RESUMO
Snakeflies (Raphidioptera) are the smallest order of holometabolous insects that have kept their distinct and name-giving appearance since the Mesozoic, probably since the Jurassic, and possibly even since their emergence in the Carboniferous, more than 300 million years ago. Despite their interesting nature and numerous publications on their morphology, taxonomy, systematics, and biogeography, snakeflies have never received much attention from the general public, and only a few studies were devoted to their molecular biology. Due to this lack of molecular data, it is therefore unknown, if the conserved morphological nature of these living fossils translates to conserved genomic structures. Here, we present the first genome of the species and of the entire order of Raphidioptera. The final genome assembly has a total length of 669 Mbp and reached a high continuity with an N50 of 5.07 Mbp. Further quality controls also indicate a high completeness and no meaningful contamination. The newly generated data was used in a large-scaled phylogenetic analysis of snakeflies using shared orthologous sequences. Quartet score and gene concordance analyses revealed high amounts of conflicting signals within this group that might speak for substantial incomplete lineage sorting and introgression after their presumed re-radiation after the asteroid impact 66 million years ago. Overall, this reference genome will be a door-opening dataset for many future research applications, and we demonstrated its utility in a phylogenetic analysis that provides new insights into the evolution of this group of living fossils.
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Fósseis , Genoma , Animais , Filogenia , Genômica , Insetos/genéticaRESUMO
Inocelliidae is one of the two extant families of the holometabolan order Raphidioptera (snakeflies), with the modern fauna represented by seven genera and 44 species. The evolutionary history of the family is little-known. Here we present the first phylogenetic and biogeographical analyses based on a worldwide sampling of taxa and datasets combined with morphological characters and mitochondrial genomes, aiming to investigate the intergeneric phylogeny and historical biogeography of Inocelliidae. The phylogenetic inference from the combined analysis of morphological and molecular data recovered the sister-group relationship between a clade of (Negha + Indianoinocellia) + Sininocellia and a clade of Fibla + the Inocellia clade (interiorly nested by Amurinocellia and Parainocellia). Amurinocellia stat.r. and Parainocellia stat.r. et emend.n. are relegated to subgeneric status within Inocellia, whereas a newly erected subgenus of Inocellia, Epinocellia subgen.n., accommodates the former Parainocellia burmana (U. Aspöck and H. Aspöck, 1968) plus a new species Inocellia (Epinocellia) weii sp.n. Further, the Inocellia crassicornis group constitutes the nominotypical subgenus Inocellia stat.n., but the Inocellia fulvostigmata group is paraphyletic. Diversification within Inocelliidae is distinguished by an Eocene divergence leading to extant genera and a Miocene radiation of species. A biogeographical scenario depicts how the diverse inocelliid fauna from East Asia could have originated from western North America via dispersal across the Beringia during the early Tertiary, and how the Miocene ancestors of Inocellia could have accomplished long-distance dispersals via the Tibet-Himalayan corridor or eastern Palaearctic to western Palaearctic. Our results shed new light specifically on the evolution of Inocelliidae and, in general, the Raphidioptera.
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Holometábolos , Animais , Aprepitanto , Ásia Oriental , Insetos/genética , FilogeniaRESUMO
The sequential breakup of the supercontinent Pangaea since the Middle Jurassic is one of the crucial factors that has driven the biogeographical patterns of terrestrial biotas. Despite decades of effort searching for concordant patterns between diversification and continental fragmentation among taxonomic groups, increasing evidence has revealed more complex and idiosyncratic scenarios resulting from a mixture of vicariance, dispersal and extinction. Aquatic insects with discreet ecological requirements, low vagility and disjunct distributions represent a valuable model for testing biogeographical hypotheses by reconstructing their distribution patterns and temporal divergences. Insects of the order Megaloptera have exclusively aquatic larvae, their adults have low vagility, and the group has a highly disjunct geographical distribution. Here we present a comprehensive phylogeny of Megaloptera based on a large-scale mitochondrial genome sequencing of 99 species representing >90% of the world genera from all major biogeographical regions. Molecular dating suggests that the deep divergence within Megaloptera pre-dates the breakup of Pangaea. Subsequently, the intergeneric divergences within Corydalinae (dobsonflies), Chauliodinae (fishflies) and Sialidae (alderflies) might have been driven by both vicariance and dispersal correlated with the shifting continent during the Cretaceous, but with strikingly different and incongruent biogeographical signals. The austral distribution of many corydalids appears to be a result of colonization from Eurasia through southward dispersal across Europe and Africa during the Cretaceous, whereas a nearly contemporaneous dispersal via northward rafting of Gondwanan landmasses may account for the colonization of extant Eurasian alderflies from the south.
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Genoma Mitocondrial , Holometábolos , Animais , Genoma Mitocondrial/genética , Holometábolos/genética , Insetos/genética , Mitocôndrias/genética , FilogeniaRESUMO
The species diversity of insects is extraordinarily rich, but still has been insufficiently explored or underestimated particularly for uncommon groups. The pleasing lacewings (Dilaridae) are a little known family of Neuroptera with distinct sexually dimorphic antennae. The species diversity of pleasing lacewings was recently found to be severely underestimated and requires a comprehensive investigation, as well as systematic reviews. Here, we report on 12 new species of the pleasing lacewing genus Dilar Rambur, 1838, from the Oriental region, namely D. forcipatus sp. nov. and D. laoticus sp. nov. from Laos (new country record of Dilar); D. malickyi sp. nov., D. phraenus sp. nov. and D. rauschorum sp. nov. from northern Thailand; D. striatus sp. nov. from northern Vietnam; D. cangyuanensis sp. nov., D. daweishanensis sp. nov., D. nujianganus sp. nov., D. weibaoshanensis sp. nov., D. yucheni sp. nov., and D. zhangweiae sp. nov. from Yunnan and Tibet, both in southwestern China. The new species of Dilar display several types of wing marking patterns, and the morphology of the male genitalia is highly diverse. A comprehensive examination of the species diversity and distribution of Dilar concluded that Yunnan (southwestern China) represents a biogeographic region with high endemism and the richest species diversity. The potential correlation between vertical distribution and geographical latitude in Dilar was also analyzed.
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The family Berothidae (beaded lacewings) is poorly known from the Indochina Peninsula. Previously, three genera, Berotha Walker, 1860, Isoscelipteron Costa, 1863, and Lekrugeria Navás, 1929, and four species were recorded from Myanmar, Thailand, and Vietnam. Here, we record the genus Asadeteva U. Aspöck H. Aspöck, 1981, in Indochina for the first time and provide a description of a new species, namely Asadeteva acutata sp. nov., from Laos and Thailand. We also describe a new species of the genus Berotha Walker, 1860, namely Berotha incurvata sp. nov. from Laos and provide the first description of the male of Lekrugeria nepalica U. Aspöck H. Aspöck, 1986, based on material from northern Myanmar.
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Insetos , Animais , Indochina , MasculinoRESUMO
A new species and a new genus of Rhachiberothidae, Rhachiella malawica gen. nov., spec. nov., are described from Malawi. The new species is characterized by a flat vertex, a long penisfilum in the male, and by a bifurcate pseudohypocauda in the female. This combination of characters requires the description of a new genus, which is the sister taxon of Mucroberotha Tjeder, 1959. This is the first record of Rhachiberothidae in Malawi. The distributions of all 14 species of Rhachiberothidae so far known are shown in three maps.
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Holometábolos , Animais , Feminino , Malaui , MasculinoRESUMO
An amendment to this paper has been published and can be accessed via the original article.
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BACKGROUND: The latest advancements in DNA sequencing technologies have facilitated the resolution of the phylogeny of insects, yet parts of the tree of Holometabola remain unresolved. The phylogeny of Neuropterida has been extensively studied, but no strong consensus exists concerning the phylogenetic relationships within the order Neuroptera. Here, we assembled a novel transcriptomic dataset to address previously unresolved issues in the phylogeny of Neuropterida and to infer divergence times within the group. We tested the robustness of our phylogenetic estimates by comparing summary coalescent and concatenation-based phylogenetic approaches and by employing different quartet-based measures of phylogenomic incongruence, combined with data permutations. RESULTS: Our results suggest that the order Raphidioptera is sister to Neuroptera + Megaloptera. Coniopterygidae is inferred as sister to all remaining neuropteran families suggesting that larval cryptonephry could be a ground plan feature of Neuroptera. A clade that includes Nevrorthidae, Osmylidae, and Sisyridae (i.e. Osmyloidea) is inferred as sister to all other Neuroptera except Coniopterygidae, and Dilaridae is placed as sister to all remaining neuropteran families. Ithonidae is inferred as the sister group of monophyletic Myrmeleontiformia. The phylogenetic affinities of Chrysopidae and Hemerobiidae were dependent on the data type analyzed, and quartet-based analyses showed only weak support for the placement of Hemerobiidae as sister to Ithonidae + Myrmeleontiformia. Our molecular dating analyses suggest that most families of Neuropterida started to diversify in the Jurassic and our ancestral character state reconstructions suggest a primarily terrestrial environment of the larvae of Neuropterida and Neuroptera. CONCLUSION: Our extensive phylogenomic analyses consolidate several key aspects in the backbone phylogeny of Neuropterida, such as the basal placement of Coniopterygidae within Neuroptera and the monophyly of Osmyloidea. Furthermore, they provide new insights into the timing of diversification of Neuropterida. Despite the vast amount of analyzed molecular data, we found that certain nodes in the tree of Neuroptera are not robustly resolved. Therefore, we emphasize the importance of integrating the results of morphological analyses with those of sequence-based phylogenomics. We also suggest that comparative analyses of genomic meta-characters should be incorporated into future phylogenomic studies of Neuropterida.
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Evolução Molecular , Holometábolos/genética , Filogenia , Animais , Sequência de Bases , Genômica , Larva/genética , Análise de Sequência de DNA , TranscriptomaRESUMO
The pleasing lacewing genus Dilar Rambur, 1838, is poorly known from South Asia, previously only three species were accounted. Here, we present a revision of Dilar from South Asia, with descriptions of five new species, namely Dilar austroindicus sp. nov., Dilar biprojectus sp. nov., Dilar clavatus sp. nov., Dilar miralobatus sp. nov., and Dilar truncatus sp. nov. In addition, Dilar nietneri Hagen, 1858, from Sri Lanka is re-described. A key to males of Dilar from South Asia is provided.
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Holometábolos , Insetos , Distribuição Animal , Animais , MasculinoRESUMO
The pleasing lacewing genus Dilar Rambur, 1838, is the only known representative of the family Dilaridae from Central Asia. Here we present a review of the five species of Dilar from Central Asia, Dilar hornei McLachlan, 1869, Dilar caesarulus H. Aspöck U. Aspöck, 1967, Dilar kirgisus H. Aspöck U. Aspöck, 1967, Dilar vartianorum H. Aspöck U. Aspöck, 1967, and Dilar dochaner H. Aspöck U. Aspöck, 1968. Remarkable intraspecific variations in male gonocoxite 10 are found in D. kirgisus and D. vartianorum. Re-descriptions of all the species are provided. Dilar indicus Monserrat, 1989, and Dilar similis Monserrat, 1989, are synonymized with D. vartianorum. A key to the species of Dilar from Central Asia is provided.
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Holometábolos , Insetos , Animais , Ásia , MasculinoRESUMO
The lacewing family Berothidae (beaded lacewings) of China is a poorly known group. It consists of two genera and 11 previously described species. In this paper, we provide a revision of the beaded lacewings from China. The genus Berotha Walker, 1860, in China contains seven species including one new species: Berotha guangdongana sp. nov. The second genus, Isoscelipteron Costa, 1863, comprises six species from China, including one new species, namely Isoscelipteron acuticaudatum sp. nov. Descriptions and re-descriptions of the species are provided as well as an identification key to the species of Berothidae from China.
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Distribuição Animal , Insetos , Animais , ChinaRESUMO
Two new snakefly species of the family Raphidiidae are described from China: Mongoloraphidia lini sp. nov. and Mongoloraphidia triangulata sp. nov. Biogeographical considerations on secondary refugial centers of the large Sino-Tibetan primary center are presented.
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Holometábolos , Insetos , Animais , ChinaRESUMO
Neuroptera (lacewings) and allied orders Megaloptera (dobsonflies, alderflies) and Raphidioptera (snakeflies) are predatory insects and together make up the clade Neuropterida. The higher-level relationships within Neuropterida have historically been widely disputed with multiple competing hypotheses. Moreover, the evolution of important biological innovations among various Neuropterida families, such as the origin, timing and direction of transitions between aquatic and terrestrial habitats of larvae, remains poorly understood. To investigate the origin and diversification of lacewings and their allies, we undertook phylogenetic analyses of mitochondrial genomes of all families of Neuropterida using Bayesian inference, maximum likelihood and maximum parsimony methods. We present a robust, fully resolved phylogeny and divergence time estimation for Neuropterida with strong statistical support for almost all nodes. Mitochondrial sequence data are typified by significant compositional heterogeneity across lineages, and parsimony and models assuming homogeneous rates did not recover Neuroptera as monophyletic. Only a model accounting for compositional heterogeneity (i.e. CAT-GTR) recovered all orders of Neuropterida as monophyletic. Significant findings of the mitogenomic phylogeny include recovering Raphidioptera as sister to Megaloptera plus Neuroptera. The sister family of all other lacewings are the dusty-wings (Coniopterygidae), rather than Nevrorthidae. Nevrorthidae are instead returned to their traditional position as the sister group of the spongilla-flies (Sisyridae) and closely related to Osmylidae. Our divergence time analysis indicates that the Mesozoic was indeed a 'golden age' for lacewings, with most families of Neuropterida diverging during the Triassic and Jurassic and all extant families present by the Early Cretaceous. Based on ancestral character state reconstructions of larval habitat we evaluate competing hypotheses regarding the life style of early neuropteridan larvae as either aquatic or terrestrial.
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BACKGROUND: The occurrence of phlebotomine sand flies in Central Europe was questioned until they were recorded for the first time in Germany in 1999, and ten years later also in Austria. The aim of this study was to investigate sand flies collected in Austria for their carrier status of Leishmania spp. FINDINGS: From 2012 to 2013 field studies were conducted in eastern Austria. Altogether, 22 individuals of sand flies were found, all morphologically identified as Phlebotomus (Transphlebotomus) mascittii Grassi, 1908. Twelve non-engorged female specimens with no visible remnants of a blood meal in their bodies were individually investigated for Leishmania spp. by ITS-1 real-time PCR. One out of these was positive for Leishmania, identified as Leishmania infantum by DNA sequencing. This finding suggests that L. infantum is not excreted by P. mascittii and possibly can establish an infection within P. mascittii. Interestingly, an asymptomatic dog living on the farm where this sand fly had been caught was also Leishmania-positive. CONCLUSIONS: This study provides new data on the suspected vector capacity of P. mascittii, being the northernmost sand fly species in Europe and in most central European regions the only sand fly species found. Proven vector capacity of P. mascittii for Leishmania spp. would be of significant medico-veterinary importance, not only with respect to expanding sand fly populations in Central Europe related to global warming, but also in the light of globalization and increasing movements of humans.
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Insetos Vetores/parasitologia , Leishmania infantum/fisiologia , Phlebotomus/classificação , Phlebotomus/parasitologia , Distribuição Animal , Animais , Áustria , Feminino , Interações Hospedeiro-Parasita , MasculinoRESUMO
The lacewing family Dilaridae (pleasing lacewings) is poorly known in Southeast Asia, currently with only five described species. In this paper, we provide a revision of the species of the genus Dilar Rambur, 1838 from Southeast Asia. Eleven species of Dilar are recorded in this region, with seven species herein described as new to science, i.e. Dilar abnormis Zhang, Liu & Winterton, sp. nov., Dilar lineatus Zhang, Liu & Winterton, sp. nov., Dilar loeinensis Zhang, Liu, Winterton, sp. nov., Dilar ohli Zhang, Liu, Aspöck & Aspöck, sp. nov., Dilar rotundatus Zhang, Liu & Winterton, sp. nov., Dilar sumatranus Zhang, Liu, Aspöck & Aspöck, sp. nov., and Dilar zimmermannae Zhang, Liu, Aspöck & Aspöck, sp. nov. Re-descriptions of Dilar grandis (Banks, 1931), and Dilar marmoratus (Banks, 1931) are also provided. Dilaridae are recorded in Indonesia (Sumatra), Myanmar, and northern Thailand for the first time. A key to the Dilar species from Southeast Asia is given.
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Insetos/classificação , Distribuição Animal , Estruturas Animais/anatomia & histologia , Estruturas Animais/crescimento & desenvolvimento , Animais , Sudeste Asiático , Tamanho Corporal , Feminino , Insetos/anatomia & histologia , Insetos/crescimento & desenvolvimento , Masculino , Tamanho do ÓrgãoRESUMO
A revision of species of the genus Dilar Rambur, 1838, from the southern part of mainland China is presented. Twenty species are recorded in this region, with 12 species described as new to science. Dilar spectabilis Zhang, Liu, H. Aspöck & U. Aspöck, 2014, is recorded in this region for the first time. The other seven valid species previously recorded in this region are re-described. Dilar wangi Yang, 1992, is herein treated as a junior synonym of Dilar montanus Yang, 1992. A key to all species of the genus Dilar so far recorded from China is provided.
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Insetos/classificação , Distribuição Animal , Animais , China , Feminino , Insetos/anatomia & histologia , Masculino , Especificidade da EspécieRESUMO
Fauna Europaea provides a public web-service with an index of scientific names of all living European land and freshwater animals, their geographical distribution at country level (up to the Urals, excluding the Caucasus region), and some additional information. The Fauna Europaea project covers about 230,000 taxonomic names, including 130,000 accepted species and 14,000 accepted subspecies, which is much more than the originally projected number of 100,000 species. This represents a huge effort by more than 400 contributing specialists throughout Europe and is a unique (standard) reference suitable for many users in science, government, industry, nature conservation and education. For Neuropterida, data from three Insect orders (Raphidioptera, Megaloptera, Neuroptera), comprising 15 families and 397 species, are included.
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Three species of the genus Dilar Rambur, 1838, are recorded and described from Tibet in southwestern China, including Dilar geometroides H. Aspöck & U. Aspöck, 1968, Dilar harmandi (Navás, 1909), and Dilar tibetanus Yang, 1987, with the former two species recorded in Tibet for the first time. All three species are redescribed. Dilar aspersus Yang, 1988, and Dilar pusillus Yang, 1992, are synonymized with Dilar geometroides Aspöck & Aspöck, 1968, and Dilar tibetanus Yang, 1987, respectively. A key to the Dilar species from Tibet is provided.