sp. nov. (Thailand), and P. (S.) haucki sp. nov. (Thailand). We also bring new data on P. (S.) corporaali Reichensperger, 1927 (Java) and P. (S.) madurensis Wasmann, 1913 (India). Besides formal descriptions, we provide photographs of the habitus in the dorsal and dorsolateral view, antennal club, head crest, and male genitalia if the male is available. Based on the comparison of new and earlier described species, we show that the antennae are highly diverse within the Scaphipaussus. Considering other characters, some species are placed in Scaphipaussus, but they differ from putative relatives in the antennal morphology. The presence of the frontal protuberances and crests is a more reliable character. Additional species show that Scaphipaussus is most diverse in southeastern Asia, especially in Indo-Burma. Concerning its supposed late Miocene origin, the group underwent rapid radiation. The species diversity of Scaphipaussus almost doubled in the last decade, and it is highly probable that further species will be described in the future.

Eocene aposematic patterns persist in modern European Lycidae beetles despite the absence of co-mimics.

Ancient aposematic signals might have evolved under different ecological circumstances. Using European Cenozoic amber and phylogenetic reconstruction, we evaluated the evolution of net-winged beetle aposematism. We describe Priabonian Hiekeolycus winkleri sp. nov. from Baltic amber, review known fossil species, and suggest earlier high diversity and morphological conservativeness of European Lycidae since the Eocene. We hypothesize the presence of red and black/red aposematic patterns in Eocene Europe. The analyses suggest the Oligocene to Miocene dispersal of additional species from East Asia and their advergence to autochthonous patterns. Recently dispersed lycids have retained similarities with their East Asian relatives. Net-winged beetles are rare in Europe after the Quaternary climatic oscillations, and we hypothesize a currently relaxed selection for shared aposematic signals. Neophobia, and eventually inborn rejection of brightly colored prey, putatively preserved ancient aposematism under changing conditions. Evidence from paleontology and phylogenetics can provide insight into the long-term persistence of old adaptations under changing conditions.

Enigmatic Campyloxenus: Shedding light on the delayed origin of bioluminescence in ancient Gondwanan click beetles.

Gondwanan elaterids, previously thought to be unrelated, include bioluminescent Campyloxenus earlier placed in bioluminescent Pyrophorinae. Genomic data suggest close relationships between Gondwanan groups. We maintain Morostomatinae and Hapatesinae and redefine Pityobiinae with Nearctic Pityobiini, Gondwanan Parablacini stat. nov., Campyloxenini stat. nov., and Tibionemini trib. nov. Their ancestors putatively underwent differentiation in Gondwana during the Cretaceous separation of southern continents. In contrast with their age, extant groups are species poor. Campyloxenus represents a recent origin of bioluminescence, no older than ∼53 my. Its large pronotal lanterns differ from Pyrophorini and resemble color patches of sympatric beetle co-mimics. This discovery highlights the fourth or fifth origin of bioluminescence in Elateroidea, alongside the lampyroid clade, click beetles Pyrophorini, Alampoides and Coctilelater in Anaissini (Pyrophorinae), and Balgus schnusei (Thylacosterninae). While our phylogenetic findings illuminate the phylogenetic aspects, the complete story awaits further field observations and in-depth genomic analyses of biochemical pathways used by bioluminescent elateroids.

New genus of erotine net-winged beetles, Damzenium gen. nov. (Coleoptera: Lycidae), from Eocene Rovno amber.

A new genus and a new species of erotine net-winged beetles, Damzenium rivnense gen., sp. nov., are described from late Eocene Rovno amber of Ukraine. The new taxon is placed in the tribe Erotini (Lycidae: Erotinae) based on the general appearance, structure of pronotal carinae and elytral costae. Its elytral primary costa 3 is very stout and forms the humeral edge. Primary costa 4 is weak and we have not been able to clearly identify regular four rows of cells in the wide external interstice. Similar modification of elytral longitudinal costae is known from the related tribe Dictyopterini. This discovery increases the number of Baltic and Rovno amber taxa of Lycidae to six genera and seven species. Remarkably, five genera and six species of these belong in the subfamily Erotinae, which has never been reported from other amber deposits. This quite species-poor group of net-winged beetles is still represented almost exclusively in the Holarctic fauna.

Analysis of the Holarctic Dictyoptera aurora Complex (Coleoptera, Lycidae) Reveals Hidden Diversity and Geographic Structure in Müllerian Mimicry Ring.

The elateroid family Lycidae is known for limited dispersal propensity and high species-level endemism. The red net-winged beetle, Dictyoptera aurora (Herbst, 1874), differs from all relatives by the range comprising almost the entire Holarctic region. Based on a five-marker phylogeny and 67 barcode entries (cox1-5' mtDNA) from the whole range, we recovered two genetically distinct species within traditionally defined D. aurora and resurrected the name D. coccinata (Say, 1835) as the oldest available synonym for Nearctic populations. Yet, no reliable morphological trait distinguishes these species except for minute differences in the male genitalia. D. coccinata is a monophylum resulting from a single Miocene dispersal event, ~15.8 million years ago, and genetic divergence implies long-term isolation by the Bering Strait. Far East Asian and west European populations are also genetically distinct, although to a lower extent. Two independent colonization events established the Fennoscandian populations after the last glacial maximum. Besides intrinsic factors, the high morphological similarity might result from stabilizing selection for shared aposematic signals. The rapidly accumulating barcode data provide valuable information on the evolutionary history and the origins of regional faunas.

Phylogenomic and mitogenomic data can accelerate inventorying of tropical beetles during the current biodiversity crisis.

Conservation efforts must be evidence-based, so rapid and economically feasible methods should be used to quantify diversity and distribution patterns. We have attempted to overcome current impediments to the gathering of biodiversity data by using integrative phylogenomic and three mtDNA fragment analyses. As a model, we sequenced the Metriorrhynchini beetle fauna, sampled from ~700 localities in three continents. The species-rich dataset included ~6500 terminals, ~ 1850 putative species delimited at 5% uncorrected pairwise threshold, possibly ~1000 of them unknown to science. Neither type of data could alone answer our questions on biodiversity and phylogeny. The phylogenomic backbone enabled the integrative delimitation of robustly defined natural genus-group units that will inform future research. Using constrained mtDNA analysis, we identified the spatial structure of species diversity, very high species-level endemism, and a biodiversity hotspot in New Guinea. We suggest that focused field research and subsequent laboratory and bioinformatic workflow steps would substantially accelerate the inventorying of any hyperdiverse tropical group with several thousand species. The outcome would be a scaffold for the incorporation of further data from environmental sequencing and ecological studies. The database of sequences could set a benchmark for the spatiotemporal evaluation of biodiversity, would support evidence-based conservation planning, and would provide a robust framework for systematic, biogeographic, and evolutionary studies.

Conspicuousness, phylogenetic structure, and origins of Müllerian mimicry in 4000 lycid beetles from all zoogeographic regions.

Biologists have reported on the chemical defences and the phenetic similarity of net-winged beetles (Coleoptera: Lycidae) and their co-mimics. Nevertheless, our knowledge has remained fragmental, and the evolution of mimetic patterns has not been studied in the phylogenetic context. We illustrate the general appearance of ~ 600 lycid species and ~ 200 co-mimics and their distribution. Further, we assemble the phylogeny using the transcriptomic backbone and ~ 570 species. Using phylogenetic information, we closely scrutinise the relationships among aposematically coloured species, the worldwide diversity, and the distribution of aposematic patterns. The emitted visual signals differ in conspicuousness. The uniform coloured dorsum is ancestral and was followed by the evolution of bicoloured forms. The mottled patterns, i.e. fasciate, striate, punctate, and reticulate, originated later in the course of evolution. The highest number of sympatrically occurring patterns was recovered in New Guinea and the Andean mountain ecosystems (the areas of the highest abundance), and in continental South East Asia (an area of moderate abundance but high in phylogenetic diversity). Consequently, a large number of co-existing aposematic patterns in a single region and/or locality is the rule, in contrast with the theoretical prediction, and predators do not face a simple model-like choice but cope with complex mimetic communities. Lycids display an ancestral aposematic signal even though they sympatrically occur with differently coloured unprofitable relatives. We show that the highly conspicuous patterns evolve within communities predominantly formed by less conspicuous Müllerian mimics and, and often only a single species displays a novel pattern. Our work is a forerunner to the detailed research into the aposematic signalling of net-winged beetles.

Biodiversity Inventory and Distribution of Metriorrhynchina Net-Winged Beetles (Coleoptera: Lycidae), with the Identification of Generic Ranges.

We reviewed the species-level classification of Metriorrhynchina net-winged beetles to make the group accessible for further studies. Altogether, 876 valid species are listed in a checklist along with known synonyms, combinations, and distribution data. The compilation of geographic distribution showed that Metriorrhynchina is distributed mainly in the Australian region with very high diversity in the islands at the northern edge of the Australian craton, i.e., in the Moluccas and New Guinea (54 and 423 spp. respectively). The neighboring northern part of the Australian continent houses a majority of known Australian species (112 spp.) and the diversity of net-winged beetles gradually decreases to the south (43 spp.). The fauna of Sulawesi is highly endemic at the generic level (4 of 10 genera, 67 of 84 spp.). Less Metriorrhynchina occur in the Solomon Islands and Oceania (in total 22 spp.). The Oriental Metriorrhynchina fauna consists of a few genera and a limited number of species, and most of these are known from the Philippines (51 of 94 Oriental spp.). We identified a high species level turn-over between all neighboring landmasses. The genus-level endemism is high in Sulawesi (4 genera) and New Guinea (11 genera), but only a single genus is endemic to Australia. During the compilation of the checklist, we identified some homonyms, and we propose the following replacement names and a new synonym: Metriorrhynchus pseudobasalis, nom. nov. for M. basalis Lea, 1921 nec M. basalis Bourgeois, 1911; Metriorrhynchus pseudofunestus, nom. nov. for M. funestus Lea, 1921 nec M. funestus (Guérin-Méneville, 1838), Trichalus pseudoternatensis, nom. nov. for T. ternatensis Kleine, 1930 nec T. ternatensis Bourgeois, 1900, Procautires subparallelus, nom. nov. for P. parallelus (Pic, 1926) nec P. parallelus (Bourgeois, 1883), and Cautires pseudocorporaali, nom. nov. for C. corporaali (Pic, 1921: 12), (formerly Odontocerus and Cladophorus) nec C. corporaali (Pic, 1921) (formerly Bulenides, later Cautires). Diatrichalus biroi Kleine, 1943, syn. nov. is proposed as a junior subjective synonym of D. subarcuatithorax (Pic, 1926). Altogether, 161 new combinations are proposed, and 47 species earlier placed in Xylobanus Waterhouse, 1879 transferred from Cautirina to Metriorrhynchina incertae sedis. The study clarifies the taxonomy of Metriorrhynchini and should serve as a restarting point for further taxonomic, evolutionary, and biogeographic studies.

Interactions in multi-pattern Müllerian communities support origins of new patterns, false structures, imperfect resemblance and mimetic sexual dimorphism.

Mimicry is a hot spot of evolutionary research, but de novo origins of aposematic patterns, the persistence of multiple patterns in Müllerian communities, and the persistence of imperfect mimics still need to be investigated. Local mimetic assemblages can contain up to a hundred of species, their structure can be a result of multiple dispersal events, and the gradual build-up of the communities. Here, we investigate the structure of lowland and mountain mimetic communities of net-winged beetles by sampling the Crocker Range in north-eastern Borneo and neighbouring regions. The local endemics evolved from the Bornean lowland fauna which is highly endemic at the species level. We inferred that metriorrhynchine net-winged beetles evolved in high elevations yellow/black and reticulate aposematic high-contrast signals from a widespread low-contrast brown/black pattern. As the mountain range is ~ 6 million years old, and these patterns do not occur elsewhere, we assume their in situ origins. We demonstrate that a signal with increased internal contrast can evolve de novo in a mimetic community and can persist despite its low frequency. Additionally, a similar aposematic signal evolves from different structures and its similarity is imperfect. The community with multiple patterns sets conditions for the evolution of aposematic sexual dimorphism as demonstrated by the yellow/black male and reticulate female pattern of Micronychus pardus. These insights elucidate the complex character of the evolution of mimetic signalling in the dynamically diversifying biota of high tropical mountains.

Click Beetle Mitogenomics with the Definition of a New Subfamily Hapatesinae from Australasia (Coleoptera: Elateridae).

Elateridae is a taxon with very unstable classification and a number of conflicting phylogenetic 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 and Pityobiinae. Therefore, Hapatesinae, a new subfamily, is proposed for the Australian genera Hapatesus (21 spp.) and Toorongus Neboiss, 1957 (4 spp.). Parablacinae, Pityobiinae, and Hapatesinae have a putative Gondwanan origin as the constituent genera are known from the Australian region (9 genera) and Neotropical region (Tibionema Solier, 1851), and only Pityobius LeConte, 1853, occurs in the Nearctic region. Another putative Gondwanan lineage, the Afrotropical Morostomatinae, forms either a serial paraphylum with the clade of Parablacinae, Pityobiinae, and Hapatesinae or 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 herein transfer the group to Elaterinae as Eudicronychini, a new status. The mitochondrial genomes provide a sufficient signal for the placement of most lineages, but the deep bipartitions need to be compared with phylogenomic analyses.