18S rRNA secondary structure and phylogenetic position of Peloridiidae (Insecta, hemiptera)

A secondary structure model for 18S rRNA of peloridiids, relict insects with a present-day circumantarctic distribution, is constructed using comparative sequence analysis, thermodynamic folding, a consensus method using 18S rRNA models of other taxa, and support of helices based on compensatory substitutions. Results show that probable in vivo configuration of 18S rRNA is not predictable using current free-energy models to fold the entire molecule concurrently. This suggests that refinements in free-energy minimization algorithms are needed. Molecular phylogenetic datasets were created using 18S rRNA nucleotide alignments produced by CLUSTAL and rigorous interpretation of homologous position based on certain secondary substructures. Phylogenetic analysis of a hemipteran data matrix of 18S rDNA sequences placed peloridiids sister to Heteroptera. Resolution of affiliations between the three main euhemipteran lineages was unresolved. The peloridiid 18S RNA model presented here provides the most accurate template to date for aligning homologous nucleotides of hemipteran taxa. Using folded 18S rRNA to infer homology of character as morpho-molecular structures or nucleotides and scoring particular sites or substructures is discussed.     

The sperm structure of Embioptera (Insecta) and phylogenetic considerations

The sperm structure of two species of Embioptera, Embia savignyi Westwood 1837 and Aposthonia japonica (Okajima 1926), was studied. Spermatozoa of both species exhibit a monolayered acrosome and a layer of material surrounding the sperm cells for most of their length. The presence of a 9+9+2 axoneme provided with accessory microtubules with 16 protofilaments, two accessory bodies and two crystallized mitochondrial derivatives are characters shared with other polyneopteran taxa. The supposed close relationship between Embioptera and Phasmatodea is not supported by characters of the sperm ultrastructure.     

Tibial combs in the Cydnidae (Hemiptera: Heteroptera) and their functional, taxonomic and phylogenetic significance

Tibial combs in representatives of the family Cydnidae are described in detail for the first time. The structure was studied in 98 species of 58 genera representing all the subfamilies, among them 16 species were investigated using scanning electron microscopic (SEM) techniques. In addition, Parastrachia japonensis (Scott, 1880) of the family Parastrachiidae, and two species of Dismegistus Amyot and Serville, 1843 (a genus of uncertain systematic position within Pentatomoidea) have also been studied. Morphological terminology is proposed for all the structures connected with tibial combs and the term 'the tibial comb complex' is suggested; its functional, taxonomic and phylogenetic significance is also discussed. The genera of Cydnidae can be classified into two groups depending on differences in the tibial comb complex, when it is present; moreover, its absence in the tribe Scaptocorini (Cephalocteinae) is demonstrated for the first time and is regarded as an autapomorphy of this tribe. Data on the occurrence of tibial combs in other families of true bugs (Hemiptera: Heteroptera) are also briefly reviewed, and an anagenetic trend in their evolution in terrestrial Heteroptera is hypothesized. The presence of tibial combs on all legs is regarded as the most plesiomorphic state.     

Wing base structure supports Coleorrhyncha + Auchenorrhyncha (Insecta: Hemiptera)

The phylogenetic placement of the moss bugs (Insecta: Hemiptera: Coleorrhyncha) has been highly controversial. Many apparent morphological apomorphies support the close relationship between Coleorrhyncha and Heteroptera (=true bugs). However, a recent phylogenomic study strongly supported a sister group relationship between Coleorrhyncha and Auchenorrhyncha (planthoppers, leafhoppers, treehoppers, spittlebugs, and cicadas). To test these two alternative hypotheses, we examined the fore‐ and hindwing base structure of the only known extant macropterous species of Coleorrhyncha using binocular and confocal laser scanning microscopes and analyzed the data selected from the wing base phylogenetically. When full morphological data including the wing base characters were analyzed, the sister group relationship between Coleorrhyncha + Heteroptera was supported, agreeing with previous consensus based on morphology. In contrast, when only wing base characters were analyzed separately, the clade Coleorrhyncha + Auchenorrhyncha was recovered, in agreement with the result from the phylogenomic study. The membranous condition of the proximal median plate in the forewing was identified as a potential synapomorphy of the latter grouping, and the absence of the tegula was excluded as a potential synapomorphy of Coleorrhyncha and Heteroptera.     

The phylogenetic position of the oxymonad Saccinobaculus based on SSU rRNA

The oxymonads are a group of structurally complex anaerobic flagellates about which we know very little. They are found in association with complex microbial communities in the guts of animals. There are five recognized families of oxymonads; molecular data have been acquired for four of these. Here, we describe the first molecular data from the last remaining group, represented by Saccinobaculus, an organism that is found exclusively in the hindgut of the wood-eating cockroach Cryptocercus. We sequenced small subunit ribosomal RNA (SSU rRNA) from total gut DNA to describe Saccinobaculus SSU rRNA diversity. We also sequenced SSU rRNA from manually isolated cells of the two most abundant and readily identifiable species: the type species Saccinobaculus ambloaxostylus and the taxonomically contentious Saccinobaculus doroaxostylus. We inferred phylogenetic trees including all five known oxymonad subgroups in order to elucidate the internal phylogeny of this poorly studied group, to resolve some outstanding issues of the taxonomy and identification of certain Saccinobaculus species, and to investigate the evolution of character states within it. Our analysis recovered strong support for the existence of the five subgroups of oxymonads, and consistently united the subgroups containing Monocercomonoides and Streblomastix, but was unable to resolve any further higher-order branching patterns.     

New shore bug (Hemiptera, Heteroptera, Saldidae) from the Early Cretaceous of China with phylogenetic analyses

A new genus with a new species of Saldidae, Brevrimatus pulchalifergen. et sp. n., is described and illustrated. The fossil specimen was found from the Early Cretaceous Yixian Formation of Duolun County, Inner Mongolia, China. Phylogenetic analyses within Saldidae were performed, and the results indicate Brevrimatus pulchalifergen. et sp. n. should be assigned to the subfamily Chiloxanthinae.     

The rhabdom structure in the ommatidia of the Heteroptera (Insecta), and its phylogenetic significance

In its plesiomorphic state the insect ommatidium consists of eight retinula cells forming a fused rhabdom. It has long been observed that, in contrast to this pattern, Heteroptera have open rhabdoms. However, there has so far been no comprehensive and comparative study of heteropteran ommatidia. For this reason, we investigated the rhabdom structure in 36 species from all higher groups of Heteroptera, as well as from Coleorrhyncha and Auchenorrhyncha as outgroup representatives. In addition we surveyed the data of earlier authors, which brings the number of examined species to a total of more than 70. All examined Heteroptera do have open rhabdoms, with a system of six peripheral and two central rhabdomeres. Outgroup comparison shows that the open rhabdom is an autapomorphy of the Heteroptera. As for the rhabdom structure within the Heteroptera, we found further autapomorphic patterns in Corixidae (Nepomorpha), Gerromorpha, and Leptopodomorpha. Finally, the Cimicomorpha and Pentatomomorpha share a special pattern of the two central rhabdomeres, which we call V-pattern. This is a new synapomorphy of these two taxa. Accepted: 22 November 1999     

Barcoding bugs: DNA-based identification of the true bugs (Insecta: Hemiptera: Heteroptera)

Background

DNA barcoding, the analysis of sequence variation in the 5′ region of the mitochondrial cytochrome c oxidase I (COI) gene, has been shown to provide an efficient method for the identification of species in a wide range of animal taxa. In order to assess the effectiveness of barcodes in the discrimination of Heteroptera, we examined 344 species belonging to 178 genera, drawn from specimens in the Canadian National Collection of Insects.

Methodology/Principal Findings

Analysis of the COI gene revealed less than 2% intra-specific divergence in 90% of the taxa examined, while minimum interspecific distances exceeded 3% in 77% of congeneric species pairs. Instances where barcodes fail to distinguish species represented clusters of morphologically similar species, except one case of barcode identity between species in different genera. Several instances of deep intraspecific divergence were detected suggesting possible cryptic species.

Conclusions/Significance

Although this analysis encompasses 0.8% of the described global fauna, our results indicate that DNA barcodes will aid the identification of Heteroptera. This advance will be useful in pest management, regulatory and environmental applications and will also reveal species that require further taxonomic research.     

基于DNA条形码的中国普缘蝽属分类研究(半翅目:异翅亚目)

通过扩增中国普缘蝽属Plinachtus Stl(半翅目:异翅亚目:缘蝽科)已知的4个种:刺肩普缘蝽P.dissimilis Hsiao,1964,钝肩普缘蝽P.bicoloripes Scott,1874,黑普缘蝽P.acicularis(Fabricius,1803)和棕普缘蝽P.basalis(Westwood,1842)的COI(1338bp)序列,计算其种间/种内的遗传距离,并进行基于距离法、最大简约法和贝叶斯法的分析,结果均支持将刺肩普缘蝽和钝肩普缘蝽合并为1个种。     

Influence of data conflict and molecular phylogeny of major clades in Cimicomorphan true bugs (Insecta: Hemiptera: Heteroptera)

Cimicomorpha, which consists of 16 families representing more than 19,400 species, is the largest infraorder in Heteroptera, Insecta. We present the first molecular phylogenetic investigation of family relationships of Cimicomorpha, including 46 taxa from 12 of 16 Cimicomorphan families. Three genes, with a total of 3277 bp of sequence data (nuclear 18S rDNA: 2022 bp, 28S rDNA: 755 bp, and mitochondrial 16S rDNA: 498 bp) were analyzed. Data partitions were analyzed separately and in combination, by employing ML (maximum likelihood), MP (maximum parsimony), and Bayesian methods. As saturation was detected in substitutions of 16S rDNA, influence of data conflict in combined analyses was further explored by three methods: the incongruence length difference (ILD) test, the partitioned Bremer support (PBS), and the partition addition bootstrap alteration approach (PABA). PBS and PABA approaches suggested that 16S rDNA was not very suitable for addressing relationships at this level in Cimicomorpha. Our results also supported the nabid-cimicoid lineage for Cimicoidea proposed by Schuh and Stys [Schuh, R.T., Stys, P., 1991. Phylogenetic analysis of Cimicomorphan family relationships (Heteroptera). J. NY Entomol. Soc. 99 (3), 298-350]. Data incongruence and the utility of the three genes were briefly discussed.     

Ovary structure and transovarial transmission of endosymbiotic microorganisms in Marchalina hellenica (Insecta,Hemiptera, Coccomorpha: Marchalinidae)

Szklarzewicz, T., Kalandyk‐Kolodziejczyk, M., Kot, M. and Michalik, A. 2011. Ovary structure and transovarial transmission of endosymbiotic microorganisms in Marchalina hellenica (Insecta, Hemiptera, Coccomorpha: Marchalinidae). —Acta Zoologica (Stockholm) 00 :1–9. The paired ovaries of Marchalina hellenica are composed of about 200 ovarioles of telotrophic type. In each ovariole, a trophic chamber, vitellarium and ovariolar stalk can be distinguished. The tropharia comprise trophocytes and early previtellogenic oocytes (termed arrested oocytes) or trophocytes only. The arrested oocytes are not capable of further development. In the vitellaria, single oocytes develop that are connected to the tropharium by means of broad nutritive cords. The number of germ cells (trophocytes and oocytes) constituting ovarioles is not constant and may range between 25 and 32. Numerous endosymbiotic bacteria occur in the cytoplasm of trophocytes. The endosymbionts are transported via nutritive cords to the developing oocyte. The obtained results are discussed in a phylogenetic context.     

18S rRNA hyper-elongation and the phylogeny of Euhemiptera (Insecta: Hemiptera)

The small subunit of nuclear ribosomal RNA (SSU nrRNA), whose sedimentation is mostly 18S in eukaryotes, is considered a relatively conservative marker for resolving phylogenetic relationship at the order level or higher. Length variation in SSU nrDNA is common, and can be rather large in some groups. In studies of Hexapoda phylogeny, the SSU nrDNA has been repeatedly used as a marker. Sternorrhyncha has been rarely included. The lengths of SSU nrDNAs of sternorrhynchids, the basal group of Hemiptera identified in the previous study are 0.3-0.6 kb longer than the usual ones in Hexapoda (1.8-1.9 kb). To use the entire SSU nrDNA sequences or the length-variable parts could cause alignment trouble and therefore affect phylogenetic results, as shown in this study of Euhemiptera phylogeny. Two problems are particularly noticeable. One is that two hyper-variable regions flanking a short length-conservative region could become overlapped in the alignment. This will destroy the positional homology over a larger range. The other is that, when a base pair in a stem of the secondary structure is located near the length-variable regions (LVRs), the simultaneous positional homology of these two bases in the pair is always lost in the alignment results. In this study, the secondary structure model of Hexapoda SSU nrRNA was slightly adjusted and the LVR distributions in it were finely positioned. The noise caused by the hyper LVRs was eliminated and the simultaneous homology for the paired bases was recovered based on the secondary structure model. These corrections improved the quality of the data matrix and hence improved the resolving behavior of the algorithm used. This study provided more convincing evidence for resolving the Euhemiptera suborders phylogeny as (Archaeorrhyncha+(Clypeorrhyncha+(Coleorrhyncha+Heteroptera))). This result provided a more solid background for outgroup determination according to the phylogenetic studies inside each suborder. The problems caused by LVRs have seldom been well addressed. As phylogenetic reconstruction depends more on the data matrix itself than on the algorithm, and length variation of SSU/LSU rRNA exists more or less in any group, it is necessary to closely investigate the effect of rRNA length variation on alignment and phylogenetic reconstruction in more groups.     

The female postabdomen of the enigmatic Nannochoristidae (Insecta: Mecopterida) and its phylogenetic significance

Hünefeld, F. and Beutel, R.G. 2011. The female postabdomen of the enigmatic Nannochoristidae (Insecta: Mecopterida) and its phylogenetic significance. —Acta Zoologica (Stockholm) 00: 1–8. External and internal features of the female postabdomen of Nannochorista neotropica are described in detail. The conditions found in females of Nannochoristidae come closest to the ground plan of Mecopterida. This lineage is characterised by telescoping postabdominal segments, a presumptive autapomorphic feature that is modified in some antliophoran groups, but displayed by the nannochoristid species in a typical manner. More potential autapomorphies of Mecopterida, all present in Nannochoristidae, are the neo‐formation of an intersegmental muscle, a transverse muscle spanning between the genital appendages of segment VIII, a muscle connecting these appendages and the genital chamber and the loss of an intersegmental muscle. Plesiomorphic features of Nannochoristidae are the presence of paired genital appendages on segments VIII and IX. Information on the egg‐depositing substrates of the females is not available. The telescoping postabdomen is suitable for oviposition in soft substrates such as moist soil, or rotten plant materials in the riparian zone, and this is possibly a ground‐plan feature of Mecopterida. The results of recent phylogenetic analyses based on morphological data support a placement of Nannochoristidae in Antliophora, whereas the exact position of the group remains ambiguous. No characters of the female postabdomen were found supporting the monophyly of Mecoptera as conventionally circumscribed, that is Nannochoristidae + Boreidae + Pistillifera.     

大黑毛肩长蝽(半翅目:异翅亚目:地长蝽科)线粒体基因组及地长蝽科系统发育地位探讨

为了更加深入地了解地长蝽科的基因组水平特征,测序并分析了大黑毛肩长蝽Neolethaeus assamensis(半翅目:异翅亚目:地长蝽科:毛肩族)的线粒体基因组序列。大黑毛肩长蝽线粒体基因组是双链共价环状DNA分子,长度为17097 bp,编码13个蛋白质编码基因,22个t RNA基因和2个r RNA基因,基因排列方式同果蝇Drosophila yakuba一致。大黑毛肩长蝽线粒体基因组内存在2个大的非编码区。一个是控制区,另一个是位于ND6和Cyt B之间的串联重复区域,TRR4.4。控制区内包含7类共9个结构显著的区域,如一个茎环结构,3个非串联的重复序列以及其他5个结构区域。TRR4.4长802 bp,包括4个184 bp的重复单元和1个66 bp的部分重复单元。TRR4.4的重复单元与控制区中TRR2.7的重复单元在长度、方向以及核苷酸组成等方面几乎完全一致。22个t RNA全部能够折叠为典型的三叶草二级结构。16S r RNA二级结构包含6个结构域(结构域III在节肢动物中缺失)和44个茎环结构,12S r RNA二级结构包含3个结构域和28个茎环结构。基于蝽次目15个线粒体基因组数据分析得到的系统发育结果,支持地长蝽科位于长蝽总科基部分支的观点。     

Phylogenomic reconstruction illuminates the evolutionary history of freshwater to marine transition in the subfamily Haloveliinae (Hemiptera: Heteroptera: Veliidae)

Water crickets of the subfamily Haloveliinae are semi-aquatic bugs occurring in freshwater and marine habitats throughout the Indo-Australian region, presently including six genera with more than 80 extant species. Whether lineage diversification in Haloveliinae is associated with the utilization of new ecological niches caused by transition events between freshwater and marine habitats remains poorly understood. We investigate the evolutionary history of Haloveliinae using large-scale phylogenomic datasets and a set of novel redefined morphological characters based on 24 ingroup taxa representing all recognized genera. Our phylogenetic results based on the novel datasets definitively indicate that the freshwater genus Strongylovelia Esaki as currently defined is paraphyletic and supports the establishment of a new genus: Metavelia gen. nov., including three congeneric species: Metavelia patiooni comb. nov. (type species), Metavelia priori comb. nov. and Metavelia albicolli comb. nov. Reconstruction of ancestral habitats suggests a freshwater origin for the Haloveliinae. Divergence time estimations reveal that the origin of the monophyletic marine clade occurred at around 83 Ma (95% highest posterior density: 71–98 Ma) in the Late Cretaceous, involving a single transition event from freshwater to marine habitats. This time coincides with the period of high global sea levels in the Late Cretaceous. During this period, the marine incursions caused by the massive sea level rise flooded the continental margins, especially in Southeast Asia, where ancestral Haloveliinae were probably distributed. The appearance of new marine habitats after the marine incursions (e.g., intertidal, mangroves and estuarine) probably led to a subsequent establishment and diversification of the marine lineages.     

Phylogeny and diversification of the true water bugs (Insecta: Hemiptera: Heteroptera: Nepomorpha)

Climate fluctuations and tectonic reconfigurations associated with environmental changes play large roles in determining patterns of adaptation and diversification, but studies documenting how such drivers have shaped the evolutionary history and diversification dynamics of limnic organisms during the Mesozoic are scarce. Members of the heteropteran infraorder Nepomorpha, or aquatic bugs, are ideal for testing the effects of these determinants on their diversification pulses because most species are confined to aquatic environments during their entire life. The group has a relatively mature taxonomy and is well represented in the fossil record. We investigated the evolution of Nepomorpha based on phylogenetic analyses of morphological and molecular characters sampled from 115 taxa representing all 13 families and approximately 40% of recognized genera. Our results were largely congruent with the phylogenetic relationships inferred from morphology. A divergence dating analysis indicated that Nepomorpha began to diversify in the late Permian (approximately 263 Ma), and diversification analyses suggested that palaeoecological opportunities probably promoted lineage diversification in this group.     

Revisiting habitat and lifestyle transitions in Heteroptera (Insecta: Hemiptera): insights from a combined morphological and molecular phylogeny

Heteroptera, the true bugs, are part of the largest clade of non-holometabolous insects, the Hemiptera, and include > 42 000 described species in about 90 families. Despite progress in resolving phylogenetic relationships between and within infraorders since the first combined morphological and molecular analysis published in 1993 (29 taxa, 669 bp, 31 morphological characters), recent hypotheses have relied entirely on molecular data. Weakly supported nodes along the backbone of Heteroptera made these published phylogenies unsuitable for investigations into the evolution of habitats and lifestyles across true bugs. Here we present the first combined morphological and molecular analyses of Heteroptera since 1993, using 135 taxa in 60 families, 4018 aligned bp of ribosomal DNA and 81 morphological characters, and various analytical approaches. The sister-group relationship of the predominantly aquatic Nepomorpha with all remaining Heteroptera is supported in all analyses, and a clade formed by Enicocephalomorpha, Dipsocoromorpha and Gerromorpha in some. All analyses recover Leptopodomorpha + (Cimicomorpha + Pentatomomorpha), mostly with high support. Parsimony- and likelihood-based ancestral state reconstructions of habitats and lifestyles on the combined likelihood phylogeny provide new insights into the evolution of true bugs. The results indicate that aquatic and semi-aquatic true bugs invaded these habitats three times independently from terrestrial habitats in contrast to a recent hypothesis. They further suggest that the most recent common ancestor of Heteroptera was predacious, and that the two large predominantly phytophagous clades (Trichophora and Miroidea) are likely to have derived independently from predatory ancestors. We conclude that by combining morphological and molecular data and employing various analytical methods our analyses have converged on a relatively well-supported hypothesis of heteropteran infraordinal relationships that now requires further testing using phylogenomic and more extensive morphological datasets.