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.
Net‐winged beetles (Coleoptera: Lycidae) are a diverse group of elateroids known for aposematism and neoteny. Phylogenetic analyses of morphological and molecular data have revealed different results ...with respect to within‐group relationships. In this study, we recovered a highly supported phylogenomic phylogeny and identified seven subfamilies: Dexorinae stat.n., Calochrominae stat.n., Erotinae, Ateliinae, Lycinae, Lyropaeinae stat.n. and Metriorrhynchinae stat.n. Our results suggest that female neoteny evolved multiple times. Therefore, the development of similar morphological modifications in neotenics may be linked and may have produced characteristics such as body miniaturization, structural simplification, i.e. reduction of mouthparts, fewer antennomeres and palpomeres, uniquely shaped terminal palpomeres, shortened elytra, the loss of coadaptation between the elytra and pronotum, and others. Additional traits evolved in parallel due to similarities in biology, function and sexual selection. These characteristics include mimetic similarities, the presence of the rostrum, pronotal carinae and elytral costae, and the structure of male genitalia. By comparing the phylogenomic topology with the evolution of morphological characters, we were able to identify evolutionary trends in lycids and compare them with analogues for other neotenic elateroids. These traits have not been accepted as homoplasies due to the ambiguous phylogenetic signal from Sanger sequencing markers.
This study presents the first densely sampled phylogenomic analysis of net‐winged beetles (Coleoptera: Lycidae) and compares results with morphology‐based hypotheses.
Multiple origins of female neoteny were recovered and the parallel morphological evolution in males is discussed in detail.
Seven major clades – Dexorinae, Erotinae, Calochrominae, Ateliinae, Lyropaeinae, Lycinae and Metriorrhynchinae – are given subfamily rank in the new classification based on the genomic analysis.
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.
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.
Plastoceridae Crowson, 1972, Drilidae Blanchard, 1845 and Omalisidae Lacordaire, 1857 (Elateroidea) are families of the Coleoptera with obscure phylogenetic relationships and modified morphology ...showing neotenic traits such as soft bodies, reduced wing cases and larviform females. We shotgun sequenced genomes of Plastocerus, Drilus and Omalisus and incorporated them into data matrices of 66 and 4202 single-copy nuclear genes representing Elateroidea. Phylogenetic analyses indicate their terminal positions within the broadly defined well-sclerotized and fully metamorphosed Elateridae and thus Omalisidae should now be considered as Omalisinae stat. nov. in Elateridae Leach, 1815. The results support multiple independent origins of incomplete metamorphosis in Elateridae and indicate the parallel evolution of morphological and ecological traits. Unlike other neotenic elateroids derived from the supposedly pre-adapted aposematically coloured and unpalatable soft-bodied elateroids, such as fireflies (Lampyridae) and net-winged beetles (Lycidae), omalisids and drilids evolved from well-sclerotized click beetles. These findings suggest sudden morphological shifts through incomplete metamorphosis, with important implications for macroevolution, including reduced speciation rate and high extinction risk in unstable habitats. Precise phylogenetic placement is necessary for studies of the molecular mechanisms of ontogenetic shifts leading to profoundly changed morphology.
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.
We analyzed the phylogeny and aposematic patterns of closely related
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
, and categorize their low-contrast aposematic patterns. The warning signal of
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
consisted of 7-85 metriorrhynchine species assigned to 3-10 colour patterns.
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.
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
Candèze, 1863, is a sister of Parablacinae and Pityobiinae. Therefore, Hapatesinae, a new subfamily, is proposed for the Australian genera
(21 spp.) and
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 (
Solier, 1851), and only
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.
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.
The geologically‐complex Indo–Australian–Melanesian archipelago (IAMA) hosts extraordinarily high levels of species richness and endemism and has long served as a natural laboratory for studying ...biogeography and evolution. Nonetheless, its geological history and the provenance and evolution of its biodiversity remain poorly understood. Here, we provide a geological scenario for the IAMA informed by a time‐calibrated molecular phylogeny of 1006 species of Trigonopterus weevils – an exceptionally diverse radiation of regionally‐endemic flightless beetles. Moreover, we performed a statistical biogeographic analysis and examined timing and patterns in the accumulation of lineages residing in a priori‐defined geographic units comprising the IAMA. We estimate that Trigonopterus originated in Australia during the early Paleogene. Subsequent rapid diversification in the area of the present‐day Papuan Peninsula suggests the presence of proto‐Papuan islands by the middle Eocene; the New Guinea North Coast Ranges were colonized in the late Eocene, followed by the New Guinea Highlands and the Bird's Head Peninsula. We inferred the presence of terrestrial habitat in the North Moluccas and Sulawesi in the late Oligocene and the subsequent rapid colonization of Sundaland and the Lesser Sunda Islands. New Caledonia and Samoa were colonized from the Papuan Peninsula, and their faunas also diverged in the late Oligocene. These biota‐informed time estimates are compatible with geological data from the region and shed new light on IAMA paleogeography, even where geological evidence has been lost to erosion. Beetle evolution thus appears to have closely tracked the geological evolution of the IAMA, revealing a uniquely well‐resolved view of regional biogeography.
Abstract
The Lycini (Elateroidea: Lycidae) contains > 400 species placed in four typologically based genera and numerous subgenera. We assembled a mito-ribosomal dataset representing ~100 species ...from the whole range and recovered a phylogeny rejecting Lycus and Lycostomus as polyphyletic assemblages. The male-specific wide elytra and elytral thorns are identified in unrelated Neolycus and Lycus. The morphological similarity based on sexual dimorphism and aposematic patterns defined terminal clades and misled the genus-rank classification. We delimit Neolycus, Rhyncheros reinst. name (= Thoracocalon syn. nov. = Lyconotus syn. nov.), LipernesLycostomus, Haplolycus and Lycus. Demosis and six subgenera of Lycus are synonymized with Lycus. Celiasis Laporte, 1840 is kept in the classification as a nomen dubium until any specimen is available. The deep lineages are known from the Americas and Asia. Africa was colonized by Lycus and Haplolycus. Each specific aposematic pattern occurs in a limited range, and the similar body shape and coloration evolved in unrelated sympatrically occurring lineages. High intraspecific polymorphism is putatively a result of the adaptation of various populations to local mimetic assemblages. Therefore, the delimitation of many phenotypically diverse species should be investigated.