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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.
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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...
Citations
... Since the Burma Terrane was likely an isolated island for large periods of time, numerous taxa were probably also island endemics. This hypothesis is confirmed by numerous taxa being known exclusively from Burmese amber, including the insect families from various orders [90][91][92][93][94][95] . ...
The click beetles (Elateridae) represent the major and well-known group of the polyphagan superfamily Elateroidea. Despite a relatively rich fossil record of Mesozoic Elateridae, only a few species are described from the Upper Cretaceous Burmese amber. Although Elateridae spend most of their lives as larvae, our knowledge on immature stages of this family is limited, which is especially valid for the fossils. So far, only a single larval click beetle has been reported from Burmese amber. Here, we describe two larval specimens from the same deposit which based on their morphology unambiguously belong to the predominantly Southern Hemisphere subfamily Pityobiinae, being the most similar to the representatives of tribe Tibionemini. However, since the larvae of the closely related bioluminescent Campyloxenini have not yet been described, we place our specimens to Tibionemini only tentatively. One species of Pityobiinae was recently described from Burmese amber based on adults, and we discuss if it can be congeneric with the here-reported larvae. Recent representatives of the Tibionemini + Campyloxenini clade are known from South America and New Zealand, and this group is hypothesized to have a Gondwanan origin. Hence, the newly discovered Burmese amber larvae may further contribute to a recently highly debated hypothesis that biota of the resin-producing forest on the Burma Terrane, which was probably an island drifting northward at the time of amber deposition, had at least partly Gondwanan affinities. The discovery of enigmatic click beetle larvae in the Upper Cretaceous Burmese amber sheds further light on the palaeodiversity and distribution of the relatively species-poor Gondwanan clade of click beetles, which contain a recent bioluminescent lineage, as well as on the taxonomic composition of the extinct Mesozoic ecosystem.
... The mesmerizing dances of fireflies on warm summer nights have fascinated generations of our ancestors and sparked centuries of scientific enquiry. With approximately 2500 described species [1], the fireflies (Lampyridae) are by far the most widespread and abundant bioluminescent beetles, sharing their ability to produce light with five related elateroid families-some click beetles (Elateridae) from the Neotropical region and small Melanesian islands, Sinopyrophoridae (Sinopyrophorus Bi & Li) endemic to south China, predominantly New World glow-worms (Phengodidae), the Old World star-worms (Rhagophthalmidae) and the recently described fossil family Cretophengodidae [2][3][4][5][6]. Bioluminescence in fireflies is known to fulfil a dual functionin communication [7] and as an aposematic antipredator mechanism [8]. ...
... Although it had no apparent light organ, we can hypothesize that the species was bioluminescent like all known extant phengodid species, at least as immature stages and larviform females. Li et al. [4] described a transitional bioluminescent elateroid beetle family (Cretophengodidae, represented by Cretophengodes azari Li, Kundrata, Tihelka & Cai) from the same amber deposit. It belongs to the bioluminescent lampyroid clade, representing a transitional fossil linking the soft-bodied Phengodidae+Rhagophthalmidae clade and the primitively hard-bodied elateroids. ...
The beetle superfamily Elateroidea comprises the most biodiverse bioluminescent insects among terrestrial light-producing animals. Recent exceptional fossils from the Mesozoic era and phylogenomic studies have provided valuable insights into the origin and evolution of bioluminescence in elateroids. However, due to the fragmentary nature of the fossil record, the early evolution of bioluminescence in fireflies (Lampyridae), one of the most charismatic lineages of insects, remains elusive. Here, we report the discovery of the second Mesozoic bioluminescent firefly, Flammarionella hehaikuni Cai, Ballantyne & Kundrata gen. et sp. nov., from the Albian/Cenomanian of northern Myanmar (ca 99 Ma). Based on the available set of diagnostic characters, we interpret the specimen as a female of stem-group Luciolinae. The fossil possesses deeply impressed oval pits on the apices of antennomeres 3–11, representing specialized sensory organs likely involved in olfaction. The light organ near the abdominal apex of Flammarionella resembles that found in extant light-producing lucioline fireflies. The growing fossil record of lampyrids provides direct evidence that the stunning light displays of fireflies were already established by the late Mesozoic.
... This clade was originally proposed to accommodate Lampyridae, Phengodidae, Rhagophthalmidae, and a recently described enigmatic bioluminescent genus, Sinopyrophorus Bi & Li in Bi et al. (2019), which was originally placed in Elateridae (Bi et al. 2019) but was treated as a separate family by Kusy et al. (2021). Later, Li et al. (2021) suggested that the newly described Mesozoic Cretophengodidae also belonged to the 'lampyroid clade' , hypothesizing its close relationships with Phengodidae and Rhagophthalmidae. Given that the interrelationships between the Elateridae and 'lampyroid' group of families are not yet fully understood (Douglas et al. 2021, Kusy et al. 2021, further phylogenomic studies are needed with an expanded taxon sampling of all above-mentioned families. ...
Phengodidae (Coleoptera: Elateroidea), commonly known as glowworm beetles, are a small family of bioluminescent and paedomorphic beetles. There are few phylogenetic studies of Phengodidae, and these are mostly discordant, especially when comparing morphology-based and molecular-based phylogenetic hypotheses. Here, we used the anchored hybrid enrichment approach to undertake the first phylogenomic analysis of Phengodidae (≤358 loci and 39 taxa) and evaluate the higher-level classification of the group. In agreement with previous molecular studies, we recovered Phengodidae as sister to Rhagophthalmidae, and the Old World Cydistinae as sister to all New World Phengodidae. In contrast to previous hypotheses, both Phengodinae and Mastinocerinae were each recovered as monophyletic. Cenophengus was found to be sister to Mastinocerinae, in contrast to some previous hypotheses that placed it as sister to all New World Phengodidae. Considering its morphological divergence, we here establish Cenophenginae subfam. nov. Despite the largest and most comprehensive sampling of Phengodidae in any molecular-based study to date, we had only limited success in revealing the relationships among genera within the most species-rich subfamily, Mastinocerinae. Further studies should focus on the phylogeny and classification of this taxonomically neglected subfamily, on the phylogenetic
placement of enigmatic Penicillophorinae, and on seeking morphological support for the main clades of Phengodidae.
... clade (Li, Kundrata, Tihelka, et al., 2021). Fossils of soft-bodied 'lampyroids' are rare in the Cretaceous fossil record. ...
... Almost 60 species classified in 13 genera have been described from burmite, and single species from Spanish amber, Azerbaijan Agdzhakend amber, and the Koonwarra Fossil Bed in Australia (e.g., Fanti & Müller, 2022;Hsiao et al., 2021;Qu et al., 2023). Li, Kundrata, Packova, et al. (2021) Deng et al. (2019), and several more species followed (e.g., Li, Kundrata, Tihelka, et al., 2021;. ...
Recent progress in beetle palaeontology has incited us to re-address the evolutionary history of the group. The Permian †Tshekardocoleidae had elytra that covered the posterior body in a loose tent-like manner. The formation of elytral epipleura and a tight fit of elytra and abdomen were important evolutionary transformations in the Middle Permian, resulting in a tightly enclosed subelytral space. Permian families were likely associated with dead wood of gymnospermous trees. The end-Permian extinction event resulted in a turnover in the composition of beetle faunas, especially a decline of large-bodied wood-associated forms. Adephaga and Myxophaga underwent a first wave of diversification in the Triassic. Polyphaga are very rare in this period. The first wave of diversification of this suborder occurs in the Jurassic, with fossils of Elateriformia, Staphyliniformia and Cucujiformia. The Cretaceous fossil record has been tremendously enriched by the discovery of amber inclusions. Numerous fossils represent all major polyphagan lineages and also the remaining suborders. Improved analytical methods for documenting and placing extinct taxa are discussed. Different factors have played a role in the diversification of beetles. The enormous number of species associated with flowering plants, and timing and patterns of diversification in phytophagous lineages indicate that the angio-sperm radiation played a major role in beetle macroevolution. Moreover, the evolution of intimate partnerships with symbionts and the acquisition of novel genes-obtained from fungi and bacteria via horizontal gene transfers-facilitated the use of plant material as a food source and were key innovations in the diversification of plant-feeding beetles.
... Several soft-bodied families, however, lack any fossil record at all. It is interesting, though, that from three described extinct families (with unresolved affinities within the superfamily), one is soft-bodied (Berendtimiridae; Winkler, 1987) and remaining two represent some intermediate stages (Mysteriomorphidae and Cretophengodidae;Alekseev &Ellenberger, 2019 andPeris et al., 2020 for the first, Li et al., 2021a for the latter). The genus Anoeuma Li et al., 2021 from Burmese amber, placed as incertae sedis within the superfamily Elateroidea, is also soft-bodied (Li et al., 2021b). ...
... The fossil family Cretophengodidae, which is supposedly closely related to Phengodidae and Rhagophthalmidae, is known exclusively from Burmese amber (Li et al., 2021a). The fact that Cretocydistus gen. ...
Elateroidea superfamily holds a huge diversity, morphological variation and a myriad of habitat specializations. The presence of bioluminescence and paedomorphosis renders the group as an interesting model for several studies. The “lampyroid” clade manifests both features, in a variety of light displays and body-forms, but the small fossil records hampers any advance in understanding the origin of these characteristics, as well as the biogeographic history of the group. We present here the description of a new fossil species, Cretocydistus wittmeri gen. et sp. nov. from the mid-Cretaceous of northern Myanmar, the first fossil of the family Phengodidae. We putatively place the genus in the subfamily Cydistinae, which extant species are distributed in Asia Minor, the Levant, and Iran. We also discuss how the discovery of this fossil taxa influences the study of the family and the “lampyroid” clade evolutionary history and biogeography.
... Bioluminescence in living things requires a chemical reaction, but unlike that of a burning wood, it involves the production of light without producing a burning heat. Firefly is among the most known animal for bioluminescence however bioluminescence can be observed in a variety of different organisms such as worms, insects, vertebrates, aquatic organisms of different taxa, plant and fungi as well as extinct species (1)(2)(3)(4)(5)(6). When we look at the concepts related to light production in living things in more detail, we see that the forms of radiation seen in the world are generally referred to as luminescence. ...
... 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. ...
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. ...
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 . ...
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]. ...
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.
