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Sinopyrophorus schimmeli Bi & Li, gen. et sp. nov. Male 17 sternite VIII 18 tergite VIII 19 tergites IX-X with sternite IX 20 aedeagus. Female 21 sternite VIII 22 ovipositor (dorsal view) 23 internal genital tract. Abbreviations: eco, the entry of the common ovdiduct; sgd, spermathecal gland duct. a, dorsal view; b, ventral view; c, lateral view. Scale bars: 1 mm; not to scale (23).
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The new subfamily Sinopyrophorinae within Elateridae is proposed to accommodate a bioluminescent species, Sinopyrophorus schimmeli Bi & Li, gen. et sp. nov., recently discovered in Yunnan, China. This lineage is morphologically distinguished from other click-beetle subfamilies by the strongly protruding frontoclypeal region, which is longitudinally...
Citations
... Within Coleoptera, bioluminescence can be found almost exclusively within the so-called "elaterid-lampyroid clade", including Elateridae, Lampyridae, Phengodidae, Rhagophthalmidae, and Sinopyrophoridae, and probably the extinct Cretophengodidae (Oba et al. 2011;Fallon et al. 2018;Bi et al. 2019;Li et al. 2021b;Kusy et al. 2021;Powell et al. 2022). In Phengodidae, all known larvae and females are bioluminescent, as are males of some species (Costa and Zaragoza-Caballero 2010). ...
Rhagophthalmidae are a small beetle family known from the eastern Palaearctic and Oriental realms. Rhagophthalmidae are closely related to railroad worms (Phengodidae) and fireflies (Lampyridae) with which they share highly modified paedomorphic females and the ability to emit light. Currently, Rhagophthalmidae include 66 species classified in the following 12 genera: Bicladodrilus Pic, 1921 (two spp.), Bicladum Pic, 1921 (two spp.), Dioptoma Pascoe, 1860 (two spp.), Diplocladon Gorham, 1883 (two spp.), Dodecatoma Westwood, 1849 (eight spp.), Falsophrixothrix Pic, 1937 (six spp.), Haplocladon Gorham, 1883 (two spp.), Menghuoius Kawashima, 2000 (three spp.), Mimoochotyra Pic, 1937 (one sp.), Monodrilus Pic, 1921 (two spp. in two subgenera), Pseudothilmanus Pic, 1918 (two spp.), and Rhagophthalmus Motschulsky, 1854 (34 spp.). The replacement name Haplocladon gorhami Kundrata, nom. nov. is proposed for Diplocladon hasseltii Gorham, 1883b (described in subgenus Haplocladon) which is preoccupied by Diplocladon hasseltii Gorham, 1883a. The genus Reductodrilus Pic, 1943 is tentatively placed in Lampyridae: Ototretinae. Lectotypes are designated for Pseudothilmanus alatus Pic, 1918 and P. marginalis Pic, 1918. Interestingly, in the eastern part of their distribution, Rhagophthalmidae have remained within the boundaries of the Sunda Shelf and the Philippines demarcated by the Wallace Line, which separates the Oriental and Australasian realms. This study is intended to be a first step towards a comprehensive revision of the group on both genus and species levels. Additionally, critical problems and prospects for rhagophthalmid research are briefly discussed.
... The typical representatives have a compact body and a pro-mesothoracic clicking mechanism (Costa et al., 2010); however, this group also includes several soft-bodied lineages (Kundrata and Bocak, 2019). Despite recent progress in understanding the composition, phylogeny, and classification of Elateridae, all these aspects remain open to further study (Kundrata et al., 2018a;Bi et al., 2019;Kundrata et al., 2019b;Kusy et al., 2021;Douglas et al., 2021). The most recent study on the phylogeny of Elateridae (Douglas et al., 2021) showed that the Lampyridae and related bioluminescent families may be in fact derived click beetles, which means that the widely delimited Elateridae clade comprises more than 13 500 extant species worldwide (Costa et al., 2010;Douglas et al., 2021). ...
Although the Mesozoic Era played an important role in the
evolution and diversification of Elateridae, the Cretaceous click-beetle
fauna remains very poorly known. Here we describe Cretopachyderes burmitinus gen. et sp. nov. based on
a single specimen from the mid-Cretaceous Burmese amber. This species is
remarkable for its extremely long posterior angles of pronotum, which is a
unique character among fossil Elateridae. We discuss the diagnostic
characters of Cretopachyderes gen. nov. and tentatively place it to subfamily Agrypninae
close to extant genus Pachyderes Guérin-Méneville, 1829.
... The elaterid subfamily Sinopyrophorinae, based on Sinopyrophorus schimmeli Bi & Li in Bi et al. (2019) from Yunnan, China, represents the first bioluminescent click beetle from Asia. In a cladogram based on 14 mitochondrial genes and a Maximum Likelihood analysis, the group formed part of a clade including Hemiopinae and Oestodinae and far removed from other bioluminescent elaterids (Balgus within Lissominae and a terminal agrypnine clade). ...
The hind wings of all known families and most subfamilies of Coleoptera are illustrated, annotated and discussed utilising the terminology of Kukalová-Peck and Lawrence (2004), with a few changes in nomenclature suggested by the senior author. The beetle families are discussed in 21 groups, based on recent classifications of Coleoptera. For each of these groups, the most recent works on phylogeny and classification are reviewed, and the wing characters are discussed to determine if some of the wing features might support or refute relationships based on recent molecular and morphological analyses. Part 1 includes a general discussion of wing structure divided into the following sections: hind wing fields, veinal systems (including the history of wing nomenclature), wing folding, wing edge and embayments, hinges and bending zones, cross-veins and braces, cells and other landmarks. It is followed by discussion of the first 14 groups (Archostemata to Elateroidea), 15 figures supporting general discussions, and 426 labelled wing images of the discussed groups, representing 380 genera.
... Efforts to resolve the subfamily level phylogeny of click-beetles using morphology [8,9] have had little success. Phylogenies using few genes have recovered some subfamilies as classically defined and have provided new insights [10][11][12][13][14][15]. However, these studies often lacked resolution or statistical support for subfamily level relationships, and morphological studies omitted exemplars of elateroid families now known to belong within Elateridae. ...
... Most DNA studies found some or all lampyroids as sister to Elateridae [6 in part,12,14 in part,16], or part of Elateridae [2,3,6 in part]. Both morphological and molecular studies agree on the monophyly of Cardiophorinae and Cardiophorinae + Negastriinae [9,12,15,18] and on the monophyly of Agrypninae [9][10][11][12][13]15,19], including former Drilidae when studied. The largest subfamily, Elaterinae, are monophyletic in all DNA-based studies [6,10,13,14,20] with incorporation of Eudicronychinae, where tested. ...
... Most DNA studies found some or all lampyroids as sister to Elateridae [6 in part,12,14 in part,16], or part of Elateridae [2,3,6 in part]. Both morphological and molecular studies agree on the monophyly of Cardiophorinae and Cardiophorinae + Negastriinae [9,12,15,18] and on the monophyly of Agrypninae [9][10][11][12][13]15,19], including former Drilidae when studied. The largest subfamily, Elaterinae, are monophyletic in all DNA-based studies [6,10,13,14,20] with incorporation of Eudicronychinae, where tested. ...
Click-beetles (Coleoptera: Elateridae) are an abundant, diverse, and economically important beetle family that includes bioluminescent species. To date, molecular phylogenies have sampled relatively few taxa and genes, incompletely resolving subfamily level relationships. We present a novel probe set for anchored hybrid enrichment of 2260 single-copy orthologous genes in Elateroidea. Using these probes, we undertook the largest phylogenomic study of Elateroidea to date (99 Elateroidea, including 86 Elateridae, plus 5 non-elateroid outgroups). We sequenced specimens from 88 taxa to test the monophyly of families, subfamilies and tribes. Maximum likelihood and coalescent phylogenetic analyses produced well-resolved topologies. Notably, the included non-elaterid bioluminescent families (Lampyridae + Phengodidae + Rhagophthalmidae) form a clade within the otherwise monophyletic Elateridae, and Sinopyrophoridae may not warrant recognition as a family. All analyses recovered the elaterid subfamilies Elaterinae, Agrypninae, Cardiophorinae, Negastriinae, Pityobiinae, and Tetralobinae as monophyletic. Our results were conflicting on whether the hypnoidines are sister to Dendrometrinae or Cardiophorinae + Negastriinae. Moreover, we show that fossils with the eucnemid-type frons and elongate cylindrical shape may belong to Eucnemidae, Elateridae: Thylacosterninae, ancestral hard-bodied cantharoids or related extinct groups. Proposed taxonomic changes include recognition of Plastocerini as a tribe in Dendrometrinae and Hypnoidinae stat. nov. as a subfamily within Elateridae.
... The click-beetles (Elateridae) are the major family in Elateroidea, comprising more than 10,000 described species worldwide [1]. Despite the efforts of numerous studies using morphological or molecular data, the classification and phylogenetic relationships within the family remain far from fully understood [2][3][4][5][6][7][8][9][10][11]. Taking this into consideration, further development of click-beetle systematics and understanding their evolution would certainly benefit from integrating modern molecular-based methods and morphology • Genus Agrypnus Eschscholtz, 1829 * Agrypnus Eschscholtz, 1829: 32 [117]. ...
The Elateridae (click-beetles) are the largest family in Elateroidea; however, their relationships, systematics and classification remain unclear. Our understanding of the origin, evolution, palaeodiversity and palaeobiogeography of Elateridae, as well as reconstruction of a reliable time-calibrated phylogeny for the group, are hampered by the lack of detailed knowledge of their fossil record. In this study, we summarize the current knowledge on all described fossil species in Elateridae, including their type material, geographic origin, age, bibliography and remarks on their systematic placement. Altogether, 261 fossil species classified in 99 genera and nine subfamilies are currently listed in this family. The Mesozoic click-beetle diversity includes 143 species, with most of them described from the Jurassic Karatau, and 118 described species are known from the Cenozoic deposits, mainly from the Eocene North American Florissant Formation and European Baltic amber. Available data on the described past diversity of Elateridae suggest that almost all fossil lineages in this group are in urgent need of revision and numerous Mesozoic species might belong to different families. Our study is intended to serve as a comprehensive basis for all subsequent research focused on the click-beetle fossil record.
... Its typical representatives are well-known for their hard, compact body and the pro-mesothoracic clicking mechanism 41,42 ; however, there are several soft-bodied lineages currently included in the family which were historically considered separate families [43][44][45] . The composition, phylogeny and natural classification of Elateridae are far from fully understood, despite the effort of numerous recent studies 43,44,46,47 . Elateridae comprise more than 10,000 extant species worldwide 42 , and the fossil record includes approximately 300 species, although the placement of many of them needs thorough investigation 48,49 . ...
... Phylogenetic analyses based chiefly on morphology have shown that their results must be treated with caution 70 . Recent molecular analyses at least partly improved our understanding the phylogeny and classification of the present-day higher taxa 43,44,46,47 ; however, this is not possible for fossils, for which we have only morphological data available. Therefore, for a systematic classification of the newly Scientific Reports | (2020) 10:20158 | https://doi.org/10.1038/s41598-020-76908-3 ...
Beetle fossils are a rich source of information about the palaeodiversity and evolutionary history of the order Coleoptera. Despite the increasing rate of fossil research on click-beetles (Coleoptera: Elateridae), the most diverse group in the superfamily Elateroidea, their fossil record has remained largely unstudied. This may be caused by the combination of their rather uniform external morphology and the suboptimal state of preservation and visibility in most fossil specimens. Here, we used X-ray micro-computed tomography to reconstruct the morphology of an interesting click-beetle from Eocene Baltic amber, which had some principal diagnostic characters obscured by opaque bubbles and body position. Our results suggest that the newly described Baltelater bipectinatus gen. et sp. nov. belongs to tribe Protelaterini within subfamily Lissominae. Since Protelaterini have a predominantly Gondwanan distribution, our discovery is of a great importance for the historical biogeography of the group. Very distinctive are the bipectinate antennae with 11 antennomeres and with rami beginning on antennomere IV, which are not found in any recent Elateridae. The discovery of a new click-beetle lineage from European Eocene amber sheds further light on the palaeodiversity and historical diversification of the family as well as on the composition of the extinct amber forest ecosystem.
... Considering that there are still no reference genomes available for other non-luminous families in Elateroidea, we constructed an additional mitogenomic phylogeny for 11 Elateroidea families (including luminous and non-luminous families) (Supplementary Table S14) to explore the phylogenetic distribution of bioluminescent within Elateroidea taxa. Our results (Fig. 2b) indicate that Lampyridae, together with other luminous families (Asian Rhagophthalmidae and South American Phengodidae), is a sister clade to world-wide Elateridae, a family with only some luminous species mainly in South America but recently also found by us in Asia 17 . They corroborate with the phylogenies inferred from 95 nuclear protein-coding genes of beetles 18 and from 13 protein-coding genes of mitogenomes and two nuclear ribosomal DNA (rDNA) (18S, 28S) 19 , and with the beetle tree 16 but differ from the phylogenies inferred from mitochondrial genes (16S, COI) and two nuclear rDNA (18S, 28S) 15 . ...
Fireflies are among the most charismatic insects for their spectacular bioluminescence, but the origin and evolution of bioluminescence remain elusive. Especially, the genic basis of luciferin (D-luciferin) biosynthesis and light patterns is largely unknown. Here, we present the high-quality reference genomes of two fireflies Lamprigera yunnana (1053 Mb) and Abscondita terminalis (501 Mb) with great differences in both morphology and luminous behavior. We sequenced the transcriptomes and proteomes of luminous organs of two species. We created the CRISPR/Cas9-induced mutants of Abdominal B gene without luminous organs in the larvae of A. terminalis and sequenced the transcriptomes of mutants and wild-types. Combining gene expression analyses with comparative genomics, we propose a more complete luciferin synthesis pathway, and confirm the convergent evolution of bioluminescence in insects. Using experiments, the function of the firefly acyl-CoA thioesterase (ACOT1) to convert L-luciferin to D-luciferin was validated for the first time. Comparisons of three-dimension reconstruction of luminous organs and their differentially expressed genes among two species suggest that two positive genes in the calcium signaling pathway and structural difference of luminous organs may play an important role in the evolution of flash pattern. Altogether, our results provide important resources for further exploring bioluminescence in insects.
... Most luminous beetles belong to Elateroidea and we know of ∼2000 firefly species (Lampyridae, Fig. 1F-J), ∼300 glow-worms or railroad-worm beetles (Phengodidae, Rhagophthalmidae), as well as >100 bioluminescent click beetle species (Elateridae; Fig. 1A-D), especially in the Neotropical region (Costa, 1975(Costa, , 1984. With the recent discovery of the first Palearctic bioluminescent clicking beetle, the number of bioluminescent beetle lineages has increased Bi et al., 2019;Fig. 1D-E). ...
... Recently, the subfamily Sinopyrophorinae Bi & Li was proposed in Elateridae for the bioluminescent Sinopyrophorus schimmeli Bi & Li. Bi et al. (2019) included homologous fragments of S. schimmeli in a phylogenetic reconstruction from earlier published click beetle rRNA and mtDNA genes (Bocakova et al., 2007;Timmermans et al., 2010Timmermans et al., , 2016Kundrata et al., 2014;Amaral et al., 2016) and used glow-worms as outgroups. They recovered Sinopyrophorus with the elaterid subfam ...
... The formal requirement for the monophyly of all named taxa would also be fulfilled by the redefinition of Elateridae sensu lato that would contain eight traditional families Elateridae, Drilidae, Omalisidae, Plastoceridae, Lampyridae, Phengodidae, Rhagophthalmidae and Sinopyrophorinae. (Kusy et al., 2018a,b;Bi et al., 2019). Considering also the logical extreme, we could accept whole Elateroidea as a single family (sans the artemotopodid clade) as recently mentioned by Muona & Taräväinen (2020). ...
Bioluminescence has been hypothesized as aposematic signalling, inter-sexual communication and a predatory strategy, but origins and relationships among bioluminescent beetles have been contentious. We reconstruct the phylogeny of the bioluminescent elateroid beetles (i.e. Elateridae, Lampyridae, Phengodidae and Rhagophthalmidae), analysing genomic data of Sinopyrophorus Bi & Li, and in light of our phylogenetic results, we erect Sinopyrophoridae Bi & Li, stat.n. as a clicking elaterid-like sister group of the soft-bodied bioluminescent elateroid beetles, that is, Lampyridae, Phengodidae and Rhagophthalmidae. We suggest a single origin of bio-luminescence for these four families, designated as the 'lampyroid clade', and examine the origins of bioluminescence in the terminal lineages of click beetles (Elateridae). The soft-bodied bioluminescent lineages originated from the fully sclerotized elateroids as a derived clade with clicking Sinopyrophorus and Elateridae as their serial sister groups. This relationship indicates that the bioluminescent soft-bodied elateroids are modified click beetles. We assume that bioluminescence was not present in the most recent common ancestor of Elateridae and the lampyroid clade and it evolved among this group with some delay, at the latest in the mid-Cretaceous period, presumably in eastern Laurasia. The delimitation and internal structure of the elaterid-lampyroid clade provides a phylogenetic framework for further studies on the genomic variation underlying the evolution of bioluminescence.
... Several other authors also investigated the chemical process of bioluminescence in beetles [13,50] and in marine snails (limpets) and luminous bacteria [29,43]. In recent times, several works can be found with reference to fungal, bacterial, marine bioluminescence and bioluminescence in insects [6,12,15,24]. Kaskova et al. have recently reported the structure of fungal oxyluciferin and its mechanism [24]. In another interesting work, Fallon et al. have found a strong support for independent origin of luciferase (and so bioluminescence) between fireflies and click beetle, which provides new insights into the evolutionary path of these insects' bioluminescence and their genes, chemical defenses, and symbionts [15]. ...
... The minimal dynamical model presented by Eqs. (6,8) represents the basic lighting mechanism of a firefly flash, considering one lighting cycle of a firefly. However, we note that the model lacks a basic feedback mechanism, which must be introduced before we can proceed. ...
A low dimensional nonlinear model based on the basic lighting mechanism of a firefly is proposed (Saikia and Bora in Nonlinear model of the firefly flash. http://export.arxiv.org/pdf/2002.01183). The basic assumption is that the firefly lighting cycle can be thought to be a nonlinear oscillator with a robust periodic cycle. We base our hypothesis on the well-known light producing reactions involving enzymes, common to many insect species, including the fireflies. We compare our numerical findings with the available experimental results which correctly predicts the reaction rates of the underlying chemical reactions. Toward the end, a time-delay effect is introduced for possible explanation of appearance of multiple-peak light pulses, especially when the ambient temperature becomes low.
... With the rise of molecular phylogenetic analyses, several soft-bodied groups, which were previously considered separate families, i.e., Drilidae, Omalisidae, and Plastoceridae, were merged with Elateridae [7,8]. However, the internal phylogenetic relationships, as well as the suprageneric classification of the group, remain open to further study [9][10][11]. ...
... Elateridae are among the most common beetle families in the fossil record [16,54,128]. However, their real palaeodiversity remains understudied [16], most probably because of their rather uniform and generally problematic external morphology which often causes problems even in identification and classification of recent lineages [1,[3][4][5][6][7][8][9][10][11]. Consequently, the systematics and classification of fossil elaterids has been in a constant state of flux. ...
... The only exception is Alaodima, which is assigned to Dimini, but there is an ongoing debate if this group belongs to Dendrometrinae or forms a separate subfamily [13,72]. The missing record of Elaterinae and Dendrometrinae genera in the early evolution of Elateridae is most probably artificial, because they belong to the basal-most splits based on the results of recent molecular phylogenetic analyses [9][10][11]. This situation may be partly caused by the hard-to-observe morphological characters defining the subfamilies and their tribes [1,4,5,78], which are usually not visible even if the fossils are well-preserved. ...
Insect fossils bear important information about the evolutionary history of the group. The fossil record of Elateridae, a large cosmopolitan beetle family, has been greatly understudied and the available data are often replete with ambiguity and uncertainty. The research of Elateridae evolution cannot be done without solid genus-group name concepts. In this study we provide an updated comprehensive summary of the fossil genera in Elateridae, including their systematic placement and information on the type species, gender, number of species, age range, and relevant bibliography. We list seven valid fossil genera in Agrypninae, one in Cardiophorinae, two in Dendrometrinae, five in Elaterinae, two in Negastriinae, one in Omalisinae, one in Pityobiinae, and 36 in Protagrypninae. Additional 19 genera are tentatively classified as Elateridae incertae sedis, and their placements are discussed. Further, we move genera Babuskaya Martins-Neto & Gallego, 2009, Cardiosyne Martins-Neto & Gallego, 2006, Fengningia Hong, 1984 and Gemelina Martins-Neto & Gallego, 2006 from Elateridae to Coleoptera incertae sedis. We also discuss the genera previously placed in Elateridae, which are currently not included in the family. The data on the fossil generic diversity suggest that Elateridae originated in the Triassic and rapidly diversified and became comparatively abundant through the Jurassic. We call for further research on the fossil Elateridae from various deposits in order to increase our knowledge on the origin, evolution, and palaeodiversity of the group.
