This sequence is from Zorotypus huxleyi [Genbank:JN192451]. The length-variable regions are indicated in red. And the unique indels are marked with green color. The D3-4 box was highlighted with thick red lines. The Da–Dj numbering system for LVRs, which has not been taken into account previously, is a supplementary system to the D1–D12 coding system. Base pairing is indicated as follows: standard canonical pairs by lines (C-G, G-C, A-U, U-A); wobble G·U pairs by dots (G·U); A·G and A·C pairs by open circles (A G, A C); other non-canonical pairs by filled circles (e.g., U•U).

This sequence is from Zorotypus huxleyi [Genbank:JN192451]. The length-variable regions are indicated in red. And the unique indels are marked with green color. The D3-4 box was highlighted with thick red lines. The Da–Dj numbering system for LVRs, which has not been taken into account previously, is a supplementary system to the D1–D12 coding system. Base pairing is indicated as follows: standard canonical pairs by lines (C-G, G-C, A-U, U-A); wobble G·U pairs by dots (G·U); A·G and A·C pairs by open circles (A G, A C); other non-canonical pairs by filled circles (e.g., U•U).

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The position of the Zoraptera remains one of the most challenging and uncertain concerns in ordinal-level phylogenies of the insects. Zoraptera have been viewed as having a close relationship with five different groups of Polyneoptera, or as being allied to the Paraneoptera or even Holometabola. Although rDNAs have been widely used in phylogenetic...

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... Kirstová et al.[29]; 2 Yoshizawa and Johnson[23];3 Ma et al.[24];4 Terry and Whiting[10];5 Terry and Whiting (unpublished);6 Wang et al.[49]; * chimaeric taxon (isolate 002 for 18S rRNA, isolate 016 for 16S rRNA and COI mtDNA). ...
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Zoraptera is a small and predominantly tropical insect order with an unresolved higher classification due to the extremely uniform external body morphology. We, therefore, conducted a multigene molecular phylogeny of extant Zoraptera and critically re-evaluated their morphological characters in order to propose a natural infraordinal classification. We recovered a highly-resolved phylogeny with two main clades representing major evolutionary lineages in Zoraptera, for which we propose family ranks. The two families exhibit striking differences in male genitalia and reproductive strategies. Each family contains two subclades (subfamilies) supported by several morphological synapomorphies including the relative lengths of the basal antennomeres, the number and position of metatibial spurs, and the structure of male genitalia. The newly proposed higher classification of Zoraptera includes the family Zorotypidae stat. revid. with Zorotypinae Silvestri, 1913 (Zorotypus stat. revid., Usazoros Kukalova-Peck and Peck, 1993 stat. restit.) and Spermozorinae subfam. nov. (Spermozoros gen. nov.), and Spriralizoridae fam. nov. with Spiralizorinae subfam. nov. (Spiralizoros gen. nov., Scapulizoros gen. nov., Cordezoros gen. nov., Centrozoros Kukalova-Peck and Peck, 1993, stat. restit., Brazilozoros Kukalova-Peck and Peck, 1993, stat. restit.), and Latinozorinae subfam. nov. (Latinozoros Kukalova-Peck and Peck, 1993, stat. restit.). An identification key and morphological diagnoses for all supraspecific taxa are provided.
... NKU-011 from China (Accession no. JQ309927) [24] and clustered together with H. tuberculatus from Japan (Accession no. GU569180) [25]. ...
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Cattle lice are obligatory blood-sucking parasites, which is the cause of animal health problems worldwide. Recently, several studies have revealed that pathogenic bacteria could be found in cattle lice, and it can act as a potential vector for transmitting louse-borne diseases. However, the cattle lice and their pathogenic bacteria in Thailand have never been evaluated. In the present study, we aim to determine the presence of bacterial pathogens in cattle lice collected from three localities of Thailand. Total genomic DNA was extracted from 109 cattle louse samples and the Polymerase Chain Reaction (PCR) of 18S rRNA was developed to identify the cattle louse. Moreover, PCR was used for screening Bartonella spp., Acinetobacter spp., and Rickettsia spp. in cattle louse samples. The positive PCR products were cloned and sequenced. The phylogenetic tree based on the partial 18S rRNA sequences demonstrated that cattle lice species in this study are classified into two groups according to reference sequences; Haematopinus quadripertusus and Haematopinus spp. closely related to H. tuberculatus. The pathogen detection revealed that Bartonella spp. DNA of gltA and rpoB were detected in 25 of 109 samples (22.93%) both egg and adult stages, whereas Acinetobacter spp. and Rickettsia spp. were not detected in all cattle lice DNA samples. The gltA and rpoB sequences showed that the Bartonella spp. DNA was found in both H. quadripertusus and Haematopinus spp. closely related to H. tuberculatus. This study is the first report of the Bartonella spp. detected in cattle lice from Thailand. The finding obtained from this study could be used to determine whether the cattle lice can serve as a potential vector to transmit these pathogenic bacteria among cattle and may affect animal to human health.
... Owing to their many peculiarities, consensus regarding a precise higher placement of Zoraptera among other polyneopteran lineages has yet to be achieved (summarized by Mashimo et al., 2014), with hypotheses supporting relationships as disparate as sister to Dictyoptera (e.g., Yoshizawa and Johnson, 2005;Wang et al., 2013), Embiodea (e.g., Engel and Grimaldi, 2000;Grimaldi and Engel, 2005;Yoshizawa, 2007Yoshizawa, , 2011, Dermaptera (Misof et al., 2014), or even outside of Polyneoptera and as sister to Paraneoptera (e.g., Hennig, 1969;Kristensen, 1981;Beutel and Weide, 2005). While studies based on Recent taxa alone have failed to resolve conclusively the phylogenetic affinities of this highly autapomorphic group, it may be hoped that the early fossil record of the order might shed some additional light on the matter (e.g., Engel and Grimaldi, 2002), particularly if stem-group zorapterans could be discovered in pre-Cretaceous deposits. ...
... For the nrDNAs, sequences were initially aligned with the program Muscle embedded within MEGA 7.0.20 (Edgar, 2004;Kumar et al., 2016). The alignments of 18S and 28S rDNAs were then manually checked and corrected based on the secondary structure models of 18S and 28S rRNAs of Cantao ocellatus (Scutelleridae: Scutellerinae), respectively (Figs S1-S3), which were reconstructed referring to the universal models of 18S rRNA (Xie et al., 2008(Xie et al., , 2009) and 28S rRNA (Gillespie et al., 2006;Wang et al., 2013) for Hexapoda. The program RNAstructure 5.8.1 based on the thermodynamic folding (Reuter and Mathews, 2010) was used to reconstruct the secondary structures. ...
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Members of the family Scutelleridae (Heteroptera: Pentatomomorpha: Pentatomoidea) are also called shield bugs because of the greatly enlarged scutellum, or jewel bugs because of the brilliant colours of many species. All scutellerids are phytophagous, feeding on various parts of their host plants. Due to lack of obvious synapomorphies and the failure to apply rigorous phylogenetic methods, the higher classification of Scutelleridae has been disputed for more than 150 years. Here we reconstructed a phylogeny of Scutelleridae based on complete sequences of 18S and 28S nuclear rDNAs and all 13 protein-coding genes of the mitochondrial genome, with the sampled taxa covering all of the currently recognized subfamilies. The monophyly of Scutelleridae was confirmed by the congruence of the results of analyses conducted using Bayesian inference, maximum likelihood and maximum parsimony. The phylogenetic relationships among subfamilies were well resolved for the first time. Furthermore, time-divergence studies estimated that the time of origin of Scutelleridae was in the Early Cretaceous (142.1–122.8 Ma), after the origin of the angiosperms. The diversification between the extant subfamilies of Scutelleridae and within the subfamilies occurred from the late Palaeocene to the late Miocene, simultaneously with the rise of the major groups of angiosperms and other phytophagous insects.
... Thus they are typically described as gregarious, colony-dwelling insects. Recent studies on Zoraptera deal with the morphology of the head (Beutel and Weide, 2005;Wipfler and Pass, 2014;Matsumura et al., 2015), the thoracic skeleto-muscular system (Friedrich and Beutel, 2008), rectal pads (Dallai et al., 2016), eggs , embryonic and postembryonic development (Mashimo et al., 2014a,b), mating behaviour (Choe, 1994a,b;1995;Dallai et al., 2013), male and female reproductive systems (Hünefeld, 2007;Dallai et al., 2011Dallai et al., , 2012aDallai et al., ,b, 2013Dallai et al., , 2014aDallai et al., ,b, 2015Matsumura et al., 2014), phylogeny (Kukalov a-Peck and Peck, 1993;Rasnitsyn, 1998;Grimaldi and Engel, 2005;Yoshizawa and Johnson, 2005;Yoshizawa, 2007Yoshizawa, , 2011Friedrich and Beutel, 2008;Ishiwata et al., 2011;Wang et al., 2013Wang et al., , 2013Wang et al., , 2016Simon, 2013, 2016;Ma et al., 2014;Mashimo et al., 2014c). Their systematic position is still controversial (see Mashimo et al., 2014c), even though the placement in Polyneoptera is confirmed (Yoshizawa and Johnson, 2005;Mashimo et al., 2014a;Wipfler and Pass, 2014). ...
... Thus they are typically described as gregarious, colony-dwelling insects. Recent studies on Zoraptera deal with the morphology of the head (Beutel and Weide, 2005;Wipfler and Pass, 2014;Matsumura et al., 2015), the thoracic skeleto-muscular system (Friedrich and Beutel, 2008), rectal pads (Dallai et al., 2016), eggs , embryonic and postembryonic development (Mashimo et al., 2014a,b), mating behaviour (Choe, 1994a,b;1995;Dallai et al., 2013), male and female reproductive systems (Hünefeld, 2007;Dallai et al., 2011Dallai et al., , 2012aDallai et al., ,b, 2013Dallai et al., , 2014aDallai et al., ,b, 2015Matsumura et al., 2014), phylogeny (Kukalov a-Peck and Peck, 1993;Rasnitsyn, 1998;Grimaldi and Engel, 2005;Yoshizawa and Johnson, 2005;Yoshizawa, 2007Yoshizawa, , 2011Friedrich and Beutel, 2008;Ishiwata et al., 2011;Wang et al., 2013Wang et al., , 2013Wang et al., , 2016Simon, 2013, 2016;Ma et al., 2014;Mashimo et al., 2014c). Their systematic position is still controversial (see Mashimo et al., 2014c), even though the placement in Polyneoptera is confirmed (Yoshizawa and Johnson, 2005;Mashimo et al., 2014a;Wipfler and Pass, 2014). ...
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The salivary glands of two species of Zoraptera, Zorotypus caudelli and Zorotypus hubbardi, were examined and documented mainly using transmission electron microscopy (TEM). The results obtained for males and females of the two species are compared and functional aspects related to ultrastructural features are discussed. The salivary glands are divided into two regions: the secretory cell region and the long efferent duct, the latter with its distal end opening in the salivarium below the hypopharyngeal base. The secretory region consists of a complex of secretory cells provided with microvillated cavities connected by short ectodermal ducts to large ones, which are connected with the long efferent duct. The secretory cell cytoplasm contains a large system of rough endoplasmic reticulum and Golgi apparatuses producing numerous dense secretions. The cells of the efferent duct, characterized by reduced cytoplasm and the presence of long membrane infoldings associated with mitochondria, are possibly involved in fluid uptaking from the duct lumen.
... Due to the improved taxon sampling and/or a better alignment of the molecular sequences, the results of different phylogenetic studies of insects during the past decade have reached a rough congruence in respect to several details [27][28][29][30][31][32] , such as the monophyly of Palaeoptera 27,30,33,34 , the ordinal relationships within Holometabola 27,[35][36][37] , and the recognition of monophyletic groups within Polyneoptera 30,32 . Although a recent study investigated the phylogeny of insects based on transcriptome data 27 , it still suffered from some unusual groupings, most notably the sister-group relationship between Psocodea and Holometabola, rendering Paraneoptera paraphyletic. ...
... Due to the improved taxon sampling and/or a better alignment of the molecular sequences, the results of different phylogenetic studies of insects during the past decade have reached a rough congruence in respect to several details [27][28][29][30][31][32] , such as the monophyly of Palaeoptera 27,30,33,34 , the ordinal relationships within Holometabola 27,[35][36][37] , and the recognition of monophyletic groups within Polyneoptera 30,32 . Although a recent study investigated the phylogeny of insects based on transcriptome data 27 , it still suffered from some unusual groupings, most notably the sister-group relationship between Psocodea and Holometabola, rendering Paraneoptera paraphyletic. ...
... In the phylogenetic results obtained from maximum parsimony (MP) analyses many clades were also supported by high or moderate bootstrap values. As a summary, our phylogenetic results show a high congruence with several previous works involving various kinds of evidence 27,[29][30][31][32][34][35][36]38,39 . This arguably provides a strong basis for an estimation of divergence times of the deeper nodes of insects. ...
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Insecta s. str. (=Ectognatha), comprise the largest and most diversified group of living organisms, accounting for roughly half of the biodiversity on Earth. Understanding insect relationships and the specific time intervals for their episodes of radiation and extinction are critical to any comprehensive perspective on evolutionary events. Although some deeper nodes have been resolved congruently, the complete evolution of insects has remained obscure due to the lack of direct fossil evidence. Besides, various evolutionary phases of insects and the corresponding driving forces of diversification remain to be recognized. In this study, a comprehensive sample of all insect orders was used to reconstruct their phylogenetic relationships and estimate deep divergences. The phylogenetic relationships of insect orders were congruently recovered by Bayesian inference and maximum likelihood analyses. A complete timescale of divergences based on an uncorrelated log-normal relaxed clock model was established among all lineages of winged insects. The inferred timescale for various nodes are congruent with major historical events including the increase of atmospheric oxygen in the Late Silurian and earliest Devonian, the radiation of vascular plants in the Devonian, and with the available fossil record of the stem groups to various insect lineages in the Devonian and Carboniferous.
... As addressed above, both morphological characters of wing base structure 21 and mitochondrial genome data 11 supported Zoraptera as sister to Embioptera. However, nuclear genes supported a close relationship between Zoraptera and Dictyoptera 28,29 . A more recent study based on genome-scale data recovered a sister-group relation between Zoraptera and Dermaptera 12 . ...
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The Polyneoptera represents one of the earliest insect radiations, comprising the majority of hemimetabolous orders, in which many species have great economic importance. Here, we sequenced eleven mitochondrial genomes of the polyneopteran insects by using high throughput pooled sequencing technology, and presented a phylogenetic reconstruction for this group based on expanded mitochondrial genome data. Our analyses included 189 taxa, of which 139 species represent all the major polyneopteran lineages. Multiple results support the monophyly of Polyneoptera, the monophyly of Dictyoptera, and the monophyly of Orthoptera. Sister taxon relationships Plecoptera + Dermaptera, and Zoraptera + Embioptera are also supported by most analyses. Within Dictyoptera, the Blattodea is consistently retrieved as paraphyly due to the sister group relationship of Cryptocercus with Isoptera. In addition, the results demonstrate that model selection, data treatment, and outgroup choice can have significant effects on the reconstructed phylogenetic relationships of Polyneoptera.
... Zoraptera are a very small pantropical group of polyneopteran insects with a still unclarified systematic position . Recent studies based on molecular data have suggested that they are closest relatives of Embioptera, Dermaptera or Dictyoptera (Wheeler et al., 2001;Terry and Whiting, 2005;Yoshizawa and Johnson, 2005;Kjer et al., 2007;Ogden and Rosenberg, 2007;Yoshizawa, 2011;Ishiwata et al., 2011;Simon et al., 2012;Wang et al., 2013;Misof et al., 2014). The presently known 40 extant species are very uniform in their external morphology. ...
... Some are from Tian et al. (2011) and Li et al. (2012); some were newly designed as shown in Table 3. Sequences of 18S rDNA, 28S rDNA, Hox and 13 protein coding genes of mitochondrial genomes were aligned by Muscle, embedded within MEGA 6.06 (Tamura et al. 2013) under the default conditions of the program. The alignments of 18S rDNA and 28S rDNA were adjusted according to the secondary structure models of heteropteran 18S rRNA and 28S rRNA (Xie et al. 2009;Wang et al. 2013), and length-variable regions of both genes were eliminated before phylogenetic reconstructing . The six Hox genes were also manually corrected according Tian et al. (2011). ...
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Pentatomomorpha is one of the most biodiverse infraorders among the true bugs (Hemiptera: Heteroptera). Phylogenetic relationships among the superfamilies within this infraorder have been uncertain, especially for the Eutrichophora. The previous studies were based on morphological characters, or just mitochondrial or nuclear genes, or only partial 18S rDNA and COI. In this study, we used maximum likelihood (ML) and Bayesian inference (BI) based on massive molecular datasets (18S rDNA, 28S rDNA, Hox and mitochondrial genes totaling 21 loci and 12,538 characters) to infer a robust phylogeny for this terrestrial group. Results strongly support the monophyly of all superfamilies; the superfamily status of Aradoidea and the following relationships: (Aradoidea + (Pentatomoidea + (Coreoidea + (Lygaeoidea + Pyrrhocoroidea)))) in Pentatomomorpha, and (Coreoidea + (Lygaeoidea + Pyrrhocoroidea)) in Eutrichophora. Our results suggest that sampling greater numbers of genes is an effective tool for resolving phylogenetic problems.
... Construction of the secondary structure models of 18S and 28S rRNAs of Pentatomoidea referred to the universal model of 18S rRNA (Xie et al., 2009(Xie et al., , 2012 and 28S rRNA (Gillespie et al., 2006;Wang et al., 2013) of insects. The numbering system for LVRs in 18S rRNA followed those used by Neefs et al. (1993) and Xie et al. (2009), and that for LVRs in 28S rRNA followed those used by Hassouna, Michot & Bachellerie (1984) and Wang et al. (2013). ...
... Construction of the secondary structure models of 18S and 28S rRNAs of Pentatomoidea referred to the universal model of 18S rRNA (Xie et al., 2009(Xie et al., , 2012 and 28S rRNA (Gillespie et al., 2006;Wang et al., 2013) of insects. The numbering system for LVRs in 18S rRNA followed those used by Neefs et al. (1993) and Xie et al. (2009), and that for LVRs in 28S rRNA followed those used by Hassouna, Michot & Bachellerie (1984) and Wang et al. (2013). RNAstructure 5.4 based on the thermodynamic folding (Reuter & Mathews, 2010) was used to reconstruct the secondary structures of the regions containing LVRs. ...
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Pentatomoidea (stink bugs and their relatives) is the third largest superfamily in Heteroptera, or the true bugs. The phylogenetic relationships among the families within Pentatomoidea remain controversial. The family Lestoniidae is morphologically highly specialized, currently including only two species endemic to Australia. Previous researchers have suggested a close relationship of Lestoniidae to either Plataspidae or Acanthosomatidae, based on morphological characters. In this study, phylogenetic tree reconstruction revealed that Lestoniidae and Acanthosomatidae form a monophyletic clade. In addition, in comparisons of the secondary structures of 18S and 28S rRNAs representing 15 families of Pentatomoidea, four length-variable regions in 18S and 28S rRNAs that can serve as autapomorphies for the clade Lestoniidae + Acanthosomatidae were recognized. Among them, E in 18S rRNA and D3-1 and D5-1 in 28S rRNA are unique in length in Lestoniidae and Acanthosomatidae. Based on the new molecular evidence and morphological evidence published by previous authors, Lestoniidae is suggested to be a highly specialized group derived from a common ancestor with Acanthosomatidae.