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General habitus of Cretophengodidae and representatives of the closely related Phengodidae and Rhagophthalmidae, under incident light. (a,b) Cretophengodes azari gen. et sp. nov., dorsal and ventral views, respectively, with arrowhead showing the photic organ. (c,d) Zarhipis sp. (Phengodidae), dorsal and ventral views, respectively. (e,f ) Rhagophthalmus sp. (Rhagophthalmidae), dorsal and ventral views, respectively. Scale bars: (a,b,e,f ) 2 mm; (c,d) 4 mm.

General habitus of Cretophengodidae and representatives of the closely related Phengodidae and Rhagophthalmidae, under incident light. (a,b) Cretophengodes azari gen. et sp. nov., dorsal and ventral views, respectively, with arrowhead showing the photic organ. (c,d) Zarhipis sp. (Phengodidae), dorsal and ventral views, respectively. (e,f ) Rhagophthalmus sp. (Rhagophthalmidae), dorsal and ventral views, respectively. Scale bars: (a,b,e,f ) 2 mm; (c,d) 4 mm.

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Bioluminescent beetles of the superfamily Elateroidea (fireflies, fire beetles, glow-worms) are the most speciose group of terrestrial light-producing animals. The evolution of bioluminescence in elateroids is associated with unusual morphological modifications, such as soft-bodiedness and neoteny, but the fragmentary nature of the fossil record di...

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... The number of genera included in Rhagophthalmidae and also their placement within Elateroidea classification vary by source (e.g., McDermott 1966;Crowson 1972;Lawrence and Newton 1995;Kawashima et al. 2010;Kundrata and Bocak 2011a). In the last decade, Elateroidea systematic research has accelerated and the classification of the superfamily has experienced many taxonomic changes (e.g., Kundrata et al. 2014Bocak et al. 2018;Kusy et al. 2018bKusy et al. , 2021, including the discoveries of two new recent families Rosa et al. 2020) and one new extinct family (Li et al. 2021b). However, only six new species of Rhagophthalmidae were described in three taxonomic papers in the same period (Ho et al. 2012;Kazantsev 2012;Yiu 2017). ...
... Kusy et al. (2021) defined the "lampyroid clade", which contains Lampyridae, Phengodidae, Rhagophthalmidae, and Sinopyrophoridae. Fossil Cretophengodidae were probably also a part of that clade (Li et al. 2021b). ...
... Cretophengodidae were described from mid-Cretaceous amber of northern Myanmar (ca. 99 Mya, Shi et al. 2012;Li et al. 2021b), and Kusy et al. (2021) reported unpublished Phengodidae from the same deposit. Kusy et al. (2021) summarized and reviewed the published molecular dating analyses of the elaterid-lampyroid clade, and showed that median estimates suggest the split of the Lampyridae, Phengodidae, and Rhagophthalmidae clade in the mid-Cretaceous. ...
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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.
... We note that the results of our morphological phylogenetic analysis must be interpreted with caution, since most deep relationships within this tree are incongruent with recent phylogenomic data, as discussed above. This likely reflects prevalent homoplasy in the morphological data set, which has been recognized as a considerable problem in elateriform morphological phylogenies (Kundrata et al. 2014, Li et al. 2021. Because the fossils possess a combination of characters from both families, we regard Mastigocoleidae as closely allied with these two, either as sister to the clade Lutrochidae + Dryopidae or as a stem group to Dryopoidea. ...
Article
With some 3,700 described species, Dryopoidea are a moderately diverse superfamily of beetles whose position within basal Polyphaga has been historically difficult to elucidate. Members of most extant dryopoid families are set apart from the majority of other polyphagans by their association with aquatic habitats, but little is known about the origin of these derived life habits and the phylogeny of the superfamily. Here we describe Mastigocoleidae Tihelka, Jäch, Kundrata & Cai fam. nov., a new family of Mesozoic dryopoids represented by fossils from the Cretaceous Yixian Formation in northeastern China (undescribed species; ~125 Ma), Crato Formation in northeastern Brazil (Mastigocoleus rhinoceros Tihelka & Cai gen. et sp. nov.; ~113 Ma), and amber from northern Myanmar (Mastigocoleus resinicola Tihelka & Cai gen. et sp. nov. and Cretaceocoleus saetosus Tihelka, Kundrata & Cai gen. et sp. nov.; ~99 Ma). Integrating the findings of recent molecular and morphological phylogenetic analyses, we recover Mastigocoleidae as an early-diverging dryopoid clade sister to the families Lutrochidae and Dryopidae, or less likely as a group of putative stem-dryopoids. Mastigocoleidae are most distinctly separated from all other dryopoid families by their whip-like antennae, with 11 antennomeres, reaching to the pronotal base, and with the scape broadest and longest, a short pedicel, and antennomeres II–XI more or less distinctively gradually tapering toward the apex. Mastigocoleidae indicate that the last common ancestor of Dryopoidea was likely terrestrial in the adult stage, and document character acquisitions associated with a specialization for aquatic life.
... Fossils of the paedomorphic lineages could provide valuable palaeobiological data to investigate evolution of paedomorphosis and, in particular, to test the higher speciation rates hypotheses. In the Elateroidea fossil record, several taxa affected by the paedomorphic syndrome were found in various families [31][32][33] . In the described fossil record of Lycidae, currently comprised by 12 known species 20,34-37 , only three species were assigned to lineages that are known to have individuals affected by the paedomorphic syndrome, i.e., Electropteron avus Kazantsev, 2013 39 . ...
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Paedomorphosis is a heterochronic syndrome in which adult individuals display features of their immature forms. In beetles, this phenomenon occurs widely in the superfamily Elateroidea, including the net-winged beetles (Lycidae), and, due to the usual flightlessness of paedomorphic females, it is hypothesized to cause speciation rates higher than in non-paedomorphic lineages. However, some fossils of paedomorphic lycids do not support this with palaeobiological data. Discovery of new Lycidae fossils attributed to the West Indian extant paedomorphic genus Cessator Kazantsev in the Dominican amber also suggests morphological stasis within this genus in the Greater Antilles. We describe Cessator anachronicus Ferreira and Ivie, sp. nov. based on adult males, as well as the first ever recorded fossil net-winged beetle larva of the same genus. We propose that the relatively young age of the studied fossils combined with the stable conditions in the forest floor of the Greater Antilles through the last tens of million years could explain the exceptionally conserved morphology in the net-winged beetles affected by the paedomorphic syndrome.
... Our molecular clock estimates corroborate the controversial idea, famously portrayed in numerous palaeontological reconstructions, that coprophagous beetles, namely geotrupids (dung beetles) and scarabaeoids (scarabs), may have been associated with Cretaceous herbivorous dinosaurs [23,116,117]. Our analyses also corroborate a Cretaceous origin of the bioluminescent lampyroid clade [118], temporally overlapping with the diversification of visually hunting predators such as anurans and stem-group birds during the KTR [119]. At the same time, some Mesozoic beetle families have their last appearance in the fossil record during the KTR, highlighting complex dynamics of transitioning from a gymnosperm-to angiosperm-dominated world [120]. ...
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Beetles constitute the most biodiverse animal order with over 380 000 described species and possibly several million more yet unnamed. Recent phylogenomic studies have arrived at considerably incongruent topologies and widely varying estimates of divergence dates for major beetle clades. Here, we use a dataset of 68 single-copy nuclear protein-coding (NPC) genes sampling 129 out of the 193 recognized extant families as well as the first comprehensive set of fully justified fossil calibrations to recover a refined timescale of beetle evolution. Using phylogenetic methods that counter the effects of compositional and rate heterogeneity, we recover a topology congruent with morphological studies, which we use, combined with other recent phylogenomic studies, to propose several formal changes in the classification of Coleoptera: Scirtiformia and Scirtoidea sensu nov., Clambiformia ser. nov. and Clamboidea sensu nov., Rhinorhipiformia ser. nov., Byrrhoidea sensu nov., Dryopoidea stat. res., Nosodendriformia ser. nov. and Staphyliniformia sensu nov., and Erotyloidea stat. nov., Nitiduloidea stat. nov. and Cucujoidea sensu nov., alongside changes below the superfamily level. Our divergence time analyses recovered a late Carboniferous origin of Coleoptera, a late Palaeozoic origin of all modern beetle suborders and a Triassic–Jurassic origin of most extant families, while fundamental divergences within beetle phylogeny did not coincide with the hypothesis of a Cretaceous Terrestrial Revolution.
... Natural bioluminescence is the emission of visible lights from a living organism due to a chemical reaction (Haddock et al. 2010). It is widely found across the ocean and terrestrial environments as well as across diverse groups of organisms, including unicellular algae, bacteria (Tanet et al. 2019), multicellular fungi (Karunarathna et al. 2020;Mishra & Srivastava 2021) and animals (Mensinger 2011;Li et al. 2021). At a deeper level, bioluminescence is currently understood as the oxidization of a light-emitting molecule known as luciferin, which is oxidized by an enzyme called luciferase or photoproteins, resulting in an intermediate high-energy substance after which the intermediate subsequently converts into oxyluciferin (Shimomura 2006;Moline et al. 2013). ...
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Seven species of bioluminescent mushrooms belonging to seven genera have been described in the tropical rainforests of China. This study contributes the eighth species, found growing on decaying bamboo in Xishuangbanna Tropical Botanical Garden (XTBG). Morphological characteristics and phylogenetic analyses using internal transcribed spacer (ITS) and ribosomal large subunit (LSU) gene regions placed the species within the genus Favolaschia. Comprehensive morphological descriptions, micro and macro photographs, and a phylogenetic tree showing the placement of the new species are provided. This is the second report of bioluminescent Favolaschia in China.
... Fossil calibrations are known to have significant impact on divergence time estimation depending on their placement and the age distribution provided (Saladin et al., 2017;Toussaint et al., 2017). Recently described fossils (e.g., Li et al., 2021) and a phylogenetic reconstruction of bioluminescent beetles (Martin et al., 2020) provides us with the opportunity to explore the time of origin for both terrestrial and aerial beetle bioluminescence with greater confidence than ever before. We used an existing large-scale phylogeny to estimate the age for the origin of bioluminescence. ...
... Fossil calibrations were placed on topologies based on morphological similarities with extant taxa discussed within the literature and their currently proposed placements. We placed one fossil at the base of Luciolinae (Protoluciola albertalleni; Kazantsev, 2015) and one at the node Phengodidae + Rhagophthalmidae (Cretophengodes azari; Li et al., 2021). We also placed a fossil at the base of the subfamily Lamprohizinae (Lamprohiza fossilis ;Beier 1952;Fanti, 2017), and a fourth at the base of an extant genus (Photinus), using the fossil Photinus kazantsevi (Alekseev, 2019). ...
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Bioluminescence is found across life and has many functions. Yet we understand very little about its timing and origins, particularly as a predator avoidance strategy. Understanding the timing between bioluminescence and predator origins has yet to be examined and can help elucidate the evolution of the ecologically important signal aposematism. Using the most prevalent bioluminescent group, fireflies, where bioluminescence primarily functions as aposematic and sexual signals, the timing for the origins of both potential predators of fireflies and bioluminescence is explored. Divergence time estimations were performed using a genomic-scale phylogenetic reconstruction Lampyridae, and multiple fossil calibration points, allowing for a robust estimate for the origin of beetle bioluminescence as both a terrestrial and aerial signal. Our results recover the origins of terrestrial beetle bioluminescence at 141 mya and aerial bioluminescence at 133 mya. These ages predate the origins of all known extant aerial predators (i.e., bats and birds) and support the much older terrestrial predators (frogs, ground beetles, lizards, snakes, and hunting spiders) as the most likely drivers of bioluminescence in beetles.
... The classification of soft-bodied elateroids based solely on morphology is precluded by the fact that many related groups are differently affected by neoteny, thus being not similar, but on the other hand, some lineages look superficially very similar although they are only distantly related 1 . The morphology-based phylogenetic analyses failed to provide us with the natural classification of the Elateroidea 13 or to determine the phylogenetic position of a recently described fossil softbodied lineage 11 . Thus, since we cannot reliably use the morphological characters to test the position of Anoeuma gen. ...
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We here report a new elateroid, Anoeuma lawrencei Li, Kundrata and Cai gen. et sp. nov., from mid-Cretaceous Burmese amber. Though superficially similar to some soft-bodied archostematans, Anoeuma could be firmly placed in the polyphagan superfamily Elateroidea based on the hind wing venation. Detailed morphological comparisons between extant elateroids and the Cretaceous fossils suggest that the unique character combination does not fit with confidence into any existing soft-bodied elateroid group, although some characters indicate possible relationships between Anoeuma and Omalisinae. Our discovery of this new lineage further demonstrates the past diversity and morphological disparity of soft-bodied elateroids.
... Beyond organism living in water [235], the numerous and mostly known examples of bioluminescence in terrestrial living beings are represented in the superfamily Elateroidea, from the order of Coleoptera. These include glow-worms fire beetles, fireflies, which have attracted interest to the evolution of their bioluminescence accompanied by changes in morphology and neoteny, stimulating also fossil-based research [250]. ...
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Organisms belonging to all life kingdoms may have the natural capacity to fluoresce. Autofluorescence events depend on the presence of natural biomolecules, namely endogenous fluorophores, with suitable chemical properties in terms of conjugated double bonds, aromatic or more complex structures with oxidized and crosslinked bonds, ensuring an energy status able to permit electronic transitions matching with the energy of light in the UV-visible-near-IR spectral range. Emission of light from biological substrates has been reported since a long time, inspiring unceasing and countless studies. Early notes on autofluorescence of vegetables have been soon followed by attention to animals. Investigations on full living organisms from the wild environment have been driven prevalently by ecological and taxonomical purposes, while studies on cells, tissues and organs have been mainly promoted by diagnostic aims. Interest in autofluorescence is also growing as a sensing biomarker in food production and in more various industrial processes. The associated technological advances have supported investigations ranging from the pure photochemical characterization of specific endogenous fluorophores to their possible functional meanings and biological relevance, making fluorescence a valuable intrinsic biomarker for industrial and diagnostic applications, in a sort of real time, in situ biochemical analysis. This review aims to provide a wide-ranging report on the most investigated natural fluorescing biomolecules, from microorganisms to plants and animals of different taxonomic degrees, with their biological, environmental or biomedical issues relevant for the human health. Hence, some notes in the different sections dealing with different biological subject are also interlaced with human related issues. Light based events in biological subjects have inspired an almost countless literature, making it almost impossible to recall here all associated published works, forcing to apologize for the overlooked reports. This Review is thus proposed as an inspiring source for Readers, addressing them to additional literature for an expanded information on specific topics of more interest.
... Hence, we found that lampyroids are modified elaterid click-beetles, so that the entire clade contains over 13,500 species. While this finding implies that major taxonomic changes are needed to Elateridae or to all these families (plus fossil Cretophengodidae [82]), we argue that such formal incorporation of lampyroids should await further study. We do not recommend changes to the rank of Sinopyrophoridae. ...
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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.
... Insecta luciferases (Group V), a luciferase type also possibly found in the cephalopod Watasenia scintillans All luminous insects (i.e., Coleoptera with around 2300 luminous species (Li et al. 2021); Diptera with a lower specific diversity but not yet exhaustively evaluated to the best of our best knowledge; ...
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Bioluminescence – i.e., the emission of visible light by living organisms - is defined as a biochemical reaction involving, at least, a luciferin substrate, an oxygen derivative, and a specialised luciferase enzyme. In some cases, the enzyme and the substrate are durably associated and form a photoprotein. While this terminology is educatively useful to explain bioluminescence, it gives a false idea that all luminous organisms are using identical or homologous molecular tools to achieve light emission. As usually observed in biology, reality is more complex. To date, 11 different luciferins have indeed been discovered, and several non-homologous luciferases lato sensu have been identified which, all together, confirms that bioluminescence emerged independently multiple times during the evolution of living organisms. While some phylogenetically related organisms may use non-homologous luciferases (e.g., at least four convergent luciferases are found in Pancrustacea), it has also been observed that phylogenetically distant organisms may use homologous luciferases (e.g., parallel evolution observed in some cnidarians, tunicates and echinoderms that are sharing a homologous luciferase-based system). The evolution of luciferases then appears puzzling. The present review takes stock of the diversity of known “bioluminescent proteins”, their evolution and potential evolutionary origins. A total of 134 luciferase and photoprotein sequences have been investigated (from 75 species and 11 phyla), and our analyses identified 12 distinct types – defined as a group of homologous bioluminescent proteins. The literature review indicated that genes coding for luciferases and photoproteins have potentially emerged as new genes or have been co-opted from ancestral non-luciferase/photoprotein genes. In this latter case, the homologous gene’s co-options may occur independently in phylogenetically distant organisms.