Genome analysis of Phrixothrix hirtus (Phengodidae) railroad worm shows the expansion of odorant-binding gene families and positive selection on morphogenesis and sex determination genes.

Among bioluminescent beetles of the Elateroidea superfamily, Phengodidae is the third largest family, with 244 bioluminescent species distributed only in the Americas, but is still the least studied from the phylogenetic and evolutionary points of view. The railroad worm Phrixothrix hirtus is an essential biological model and symbolic species due to its bicolor bioluminescence, being the only organism that produces true red light among bioluminescent terrestrial species. Here, we performed partial genome assembly of P. hirtus, combining short and long reads generated with Illumina sequencing, providing the first source of genomic information and a framework for comparative analyses of the bioluminescent system in Elateroidea. This is the largest genome described in the Elateroidea superfamily, with an estimated size of ∼3.4 Gb, displaying 32 % GC content, and 67 % transposable elements. Comparative genomic analyses showed a positive selection of genes and gene family expansion events of growth and morphogenesis gene products, which could be associated with the atypical anatomical development and morphogenesis found in paedomorphic females and underdeveloped males. We also observed gene family expansion among distinct odorant-binding receptors, which could be associated with the pheromone communication system typical of these beetles, and retrotransposable elements. Common genes putatively regulating bioluminescence production and control, including two luciferase genes corresponding to lateral lanterns green-emitting and head lanterns red-emitting luciferases with 7 exons and 6 introns, and genes potentially involved in luciferin biosynthesis were found, indicating that there are no clear differences about the presence or absence of gene families associated with bioluminescence in Elateroidea.

Rhagophthalmidae Olivier, 1907 (Coleoptera, Elateroidea): described genera and species, current problems, and prospects for the bioluminescent and paedomorphic beetle lineage.

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 Haplocladongorhami Kundrata, nom. nov. is proposed for Diplocladonhasseltii Gorham, 1883b (described in subgenus Haplocladon) which is preoccupied by Diplocladonhasseltii Gorham, 1883a. The genus Reductodrilus Pic, 1943 is tentatively placed in Lampyridae: Ototretinae. Lectotypes are designated for Pseudothilmanusalatus 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.

Anchored Phylogenomics, Evolution and Systematics of Elateridae: Are All Bioluminescent Elateroidea Derived Click Beetles?

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.

Organization and comparative analysis of the mitochondrial genomes of bioluminescent Elateroidea (Coleoptera: Polyphaga).

Mitochondrial genome organization in the Elateroidea superfamily (Coleoptera), which include the main families of bioluminescent beetles, has been poorly studied and lacking information about Phengodidae family. We sequenced the mitochondrial genomes of Neotropical Lampyridae (Bicellonycha lividipennis), Phengodidae (Brasilocerus sp.2 and Phrixothrix hirtus) and Elateridae (Pyrearinus termitilluminans, Hapsodrilus ignifer and Teslasena femoralis). All species had a typical insect mitochondrial genome except for the following: in the elaterid T. femoralis genome there is a non-coding region between NADH2 and tRNA-Trp; in the phengodids Brasilocerus sp.2 and P. hirtus genomes we did not find the tRNA-Ile and tRNA-Gln. The P. hirtus genome showed a ~1.6kb non-coding region, the rearrangement of tRNA-Tyr, a new tRNA-Leu copy, and several regions with higher AT contents. Phylogenetics analysis using Bayesian and ML models indicated that the Phengodidae+Rhagophthalmidae are closely related to Lampyridae family, and included Drilus flavescens (Drilidae) as an internal clade within Elateridae. This is the first report that compares the mitochondrial genomes organization of the three main families of bioluminescent Elateroidea, including the first Neotropical Lampyridae and Phengodidae. The losses of tRNAs, and translocation and duplication events found in Phengodidae mt genomes, mainly in P. hirtus, may indicate different evolutionary rates in these mitochondrial genomes. The mitophylogenomics analysis indicates the monophyly of the three bioluminescent families and a closer relationship between Lampyridae and Phengodidae/Rhagophthalmidae, in contrast with previous molecular analysis.

Survey of Bioluminescent Coleoptera in the Atlantic Rain Forest of Serra da Paranapiacaba in São Paulo State (Brazil) / Levantamento de Coleópteros bioluminescentes na Mata Atlântica da Serra da Paranapiacaba

Brazil is the country with the largest number of bioluminescent beetle species in the world. However, estimates suggest that this number could much be higher, since many species remain to be discovered. In this work we made a survey of the species of bioluminescent beetles in Serra de Paranapiacaba - the largest remnant of Atlantic Forest in São Paulo State. The survey was done at Intervales State Park, Carlos Botelho State Park and municipality of Tapiraí and the following species were collected: Aspisoma lineatum, Aspisoma physonotum, Aspisoma fenestrata, Cratomorphus besckey, Cratomorphus distinctus, Photinus penai, Photinus sp1, Photinus sp9, Ethra aff. malledicta or axilaris, Ethra aff. adicta , Lucidotini incertae sedis, Cladodes flabellicornis, Cladodes demoulini, Amydetes lucernuta, Bicellonycha sp8, Bicellonycha ornaticollis, Pyrogaster lunifer, Pyrogaster moestus, Pyrogaster sp2, Pyrogaster sp5, Pyrogaster sp6, Photuris lugubris, Photuris sp1, Photuris sp7, Stenophrixotrix sp1, Brasilocerus sp1, Pseudophengodes sp1, Hapsodrilus pyrotis, Hypsiophtalmus sp1, Ptesimopsia luculenta, Pyroptesis cincticollis, Pyrearinus brevicolis, Pyrearinus candelarius, Pyrearinus micatus, Pyrophorus divergens. Our data show that Serra de Paranapiacaba is the second richest area in São Paulo state, especially in elaterids, with unique species typical of this area and species common to other investigated sites such as the Biological Station of Boracéia (in Salesópolis county) and the urbanized areas in the between Campinas - Sorocaba- São Paulo, originally covered to the Atlantic Rainforest.

O Brasil é o país que possui o maior número de espécies de coleópteros bioluminescentes no mundo. Entretanto, estimativas sugerem que este número possa ser bem maior, dado que muitas espécies ainda não foram descritas. Neste trabalho foi realizado um levantamento das espécies de coleópteros bioluminescentes em três localidades na Serra da Paranapiacaba - a maior área remanescente contígua de Mata Atlântica no país, com associação ao respectivo habitat. No Parque Estadual Intervales, Parque Estadual Carlos Botelho e Tapiraí foram coletadas as seguintes espécies: (Lampyidae) Aspisoma lineatum, Aspisoma physonotum, Aspisoma fenestrata, Cratomorphus besckey, Cratomorphus distinctus, Photinus penai, Photinus sp1, Photinus sp9, Ethra aff. malledicta ou axilaris, Ethra aff. adicta, Lucidotini incertae sedis, Cladodes flabellicornis, Cladodes demoulini, Amydetes sp1, Bicellonycha sp8, Bicellonycha ornaticollis, Pyrogaster lunifer, Pyrogaster moestus, Pyrogaster sp2, Pyrogaster sp5, Pyrogaster sp6, Photuris lugubris, Photuris sp1, Photuris sp7; (Phengodidae) Stenophrixotrix sp1, Brasilocerus sp2, Pseudophengodes sp1; (Elateridae) Hapsodrilus pyrotis, Hypsiophtalmus sp1, Ptesimopsia luculenta, Pyroptesis cincticollis, Pyrearinus brevicolis, Pyrearinus candelarius, Pyrearinus micatus, Pyrophorus divergens. Estes dados mostram que esta constitui a segunda área mais rica em espécies luminescentes do Estado de São Paulo, depois da Est. Biológica de Boracéia, especialmente em elaterídeos, com espécies únicas características destas localidades e espécies comuns è outras áreas investigadas, como as da Estação Biológica de Boracéia (Salesópolis, SP) e áreas urbanizadas no triângulo Campinas - Sorocaba - São Paulo, originalmente cobertas por Mata Atlântica.

Molecular phylogeny of Neotropical bioluminescent beetles (Coleoptera: Elateroidea) in southern and central Brazil.

Bioluminescence in beetles is found mainly in the Elateroidea superfamily (Elateridae, Lampyridae and Phengodidae). The Neotropical region accounts for the richest diversity of bioluminescent species in the world with about 500 described species, most occurring in the Amazon, Atlantic rainforest and Cerrado (savanna) ecosystems in Brazil. The origin and evolution of bioluminescence, as well as the taxonomic status of several Neotropical taxa in these families remains unclear. In order to contribute to a better understanding of the phylogeny and evolution of bioluminescent Elateroidea we sequenced and analyzed sequences of mitochondrial NADH2 and the nuclear 28S genes and of the cloned luciferase sequences of Brazilian species belonging to the following genera: (Lampyridae) Macrolampis, Photuris, Amydetes, Bicellonycha, Aspisoma, Lucidota, Cratomorphus; (Elateridae) Conoderus, Pyrophorus, Hapsodrilus, Pyrearinus, Fulgeochlizus; and (Phengodidae) Pseudophengodes, Phrixothrix, Euryopa and Brasilocerus. Our study supports a closer phylogenetic relationship between Elateridae and Phengodidae as other molecular studies, in contrast with previous morphologic and molecular studies that clustered Lampyridae/Phengodidae. Molecular data also supported division of the Phengodinae subfamily into the tribes Phengodini and Mastinocerini. The position of the genus Amydetes supports the status of the Amydetinae as a subfamily. The genus Euryopa is included in the Mastinocerini tribe within the Phengodinae/Phengodidae.

Bioluminescent Coleoptera of Biological Station of Boracéia (Salesópolis, SP, Brazil): diversity, bioluminescence and habitat distribution / Coleópteros Bioluminescentes da Estação Biológica de Boracéia (Salesópolis, SP, Brasil): diversidade, bioluminescência e distribuição por habitat

Brazil hosts the richest biodiversity of bioluminescent beetles in the world. Several species are found in the Atlantic rain forest, one of the richest and most threatened tropical forests in the world. We have catalogued the biodiversity of bioluminescent species mainly of Elateroidea superfamily occurring in one of the last largest and most preserved remnants of Atlantic rain forest, located at the Biological Station of Boracéia of São Paulo University (Salesopolis, SP, Brazil). This site accounted with the largest diversity ever found for a single place in São Paulo State, with 39 species: Lampyridae (30), Phengodidae (5), Elateridae (3) and Staphylinidae (1). This fauna has unique species that were not found in any other places of Atlantic forest in São Paulo state, especially fireflies from the Lampyrinae tribes (Cratomorphini, Lamprocerini, Lucidotini), the subfamilies Amydetinae and Photurinae, and the phengodids Pseudophengodes and Brasilocerus sp.2. Most species are found in dense Ombrophyl forest or at their border, and a few ones are found dwelling in the few open fields around the forest. There is a predominance of glowing patterns in the green region among forest inhabiting species when compared with open field fireflies.

O Brasil tem a maior biodiversidade de coleópteros bioluminescentes do mundo. Muitas destas espécies são encontradas na Mata Atlantica, um dos mais ricos e ameaçados ecossitemas. Catalogamos a biodiversidade de espécies bioluminescentes da superfamília Elateroidea em um dos maiores e mais preservados remanescentes de Mata Atlântica da Serra do Mar, localizado na Estação Biológica de Boraceia (E.B.B.) em Salesopolis, SP, reserva administrada pelo Museu de Zoologia da Universidade de São Paulo. Este local contou com a maior biodiversidade de espécies bioluminescentes já encontrada num único local dentro do estado, com 39 espécies: Lampyridae (30), Phengodidae (5), Elateridae (3) e Staphylinidae (1). Esta fauna tem espécies únicas não encontradas em nenhum outro lugar investigado de Mata Atlântica no estado de São Paulo, especialmente das tribos de Lampyrinae (Cratomorphini, Lamprocerini, Lucidotini), as subfamílias Amydetinae e Photurinae, e os fengodídeos Pseudophengodes e Brasilocerus sp.2. A maioria das espécies ocorre em floresta ombrófila densa, e poucas ocorrem em campos abertos. Existe uma predominância de espécies com padrões de brilho contínuo na região do verde no interior da mata quando comparadas a espécies de vagalumes de campo aberto.

Fauna de besouros bioluminescentes (Coleoptera: Elateroidea: Lampyridae; Phengodidae, Elateridae) nos municípios de Campinas, Sorocaba-Votorantim e Rio Claro-Limeira (SP, Brasil): biodiversidade e influência da urbanização / Bioluminescent beetles (Coleoptera: Elateroidea: Lampyridae, Phengodidae, Elateridae) in the municipalities of Campinas, Sorocaba-Votorantim and Rio Claro-Limeira (SP, Brazil): biodiversity and influence of urban sprawl

O Brasil abriga a maior biodiversidade de besouros bioluminescentes do mundo. Apesar disto, levantamentos faunísticos regionais e estudos sobre o efeito da urbanização na biodiversidade de vaga-lumes são inexistentes. Este tipo de estudo pode ser especialmente importante para selecionar novos bioindicadores de ambientes noturnos. A biodiversidade de espécies bioluminescentes de Elateroidea em três grandes áreas urbanas no Estado de São Paulo: os municípios de Campinas, Sorocaba-Votorantim e Rio Claro-Limeira foi catalogada. Em Campinas, a ocorrência de espécies tem sido monitorada durante os últimos 20 anos. Foi registrada a ocorrência de 26 espécies em Campinas, 21 em Sorocaba e 19 em Rio Claro, sendo estas encontradas principalmente em áreas de Mata Atlântica, crescimentos secundários, brejos e campos. Apesar de vaga-lumes lampirídeos serem encontrados em todos os habitats, elaterídeos e fengodídeos ocorreram encontrados preferencialmente em áreas florestadas. A expansão urbana, em especial a iluminação artificial tem impacto evidente na biodiversidade deste grupo de insetos. Os lampirídeos Aspisoma lineatum Gyllendal, 1817 (Lampyrinae: Cratomorphini) e Bicellonychia lividipennis Motschulsky, 1854 (Photurinae) são as espécies mais frequentes em areas urbanizadas, ao passo que várias outras espécies de lampirídeos, elaterídeos e fengodídeos são as mais incomuns nestas áreas. Estes estudos oferecem subsídios para o uso deste grupo de vaga-lumes como potenciais bioindicadores noturnos.

Brazil hosts the richest biodiversity of bioluminescent beetles in the world. However, no surveys on the regional biodiversity have been made. Furthermore, studies on the effect of urbanization on bioluminescent beetle's biodiversity are still missing. Such kind of studies could be especially helpful to select new night environment bioindicators. Bioluminescent species of the Elateroidea superfamily occurring in three large and expanding urban areas in São Paulo State: Campinas, Sorocaba-Votorantim and Rio Claro-Limeira municipalities, were catalogued. In Campinas, the occurrence of bioluminescent beetles in different sites has been monitored during the past 20 years. There were 26 species in Campinas, 21 in Sorocaba-Votorantim and 19 in Rio Claro-Limeira, occurring mainly in conserved remnants of Atlantic Rain-forest, secondary growths, marshes and open fields. Whereas fireflies (Lampyridae) are found in almost all habitats, click beetles (Elateridae) and railroadworms (Phengodidae) occur predominantly in woody environments. Urban sprawl has an evident impact on the biodiversity of this selected group of beetles, and special attention should be given to the effect of artificial night lighting in the occurrence of these beetles. The lampyrids Aspisoma lineatum Gyllenhal, 1817 (Lampyrinae: Cratomorphini) and Bicellonychia lividipennis Motschulsky 1854 (Photurinae) are the most common species in urbanized areas, whereas many other species of lampyrids, elaterids and phengodids are rarely found in such environments. These studies offer insights on the potential use of a selected group of luminescent beetles found in this region as nocturnal environment bioindicators.