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Molecular phylogeny of the assassin bugs (Hemiptera: Reduviidae), based on mitochondrial and nuclear ribosomal genes
Abstract
The first comprehensive cladistic analysis of Reduviidae, the assassin bugs, based on molecular data is presented and discussed in the context of a recently-published morphological analysis. Assassin bugs are essential components of ecosystems, but also important in agriculture and medicine. Sampling included 94 taxa (89 Reduviidae, 5 outgroups) in 15 subfamilies and 24 tribes of Reduviidae and is based on ∼3300 base pairs of mitochondrial (16S) and nuclear (18S, 28SD2, 28SD3-5) ribosomal DNA. Partitions of the dataset were aligned using different algorithms implemented in MAFFT and the combined dataset was analyzed using parsimony, partitioned maximum likelihood and partitioned Bayesian criteria. Clades recovered in all analyses, independent of alignment and analytical method, comprise: Cimicomorpha and Reduviidae; Hammacerinae; Harpactorinae; Apiomerini; Peiratinae; Phymatinae; Salyavatinae; Triatominae; Phymatinae + Holoptilinae; the higher Reduviidae (Reduviidae excluding Hammacerinae and the Phymatine Complex); Ectrichodiinae + Tribelocephalinae; (Triatominae + Zelurus) + Stenopodainae. Hammacerinae are rejected as sister group to all remaining Reduviidae in all analyses, as is the monophyly of Reduviinae, Emesinae and Harpactorini. High support values for Triatominae imply that blood-feeding has evolved only once within Reduviidae. Stenopodainae and part of Reduviinae are discussed as close relatives to Triatominae.
... The weak point of these analyses is the poorly sample used for the Saicinae (Hwang & Weirauch, 2012;Weirauch, 2008;Weirauch & Munro, 2009). In addition, recent studies within the allied subfamily Emesinae showed that the genital characters could offer information to elucidate the relationships and limits to tribal, generic, and specific levels (Castro-Huertas et al., 2018. ...
... We can summarize the main findings of our research as follows: (1) Subfamily Saicinae, as considered nowadays, is not monophyletic, and the genera Saicireta and Oncerotrachelus should be removed from the subfamily and probably be considered as Emesinae; (2) (Villiers, 1951) or closely related to, but different (China & Miller, 1959); and in previous cladistic studies the position of Carayonia is variable: as sister-group of Oncerotrachelus conforming a clade related with Emesinae + Saicinae (Weirauch, 2008); as part of the Saicinae (Weirauch & Munro, 2009), and as sister-group of a clade form by Emesinae + Saicinae and other subfamilies (Hwang & Weirauch, 2012). ...
... evidence; suggesting that Saicinae might be paraphyletic due to the position of Oncerotrachelus(Hwang & Weirauch, 2012;Weirauch & Munro, 2009;Weirauch, 2008). Our results confirm this, and as currently defined Saicinae is polyphyletic, considering the position of the genera Oncerotrachelus and Saicireta. ...
Despite genital structures in insects being consistently important as systematic and taxonomy evidence, within assassin bugs (Heteroptera, Reduviidae) at least, the male and female genitalic structures of several subfamilies are poorly or totally unknown. The genital structure is mostly symmetric within Saicinae genera, but male genital asymmetry has been recorded in Gallobelgicus, Polytoxus, and recently Pseudosaica. The subfamily has been considered as closely related to Emesinae and Visayanocorinae but this hypothesis has never been tested using a comprehensive taxa sampling, being a constraint to test morphological traits changes or exploring characters' evolution hypotheses. Here, we compiled a morphological data set of 170 characters that includes external morphological characters and genitalia of both sexes of Saicinae which was analyzed cladistically including 55 terminals, comprising 16 genera (64% of the generic diversity), 43 species of Saicinae and 12 outgroups. Saicinae was recovered as polyphyletic, Saicireta correntina is recovered as sister-species of Empicoris armatus + Collartida (Emesinae), Oncerotrachelus, Carayonia (Visayanocorinae), and the Clade Saicinae sensu stricto. Carayonia orientalis is recovered as sister-species of Saicinae sensu stricto. Ancestral state reconstruction of symmetry of the male genitalia shows an ancestor with symmetric male genitalia, two independent emergences of asymmetrical male genitalia within Saicinae sensu stricto, and the asymmetrical endosomal sclerites appearing before the other asymmetric traits.
... Emesine, Saicinae, and Visayanocorinae have long been recognised as closely related taxa (Hwang & Weirauch, 2012;Weirauch, 2008;Weirauch & Munro, 2009;Wygodzinsky, 1966). Members of the Emesine Complex are recognised by the absence of several features, namely the ocelli, a well-developed corium on the forewing, dorsal abdominal scent-glands, and a fossula spongiosa on both the fore-and mid legs (Wygodzinsky, 1966). ...
... Table S1 provides the current classification for in-and outgroups, unique specimen identifier numbers, voucher depositories, and locality information. Vouchering procedures followed the guidelines laid out in Weirauch and Munro (2009). Specifically, voucher specimens were associated with unique specimen identifiers (USI labels) and databased using the Arthropod Easy Capture Specimen (AEC) database (https:// research.amnh.org/pbi/locality/). ...
... Protocols for extraction, amplification, PCR cleaning, and sequencing followed those described in Weirauch and Munro (2009), with the exception that occasionally abdomens were used for extraction when genomic DNA yield from a leg was too low. PCR products were cleaned using the Bio 101 Geneclean Kit or SureClean from Bioline. ...
Web‐building spiders are formidable predators, yet assassin bugs in the Emesine Complex (Hemiptera: Reduviidae: Emesinae, Saicinae, and Visayanocorinae) prey on spiders. The Emesine Complex comprises >1000 species and these web‐associated predatory strategies may have driven their diversification. However, lack of natural history data and a robust phylogenetic framework currently preclude tests of this hypothesis. We combine Sanger (207 taxa, 3865 bp) and high‐throughput sequencing data (15 taxa, 381 loci) to generate the first taxon‐ and data‐rich phylogeny for this group. We discover rampant paraphyly among subfamilies and tribes, necessitating revisions to the classification. We use ancestral character state reconstructions for 40 morphological characters to identify diagnostic features for a revised classification. Our new classification treats Saicinae Stål and Visayanocorinae Miller as junior synonyms of Emesinae Amyot and Serville, synonymizes the emesine tribes Ploiariolini Van Duzee and Metapterini Stål with Emesini Amyot and Serville, and recognises six tribes within Emesinae (Collartidini Wygodzinsky, Emesini, Leistarchini Stål, Oncerotrachelini trib.n. , Saicini Stål stat.n. , and Visayanocorini Miller stat.n. ). We show that a pretarsal structure putatively involved in web‐associated behaviours evolved in the last common ancestor of Emesini, the most species‐rich clade within Emesinae, suggesting that web‐associations could be widespread in Emesinae.
... With numerous efforts to investigate the phylogeny of Reduviidae during the past few decades, on the one hand some well-resolved phylogenetic results have been proposed [40][41][42][43][44][45][46], and on the other hand, there are still some conflicting evolutionary hypotheses in previous studies. Our phylogenetic analyses, based on concatenated mitochondrial and nuclear genes of 33 assassin bugs covering 17 subfamilies, achieved largely consistent topologies between the BI and ML methods ( Figure 2 and Figure S2). ...
... Our phylogenetic analyses, based on concatenated mitochondrial and nuclear genes of 33 assassin bugs covering 17 subfamilies, achieved largely consistent topologies between the BI and ML methods ( Figure 2 and Figure S2). The monophyletic Phymatine Complex sensu lato [44] is consistently recovered as the sister group to all the remaining reduviids ('Higher Reduviidae'), which was also supported by previous molecular analyses [41,44]. The well-supported clade consisting of Centrocnemidinae, Holoptilinae, and Phymatinae underpins the reduction of ridges and teeth on the outside of the mandibles, the elongated pygophore, the processes on the endosomal struts of the phallus, and ventral glands as the synapomorphies of Phymatine Complex sensu stricto [40]. ...
... Within the 'Higher Reduviidae', the basal phylogenetic position of Peiratinae is constant across our analyses with high confidence. The subsequent split of a clade comprising Emesine Complex and Ectrichodiinae with the synapomorphies of the mostly membranous forewings and the absence of hooks or teeth on the ridges of the mandibles is in accordance with previous molecular analyses [41,45]. Additionally, the monophyly of this clade and the sister relationship between Emesine Complex and Ectrichodiinae were also supported by earlier morphological cladistic analyses [40] and molecular analyses [41]. ...
Reduviidae, a hyper-diverse family, comprise 25 subfamilies with nearly 7000 species and include many natural enemies of crop pests and vectors of human disease. To date, 75 mitochondrial genomes (mitogenomes) of assassin bugs from only 11 subfamilies have been reported. The limited sampling of mitogenome at higher categories hinders a deep understanding of mitogenome evolution and reduviid phylogeny. In this study, the first mitogenomes of Holoptilinae (Ptilocnemus lemur) and Emesinae (Ischnobaenella hainana) were sequenced. Two novel gene orders were detected in the newly sequenced mitogenomes. Combined 421 heteropteran mitogenomes, we identified 21 different gene orders and six gene rearrangement units located in three gene blocks. Comparative analyses of the diversity of gene order for each unit reveal that the tRNA gene cluster trnI-trnQ-trnM is the hotspot of heteropteran gene rearrangement. Furthermore, combined analyses of the gene rearrangement richness of each unit and the whole mitogenome among heteropteran lineages confirm Reduviidae as a ‘hot-spot group’ of gene rearrangement in Heteroptera. The phylogenetic analyses corroborate the current view of phylogenetic relationships between basal groups of Reduviidae with high support values. Our study provides deeper insights into the evolution of mitochondrial gene arrangement in Heteroptera and the early divergence of reduviids.
... DNA of 22 bed bug specimens was subjected to standard PCR and sequencing using a thermal cycler automaton (Applied Biosystems, Foster City, CA, USA) using the primers targeting the 18S rDNA gene (Weirauch and Munro, 2009) to confirm morphological identification. DNA from C. hemipterus specimens reared in the laboratory was used as positive control. ...
... This technique was thus used unambiguously in this study to confirm the identification of species of C. hemipterus (GenBank: MN056507) (Benkacimi et al., 2020). The molecular data analysis was based on the 18S rDNA gene, which is usually used for the molecular identification of hemipterans (Weirauch and Munro, 2009). ...
... Despite being one of the largest superfamilies of true bugs and drawing the attention of medical entomologists, the evolutionary history of Reduvioidea is understudied. Phylogenetic hypotheses across this lineage include a morphological study (Weirauch 2008), Sangersequencing-based analyses (Weirauch and Munro 2009;Hwang and Weirauch 2012), as well as a phylogenomic study with limited taxon sampling (Zhang, Gordon, et al. 2016). Only the analysis of Hwang and Weirauch (2012) sampled Reduviidae relatively comprehensively and recovered many subfamilies with high support, but largely failed to resolve intersubfamilial relationships. ...
... Of the three subfamilies missing from our analyses, one is thought to be part of the Phymatinecomplex based on morphological data [Elasmodeminae (Carayon et al. 1958)], another rendered Salyavatinae paraphyletic in Sanger-based analyses [Sphaeridopinae ], and the last, Manangocorinae are monotypic, only known from the holotype, and morphology suggests that this subfamily may belong to clade N7. Topologies recovered from different data sets in our analyses are similar to but not identical with published hypotheses that attempted to resolve relationships across reduviid subfamilies (Weirauch and Munro 2009;Hwang and Weirauch 2012; and to topologies from studies that aimed to uncovering relationships within subfamilies or clades of related subfamilies [Harpactorinae/Bactrodinae ; Phymatine-complex (Masonick et al. 2017); Triatominae and relatives (Justi et al. 2014;Kieran et al. 2021); Ectrichodiinae ]. While several of the nodes along the backbone of Higher Reduviidae (N4) are unstable across our analyses, many aspects of our hypotheses are well -supported in the NT, AA, Ast, AU, and X12 analyses and can form the basis for a revised subfamily-and tribal-level classification built on diagnosable monophyletic groups. ...
Though the phylogenetic signal of loci on sex chromosomes can differ from those on autosomes, chromosomal-level genome assemblies for non-vertebrates are still relatively scarce and conservation of chromosomal gene content across deep phylogenetic scales has therefore remained largely unexplored. We here assemble a uniquely large and diverse set of samples (17 Anchored Hybrid Enrichment [AHE], 24 RNA-Seq, and 70 whole-genome sequencing [WGS] samples of variable depth) for the medically important assassin bugs (Reduvioidea). We assess the performance of genes based on multiple features (e.g., nucleotide vs. amino acid, nuclear vs. mitochondrial, and autosomal vs. X chromosomal) and employ different methods (concatenation and coalescence analyses) to reconstruct the unresolved phylogeny of this diverse (∼7,000 spp.) and old (>180 MYA) group. Our results show that genes on the X chromosome are more likely to have discordant phylogenies than those on autosomes. We find that the X chromosome conflict is driven by high gene substitution rates that impact accuracy of phylogenetic inference. However, gene tree clustering showed strong conflict even after discounting variable third codon positions. Alternative topologies were not particularly enriched for sex chromosome loci, but spread across the genome. We conclude that binning genes to autosomal or sex chromosomes may result in a more accurate picture of the complex evolutionary history of a clade.
... Despite being one of the largest superfamilies of true bugs and drawing the attention of medical entomologists, the evolutionary history of Reduvioidea is understudied. Phylogenetic hypotheses across this lineage include a morphological study (Weirauch 2008), Sangersequencing-based analyses (Weirauch and Munro 2009;Hwang and Weirauch 2012), as well as a phylogenomic study with limited taxon sampling (Zhang, Gordon, et al. 2016). Only the analysis of Hwang and Weirauch (2012) sampled Reduviidae relatively comprehensively and recovered many subfamilies with high support, but largely failed to resolve intersubfamilial relationships. ...
... Of the three subfamilies missing from our analyses, one is thought to be part of the Phymatinecomplex based on morphological data [Elasmodeminae (Carayon et al. 1958)], another rendered Salyavatinae paraphyletic in Sanger-based analyses [Sphaeridopinae ], and the last, Manangocorinae are monotypic, only known from the holotype, and morphology suggests that this subfamily may belong to clade N7. Topologies recovered from different data sets in our analyses are similar to but not identical with published hypotheses that attempted to resolve relationships across reduviid subfamilies (Weirauch and Munro 2009;Hwang and Weirauch 2012; and to topologies from studies that aimed to uncovering relationships within subfamilies or clades of related subfamilies [Harpactorinae/Bactrodinae ; Phymatine-complex (Masonick et al. 2017); Triatominae and relatives (Justi et al. 2014;Kieran et al. 2021); Ectrichodiinae ]. While several of the nodes along the backbone of Higher Reduviidae (N4) are unstable across our analyses, many aspects of our hypotheses are well -supported in the NT, AA, Ast, AU, and X12 analyses and can form the basis for a revised subfamily-and tribal-level classification built on diagnosable monophyletic groups. ...
Though the phylogenetic signal of loci on sex chromosomes can differ from those on autosomes, chromosomal-level genome assemblies for non-vertebrates are still relatively scarce and conservation of chromosomal gene content across deep phylogenetic scales has therefore remained largely unexplored. We here assemble a uniquely large and diverse set of samples (17 Anchored Hybrid Enrichment [AHE], 24 RNA-Seq, and 70 whole-genome sequencing [WGS] samples of variable depth) for the medically important assassin bugs (Reduvioidea). We assess the performance of genes based on multiple features (e.g., nucleotide vs. amino acid, nuclear vs. mitochondrial, and autosomal vs. X chromosomal) and employ different methods (concatenation and coalescence analyses) to reconstruct the unresolved phylogeny of this diverse (~7,000 spp.) and old (>180 MYA) group. Our results show that genes on the X chromosome are more likely to have discordant phylogenies than those on autosomes. We find that the X chromosome conflict is driven by high gene substitution rates that impact accuracy of phylogenetic inference. However, gene tree clustering showed strong conflict even after discounting variable third codon positions. Alternative topologies were not particularly enriched for sex chromosome loci, but spread across the genome. We conclude that binning genes to autosomal or sex chromosomes may result in a more accurate picture of the complex evolutionary history of a clade.
... It is shown that the diversification of Higher Reduviidae began at 97 Ma (81-113 Ma) in the Late Cretaceous, coinciding with two global changes in the timing of angiosperm and phytophagous insect radiation. The analysis also showed that all lineages of Reduviinae are located in the branches of Higher Reduviidae, and the same results were shown in the cladistic analysis of Reduviidae based on morphological characters (Weirauch, 2008), the molecular phylogeny within the assassin bugs obtained from mitochondrial and nuclear ribosomal genes (Weirauch and Munro, 2009), and in the phylogenetic analysis based on transcriptomic and ribosomal DNA data (Zhang et al., 2016). The fossils in this study are geologically dated to the Early Cretaceous, and there is ample evidence to support their classification as Reduviinae, which is the earliest known Reduviinae fossil in the world. ...
... This may be caused by changes in the predatory behavior of Reduviidae due to changes in the predator (Haridass and Ananthakrishnan, 1980;Weirauch and Cassis, 2006;Zhang and Weirauch, 2014;Wang and Liang, 2015;Castro-Huertas et al., 2019). The most primitive types of fossula spongiosa and stridulitrum in Reduviidae are found in the fossils of Reduviinae in this study, which further indicates that Reduviinae is probably the basal taxon in the Higher Reduviidae, rather than the branching taxon suggested in previous studies (Weirauch, 2008;Weirauch and Munro, 2009;Hwang and Weirauch, 2012;Zhang et al., 2016). The results of Weirauch (2008) based on morphological characters showed that Harpactorinae that lack fossula spongiosa in most species as a basal taxon within the Higher Reduviidae, which may need to be reconsidered. ...
This study describes a new genus Simplicivenius gen. nov. of Reduviinae with two new species, Simplicivenius tuberculosus sp. nov. and Simplicivenius rectidorsius sp. nov. from the Yixian Formation in northeastern China. The diagnosis of the genus includes the width of the scutellum about 2/3 of the width of the pronotum and the absence of longitudinal veins around the exterior cell of the membrane. This is the oldest fossil record of Reduviinae and indeed Reduviidae at present, dating back to the Early Cretaceous. This implies that Reduviinae existed before the Late Cretaceous biological mass extinction, hinting that Reduviinae may be a basal taxon within the Higher Reduviidae. Moreover, the fossils prove the correctness of the previous speculations on the ancestral state of the stridulitrum and the fossula spongiosa of the assassin bugs. The fossula spongiosa on the mid leg and fore leg are subequal in size and both occupy 1/3 of the tibia is considered the primitive type of fossula spongiosa.
... Triatominae is unusual within Reduviidae because all of its members are defined by their blood-sucking habit and show morphological adaptations associated with hostfinding and feeding on vertebrate blood (Otálora-Luna et al. 2015). Since triatomines probably evolved from a predator ancestor, feeding on insects can be considered a primitive feature and the ancient triatomines probably fed on both insect and vertebrates (Weirauch and Munro 2009). This hypothesis could explain why some triatomine species remain able to feed on other invertebrates (Sandoval et al. 2004(Sandoval et al. , 2010. ...
... Pinto, 1926 and also including several Reduviidae species. On the other hand, Weirauch (2008) using morphological traits of 21 subfamilies of Reduviidae supported the hypothesis of the subfamily Triatominae to be monophyletic, results which was corroborated by a molecular phylogeny of Reduviidae based on mitochondrial and nuclear ribosomal genes (Weirauch and Munro 2009). Hwang and Weirauch (2012), in a more extensive molecular analysis, reconstructed a phylogeny in which they observed the predatory reduviine genera Opisthacidius Berg, 1879 and Zelurus Hahn, 1826 to be closely related to Triatominae, recovered as paraphyletic. ...
The members of the subfamily Triatominae are true bugs specialized in blood-sucking. All species are potential vectors of Trypanosoma cruzi (Chagas, 1909), the causative agent of Chagas disease, although relatively few have epidemiological significance as vectors of the infection to the humans. The incidence of Chagas disease is declining, after the successful vector control campaign through Americas, but, the disease remains as a major problem to public health in Latin America. A few species of triatomines are found also in Asia and Oceania where the vector-borne transmission of T. cruzi does not occur. After Carlos Chagas discovered their importance as vectors, triatomine bugs have attracted permanent attention, and, thus, several aspects of their systematics, biology, ecology, biogeography, and evolution have been studied. Since the first species description, at the end of the eighteenth century, until the current approximately 150 extant species considered as valid, their classification remains mainly based in the traditional morphology. However, modern and diversified methods applied to studies of their characteristics, such as molecular approaches, are improving the systematics of these vectors. In the present chapter, the author summarizes the current knowledge on the taxonomy of the Triatominae.
... Unsurprisingly for a group of this size and complexity, the current classification of Reduviidae does not reflect evolutionary relationships within the group. Although the majority of subfamilies are well-supported as monophyletic groups, some are not (Hwang and Weirauch 2012;Weirauch and Munro 2009;Zhang et al. 2016a, b). The most dramatic example of a polyphyletic subfamily is Reduviinae-recognized exclusively by the absence of features present in the other subfamilies-that have been recovered in 11-13 different lineages in recent analyses (Hwang and Weirauch 2012). ...
... The triatomine sister taxon was represented by Opisthacidius Berg and Centrogonus Bergroth, two large-bodied reduviine taxa from the Neotropical and Australian regions, respectively, with Zelurus Burmeister and Neivacoris Lent & Wygodzinsky (both Neotropical) being more distantly related. Most subsequent molecular phylogenies included species of Opisthacidius, Zelurus, and/or Stenopodainae (but not the other two genera, nor additional largebodied Reduviinae) and all have found typically high support for this clade sometimes referred to as the Zelurus clade of Reduviinae + Triatominae + Stenopodain ae (Georgieva et al. 2017;Hwang and Weirauch 2012;Justi et al. 2016;Patterson and Gaunt 2010;Weirauch and Munro 2009;Zhang et al. 2016a). In all analyses that included Opisthacidius, this genus forms either the sister taxon of Triatominae (Justi et al. 2016;Patterson and Gaunt 2010) (Figs. ...
Triatominae, the kissing bugs, are one of the smaller subfamilies of the otherwise predatory hemipteran family Reduviidae, the assassin bugs. Substantial progress has been made during the past decades to resolve phylogenetic relationships between Triatominae and other reduviid subfamilies as well as relationships among the ~150 species currently classified as kissing bugs. Nevertheless, many open questions remain. While Triatominae are now shown to be part of an assassin bug clade that also comprises Stenopodainae and part of Reduviinae, it is still not conclusively established if Triatominae are monophyletic or paraphyletic, and there are uncertainties with regard to relationships between and within tribes of Triatominae. This chapter summarizes available information on the evolution of Triatominae, highlighting strengths and shortcomings of currently published phylogenetic hypotheses. It stresses the importance of densely sampled, data-rich, robust phylogenies to inform the classification of Triatominae and to severe as a framework for evolutionary investigations across the group.
... Four cimicomorph lineages, Cimicidae (bed bugs), Polyctenidae (bat bugs), Reduviidae (assassin bugs), and extinct Torirostratidae, contain the hematophagous genera within Hemiptera. Given established phylogenies within the Cimicomorpha (Schuh et al. 2009;Weirauch and Munro 2009; also see Yao et al. 2014), hematophagous parasitism originated in the Hemiptera three times: once in Reduviidae, once in the Polyctenidae + Cimicidae clade, and once in the extinct Torirostratidae. ...
... Rhodnius prolixus (common kissing bug) and Triatoma infestans (winchuka) are major vectors of Chagas disease and frequently are associated with triggering anaphylaxis (Klotz et al. 2010). The Triatominae is a monophyletic subfamily within the Reduviidae (Weirauch and Munro 2009), although the Reduviidae is phylogenetically very distant from the Cimicidae + Polyctenidae clade within Heteroptera (Schuh et al. 2009). ...
Insect parasites and parasitoids are a major component of terrestrial food webs. For parasitoids, categorization is whether feeding activity is located inside or outside its host, if the host is immobilized or allowed to grow, and if the feeding is done by one or many conspecific or heterospecific individuals, and other features. Fossil evidence for parasitism and parasitoidism consists of taxonomic affiliation, morphology, gut contents, coprolites, tissue damage and trace fossils. Ten hemimetabolous and holometabolous orders of insects developed the parasite condition whereas seven orders of holometabolous insects evolved the parasitoid life habit. Modern terrestrial food webs are important for understanding the Mid Mesozoic Parasitoid Revolution. The MMPR began in late Early Jurassic (Phase 1), in which bottom-to-top regulation of terrestrial food webs dominated by inefficient clades of predators were replaced by top-to-bottom control by trophically more efficient parasitoid clades. The MMPR became consolidated in Phase 2 by the end of the Early Cretaceous. These clades later expanded (phases 3 and 4) as parasitoids became significant ecological elements in terrestrial food webs. Bottom-to-top food webs explained by the resource concentration hypothesis characterize pre-MMPR time. During phases 1 and 2 of MMPR (Middle Jurassic to Early Cretaceous), a shift ensued toward top-to-down food webs, explained by the trophic cascade hypothesis, exemplified by hymenopteran parasitoid clades Stephanoidea and Evanioidea. Clade-specific innovations spurring the MMPR included long, flexible ovipositors (wasps), host seeking, triungulin and planidium larvae (mantispids, beetles, twisted-wing parasites, flies), and extrudable, telescoped ovipositors (flies). After the MMPR, in phases 3 and 4 (Late Cretaceous to Recent), parasitoids increased in taxonomic diversity, becoming integrated into food webs that continue to the present day.
... Primer sequences and annealing temperatures are provided in Table 2. Primers for 18S rRNA and 28S rRNA correspond to (2012) for the 5 0 region of cytochrome c oxidase subunit I (COI hereafter) and primers from Damgaard and Sperling (2001) for the 3 0 region of COI. Primers for 16S rRNA were taken from Menard et al. (2014) and Weirauch and Munro (2009). The PCR for 16S rRNA with primers from Menard et al. (2014) had lower success rate and resulted in longer sequences, in comparison to primers from Weirauch and Munro (2009). ...
... Primers for 16S rRNA were taken from Menard et al. (2014) and Weirauch and Munro (2009). The PCR for 16S rRNA with primers from Menard et al. (2014) had lower success rate and resulted in longer sequences, in comparison to primers from Weirauch and Munro (2009). The annealing temperature for the 5 0 region of COI in most cases was 45 C. If the results were unsuccessful, the PCR was repeated with the annealing temperature at 42 C to obtain more sequences. ...
The Lygus-complex is one of the most taxonomically challenging groups of Miridae (Heteroptera), and its Australian fauna is poorly studied. Here we examine the Australian taxa of the Lygus-complex using morphological and molecular methods. After a detailed morphological study of the material collected throughout Australia, Taylorilygus nebulosus is transferred to Diomocoris, with the genus recorded for the first time in this country. Taylorilygus apicalis, also widely distributed in Australia, is redescribed on the basis of Australian material. The genus Micromimetus is recorded for the first time in Australia, with M. celiae, sp. nov., M. hannahae, sp. nov., M. nikolai, sp. nov. and M. shofneri, sp. nov. described as new to science. Micromimetus pictipes is redescribed and its distributional range is increased. The monophyly of the Lygus-complex and relationships within this group were tested using cytochrome c oxidase subunit I (COI), 16S rRNA, 18S rRNA and 28S rRNA markers. The Lygus-complex has been found to be non-monophyletic. Phylogeny confirmed the monophyly of Micromimetus, and it has shown that Taylorilygus apicalis is closer to Micromimetus species than to Diomocoris nebulosus. This study is the initial step in understanding the Lygus-complex phylogeny; analyses with more taxa, more genes and morphology are needed to reveal the interrelationships within this group, and sister-group relationships of Australian taxa. http://zoobank.org/urn:lsid:zoobank.org:pub:7393D96B-2BBA-438D-A134-D372EFE7FB9E
... Thus, taxonomic obscurities and confusions in species recognition have been caused by cases of the cryptic species complex (Panzera et al., 2015;Zhao et al., 2021), male-female dimorphism in a single species (Kwadjo et al., 2010;Gil-Santana, 2017;Weirauch et al., 2017;Chen et al., 2021), and remarkable morphological polymorphism or variation among conspecific populations or seasonal generations (Stål, 1867;Distant, 1903;Moreno et al., 2006;Rivas et al., 2021;Vilaseca et al., 2021). Moreover, recent studies using molecular phylogenetic analyses have provided phylogenetic hypotheses or presumptions that claim the necessity of reexaminations of the boundaries of many genera and subfamilies in the current classification of the family (Weirauch & Munro, 2009;Hwang & Weirauch, 2012). Therefore, future studies with more representatives of subfamilies and genera might recognize more issues of the current taxonomy of Reduviidae. ...
The genus Rhynocoris Hahn, 1834 is one of the largest genera of the subfamily Harpactorinae. This genus consists of 144 described species widely distributed in the Afrotropical, Palearctic, Sino-Japanese, Oriental, and Nearctic Realms. There are three species, Rhynocoris fuscipes (Fabricius, 1787), Rhynocoris marginellus (Fabricius, 1803), Rhynocoris mendicus (Stål, 1867) recorded in Vietnam. In this study, we attempted to re-examine the Rhynocoris species collected from Vietnam based on external morphology, genital morphology, and COI phylogeny to reveal the species delimitation of this genus in Vietnam. As a result, the independence of the three Rhynocoris species from Vietnam was confirmed. Moreover, R. mendicus was revealed as being a polymorphic species with two intraspecific morphological phenotypes.
... Phimophorinae with a precise subclade of Reduviidae. However, Davis (1961) mentioned that, based on wing venation, the Phimophorinae could be included in the Phymatine complex, one of the two clades supported by both molecular and morphological phylogenies of the Reduviidae (Weirauch, 2008;Weirauch & Munro, 2009), an opinion shared by Zhang et al. (2015), although not indicating any supporting characters. Such a hypothesis currently lacks robust phylogenetic argumentation and future analyses need to clarify the placement of these enigmatic assassin bugs. ...
The second fossil assassin bug of the cryptic subfamily Phimophorinae is described and figured from the mid-Miocene amber of the Dominican Republic, on the basis of traditional optical observations and a computed micro-tomography 3D reconstruction. The fossil can be attributed to the extant, monotypic genus Phimophorus Bergroth, which is currently known only from northern South America. The fossil extends the distribution of the lineage into the Caribbean of the Early Neogene and places it among a series of examples of arthropod groups that once thrived in Hispaniola but are today not native to the islands.
... Body sizes vary between 2 and 50 mm. They are more common in tropical and subtropical regions and form a group of approximately 6900 species belonging to 23 subfamilies (Weirauch and Munro, 2009;Putshkov and Moulet, 2009). The color of Reduviidae is mostly brown and brownish yellow, but some are red, yellow, orange and black. ...
... The phylogenetic inference was calculated with RAxML (model GTR1I1G4; bootstrap random number seed, 1,234; number of bootstraps, 100; random number seed for the parsimony inferences, 123) (66). For the samples that were morphologically identified as the genus Reduvius, we employed an alternative mitochondrial marker (16S rRNA gene amplified with 16sa and 16sb primers [62]) and followed the same approach as described above. Amplicon library preparation. ...
The importance of gut microbiomes has become generally recognized in vector biology. This study addresses microbiome signatures in North American Triatoma species of public health significance (vectors of Trypanosoma cruzi) linked to their blood-feeding strategy and the natural habitat. To place the Triatoma-associated microbiomes within a complex evolutionary and ecological context, we sampled sympatric Triatoma populations, related predatory reduviids, unrelated ticks, and environmental material from vertebrate nests where these arthropods reside. Along with five Triatoma species, we have characterized microbiomes of five reduviids (Stenolemoides arizonensis, Ploiaria hirticornis, Zelus longipes, and two Reduvius species), a single soft tick species, Ornithodoros turicata, and environmental microbiomes from selected sites in Arizona, Texas, Florida, and Georgia. The microbiomes of predatory reduviids lack a shared core microbiota. As in triatomines, microbiome dissimilarities among species correlate with dominance of a single bacterial taxon. These include Rickettsia, Lactobacillus, “Candidatus Midichloria,” and Zymobacter, which are often accompanied by known symbiotic genera, i.e., Wolbachia, “Candidatus Lariskella,” Asaia, Gilliamella, and Burkholderia. We have further identified a compositional convergence of the analyzed microbiomes in regard to the host phylogenetic distance in both blood-feeding and predatory reduviids. While the microbiomes of the two reduviid species from the Emesinae family reflect their close relationship, the microbiomes of all Triatoma species repeatedly form a distinct monophyletic cluster highlighting their phylosymbiosis. Furthermore, based on environmental microbiome profiles and blood meal analysis, we propose three epidemiologically relevant and mutually interrelated bacterial sources for Triatoma microbiomes, i.e., host abiotic environment, host skin microbiome, and pathogens circulating in host blood.
... Higher-level classification of Assassin bugs was settled by Amyot and Serville, who identified many subfamilies. Published phylogenies using morphological (Davis, 1957(Davis, , 1961Weirauch, 2008), molecular (Weirauch and Munro, 2009;Hwang and Weirauch, 2012) and combined datasets (Schuh et al., 2009) show high support for the monophyly of the family Reduviidae, with Pachynomidae as their sister group. ...
... The distal part of each bed bug specimen's abdomen was transferred in a 1.5 mL tube with 180 µL of G2 lysis buffer and 20 µL of proteinase K (Qiagen, Hilden, Germany) and incubated overnight at 56 • C. DNA extraction was performed using an EZ1 DNA Tissue Kit (Qiagen), according to the manufacturer's protocol, as previously described [7]. The DNA from 13 Cimex specimens was subjected to standard PCR and sequencing using a thermal cycler automaton (Applied Biosystems, Foster City, CA, USA) with primers targeting the 18S rDNA gene [17]. The DNA from laboratory-bred C. lectularius was used as a positive control. ...
Human infestations by bed bugs have upsurged globally in recent decades, including in African countries, where recent reports pointed out an increase in infestation. Sympatric dwelling has been described for two species of bed bug parasitizing humans: Cimex hemipterus (the tropical bed bug) and C. lectularius. Identification of these two species is based on morphological characteristics, and gene sequencing, and may also rely on Matrix-Assisted Laser Desorption Ionization Time-Of-Flight Mass Spectrometry (MALDI-TOF MS). The present work aimed to assess whether MALDI-TOF MS was applicable for species level identification of immature stages of Cimex. Arthropods were collected in domestic settings in Nouakchott, Mauritania. Identification used morphological keys and MALDI-TOF MS identification was assessed for immature stages. Quantitative PCR and sequencing assays were used to detect arthropod-associated bacteria in each specimen. A total of 92 arthropods were collected, all morphologically identified as C. hemipterus (32 males, 14 females and 45 immature stages). A total of 35/45 specimens produced good quality MALDI-TOF MS spectra. Analysis allowed species level identification of all immature C. hemipterus after their spectra were entered into our in-house MALDI-TOF MS arthropod spectra database. Molecular screening allowed detection of Wolbachia DNA in each specimen. These results suggested that MALDI-TOF MS is a reliable tool for species level identification of Cimex specimens, including immature specimens. Future studies should assess this approach on larger panels of immature specimens for different Cimex species and focus on the precise staging of their different immature developmental stages.
... This was the case for large and comparatively well studied families lacking widely accepted phylogenetic resolution e.g., Aradidae, Miridae, or Pentatomidae. However, in groups with a robust backbone phylogeny in place, e.g., Reduviidae (Weirauch 2008;Weirauch and Munro 2009;Hwang and Weirauch 2012), there was no doubt that the testis follicle number of seven is not only most common across the family, but may be treated as ancestral, and was coded as such. Within Veliidae, Halovelia septentrionalis Esaki, 1924 remains the only veliid species having two ovarioles p.o. Phylogeny of Gerromorpha by Damgaard (2008) rendered Haloveliinae as a sister group to Microveliinae which in turn forms a sister group to Gerridae. ...
The structure of testes and ovaries can be described in its simplest form by the number of follicles and ovarioles they contain. Sixty-five years after the last review of the internal reproductive systems in true bugs (Heteroptera), the data accumulated today on the number of testicular follicles and ovarioles in their gonads are summarized. In addition, data on the number and type (mesadenia/ectadenia) of accessory glands are given. The hemipteran suborder Heteroptera constitutes one of the most diverse groups of non-homometabolous (‘Hemimetabola’) insects, comprising more than 40,000 described species worldwide and approximately 100 families, classified into seven infraorders. Data are available for all infraorders; however, more than 90% of studied species belong to the largest and most evolutionarily derived infraorders Cimicomorpha and Pentatomomorpha. In true bugs, in general, the number of follicles varies from one to nine (in a testis), and the number of ovarioles varies from two to 24 (in an ovary). Seven follicles per testis and seven ovarioles per ovary prevail being found in approximately 43.5% (307 species) and 24.4% (367 species) of studied species, respectively. Such a structure of testes and ovaries is considered an ancestral character state in the Heteroptera. In the evolution of this group, the number of follicles and ovarioles both increased and decreased, but the trend towards a decrease clearly prevailed.
... Characterization of Triatominae was based on the amplification of two fragments of mitochondrial genes, the large ribosomal subunit 16S (LSU-rRNA) and cytochrome B (Cytb), and one fragment of a nuclear gene, the D2 variable region of the 28S RNA gene (28S-D2). The primer sequences used are described in Additional file 1: Table S1 [30,45,46], and the methodology is described in Additional file 1: Text S1b. The characterization of the blood meal source was achieved using the set of primers for the Cytb fragment (Additional file 1: Table S1) [47], using the methodology described in Additional file 1: Text S1c. ...
Background
Chagas disease, one of the most important neglected tropical diseases in the countries of Latin America, is considered to be a particularly important public health concern in the Amazon region due to increases in the number of outbreaks of acute Chagas disease and increased local transmission in the last 20 years. However, relative to other countries, in Bolivia there is little information available on its transmission in the Amazon region. The aim of this study was to investigate the infestation of palm trees, the main habitat of Triatominae in the region, in several localities, to evaluate the danger they represent to inhabitants.
Methods
Triatominae were collected using live bait traps left overnight in six localities in Pando and Beni Departments, Bolivia. DNA extraction and sequencing were used to establish the Triatominae species (Cytb, 16S and 28S-D2 gene fragments), and the blood meal sources (Cytb fragment). Trypanosoma sp. infection was analyzed by sequencing gene fragments (GPX, GPI, HMCOAR, LAP, PDH and COII) or by mini-exon multiplex PCR.
Results
A total of 325 Rhodnius were captured (97.3% of nymphs) from the 1200 traps placed in 238 palm trees and 32 burrows/ground holes. Sequence analyses on DNA extracted from 114 insects and phylogeny analysis identified two triatomine species: Rhodnius stali (17%) and Rhodnius montenegrensis (equated to Rhodnius robustus II, 83%). These were found in palm trees of the genera Attalea (69%), Astrocaryum (13%), Copernicia (12%), Euterpe (2%) and Acrocomia (1%). The infection rate was around 30% (165 analyzed insects), with 90% of analyzed insects infected by Trypanosoma cruzi (only the TcI discrete typing unit was detected), 3% infected by Trypanosoma rangeli (first time found in Bolivian Triatominae) and 7% infected by mixed T. cruzi (TcI)-T. rangeli. Rhodnius specimens fed on Didelphidae, rodents, gecko and humans.
Conclusions
The results of this study highlight the epidemiological importance of Rhodnius in the Bolivian Amazon region. The huge geographical distribution of Rhodnius and their proximity to the human dwellings, high infection rate and frequent meals on the human population highlight a risk of transmission of Chagas disease in the region.
Graphical Abstract
... Most of their species are generally found on the soil surface, under stones and debris and on low plants (Lodos & Önder, 1986). Reduviidae are more common in tropical and subtropical zones and constitute a group of nearly 6900 species divided in 23 clear cut subfamilies (Weirauch & Munro, 2009;Putshkov & Moulet, 2009). Most species are predators of insects or other arthropods, either specialized on certain groups of prey organisms or apparently relatively unspecialized, but Triatominae are haematophagous on vertebrates and are known as vectors of Chagas Disease in humans (Weirauch, 2008). ...
This study is based upon material of Anthocoridae, Miridae, Lygaeidae, Reduviidae and Tingidae collected from different localities in Iğdır between 2013 and 2018. In this study, five species belonged to four genera from family Anthocoridae, eight species of seven genera from family Miridae, 10 species of eight genera from family Lygaeidae, one species of one genus from family Reduviidae and four species of three genera from family Tingidae were recorded from Iğdır Province. In total, 28 species from 23 genera of families Anthocoridae, Miridae, Lygaeidae, Reduviidae and Tingidae were recorded from Iğdır. Among them, Lygus rugulipennis Poppius, 1911, Nagusta goedelii (Kolenati, 1857), Monosteira unicostata (Mulsant & Rey, 1852) and Stephanitis (Stephanitis) pyri (Fabricius, 1775) were found the most abundant and widespread species on fruit orchards in Iğdır Plain.
... According to our analyses using complete mitogenomes, the closest reduviid subfamily to Triatominae would be Stenopodainae species: Oncocephalus breviscutum and Canthesancus helluo. Previous phylogenetic comparisons using morphological characters and molecular sequences showed a close relationship between Triatominae and the Stenopodainae and Reduviinae (Zelurus clade) subfamilies [3,4,7,11,[34][35][36]. Unfortunately, no Zelurus mitogenome is available to be included in the present analysis. ...
The complete mitogenome of Triatoma boliviana Martínez, Chávez, Sossa, Aranda, Vargas and Vidaurre, 2007 was assembled using next generation sequencing data. The 16,719 bp long genome contains 13 protein coding genes, 22 transfer RNAs, two ribosomal RNAs, and a control region. This mitogenome showed similar nucleotide composition, gene order and orientation than other triatomines. Molecular phylogenetic analyses based on available complete mitogenomes from Reduviidae supported that Triatominae is a monophyletic group and that T. boliviana is basal to the two main Triatomini clades: North and South American. In addition, the analysis of a fragment of the 16S mitochondrial gene among Triatomini species, including species of the dispar lineage, supports the inclusion of T. boliviana in this group.
... However, as in the trees based on the PCGs and rRNA dataset, the monophyly of the Reduviinae subfamily was also not recovered using the tRNA dataset. The paraphyly/polyphyly of the Reduviinae subfamily has been discussed by several authors based on mitochondrial and nuclear genes (Weirauch and Munro 2009;Hwang and Weirauch 2012;Zhang et al. 2016;Zhao et al. 2019). The extensive sampling of Reduviinae by Hwang and Weirauch (2012) and Justi et al. (2016) generated a phylogenyinformed framework for the eventual re-classification of the members of this subfamily. ...
Chagas' disease is transmitted mainly by members of the subfamily Triatominae (Hemiptera: Reduviidae). Among them, Triatoma infestans (Klug) is the main vector of the disease in Southern Cone of Latin America. In order to contribute to knowledge of the genetic variation between triatomine vectors, in the present study, we analyzed the intraspecific and interspecific variations of the seven mitogenomes available from Triatominae. In addition, in order to examine their evolutionary relationships with others species of Reduviidae and to estimate the divergence time of the main lineages, we constructed phylogenetic trees including mitogenome sequences of 30 species from Reduviidae. Comparative analysis between mitochondrial DNA sequences from two specimens of T. infestans revealed a total of 54 variable sites. Triatoma infestans, Triatoma dimidiata (Latreille), Triatoma rubrofasciata (De Geer), Triatoma migrans (Breddin), Rhodnius pictipes (Stål), and Panstrongylus rufotuberculatus (Champion) present similar mitogenome organization and the length differences observed among these species are primarily caused by variations in control region (CR) and intergenic spacers (IGS). The relative synonymous codon usage values (RSCU) were similar in the six species of Triatominae, and in agreement with the observed in other insects, a biased use of A and C nucleotides in the majority strand was detected. The monophyly of five subfamilies was strongly supported (Phymatinae, Peiratinae, Triatominae, Stenopodainae, and Harpactorinae), while the sampled species of Reduviinae were grouped with one specie from the Salyavatinae subfamily. The oldest subfamily is Phymatinae at 100.3 Mya (99.6-102.2 Mya) and the youngest is Triatominae and Stenopodainae at 52.6 Mya (42.5-63.7 Mya). The estimated diversification time for the Triatominae subfamily agrees with the Andean uplift geological event. An analysis with more mitogenomes from more Triatominae species would be necessary to provide sufficient evidence to support this finding.
... DNA from 28 bed bug specimens were subjected to standard PCR and sequencing in an automated DNA thermal cycler (Applied Biosystems, Foster City, CA) using as primers the 18S rDNA gene (Weirauch et al. 2009) and mitochondrial cytochrome c oxidase (COI) subunit I (Folmer et al. 1994) amplifying fragments of 951 and 710 base pairs respectively (Table 1) to confirm the morphological identification, as previously described (Benkacimi et al. 2020). DNA extracted from C. lectularius specimens reared in the laboratory was used as a positive control. ...
Bed bugs are known to carry several microorganisms. The purpose of this study was to assess the prevalence of bed bug infestation in two rural areas of Senegal and determine the species present in the population. A screening was conducted to detect some arthropod associated pathogenic bacteria in bed bugs and to evaluate the prevalence of endosymbiont carriage. One survey took place in 17 villages in Niakhar and two surveys in Dielmo and Ndiop and surroundings area in the same 20 villages. Bed bugs collected were identified morphologically and by MALDI-TOF MS tools. Microorganisms screening was performed by qPCR and confirmed by sequencing. During the survey in the Niakhar region, only one household 1/255 (0.4%) in the village of Ngayokhem was found infested by bed bugs. In a monitoring survey of the surroundings of Dielmo and Ndiop area, high prevalence was found during the two rounds of surveys in 65/314 (21%) in 16/20 villages (January–March) and 93/351 (26%) in 19/20 villages (December). All bed bugs were morphologically identified as the species Cimex hemipterus, of which 285/1,637 (17%) were randomly selected for MALDI-TOF MS analysis and bacteria screening. Among the Bacteria tested only Wolbachia (Alphaproteobacteria, Rickettsiales, Rickettsiaceae) DNA was found in 248/276 (90%) of the bedbugs. We briefly describe a high level of non-generalized bed bug infestation in rural Senegal and the diversity of Wolbachia strains carried by C. hemipterus. This study opens perspectives for raising household awareness of bed bug infestations and possibilities for appropriate control.
... The phylogenetic relationship between Psammolestes and Rhodnius is T. Belintani et al. often discussed (Monteiro et al., 2002;Weirauch and Munro, 2009;Kieran et al., 2021). The data presented here show the proximity of Psammolestes to the two studied Rhodnius species. ...
The study of geometric morphometry has an impact on Triatominae studies. Currently, several taxonomic and systematic studies use this approach. The Triatominae subfamily comprises three fossil species and 154 extant species potentially capable of transmitting Trypanosoma cruzi, the causative agent of Chagas disease. This study aims to evaluate the external female genitalia of adult triatomines using multivariate geometric morphometric approaches, not only for validation but also for systematic inferences. Specimens belonging to the genera Panstrongylus, Psammolestes, Rhodnius, and Triatoma were evaluated, in addition to two species previously included in Triatoma: T. longipennis and T. phyllosoma. The results show that the external female genitalia have operational morphology and allow characterization of the species and the genera of the Triatominae. In addition, the multivariate technique enabled delimitation of the phylogenetic relationships of the subfamily, presenting results consistent with systematic studies. It can be concluded that the external female genitalia evaluated by geometric morphometry is a useful character for the taxonomy and systematics of Triatominae.
... The number of chromosomes characterized, until now, as the fundamental karyotype for the genus Zelurus (2n = 22) is the most observed karyotype and considered as an ancestor in Triatominae (Ueshima 1966;Alevi et al. 2015Alevi et al. , 2018. New karyotype analyses are important to assess the chromosomal relationship between Zelurus and triatomines, since Zelurus is recovered as a monophyletic group with Triatominae and Stenopodainae in the Zelurus clade (Weirauch & Munro 2009;Hwang & Weirauch 2012). Morphological analysis of the testes was proposed as a tool to evaluate evolutionary relationships among Heteroptera as there is great variation among the suborder (Gomes et al. 2013;Pereira et al. 2015). ...
Based on specimens of Zelurus ochripennis (Stål, 1854) collected in the entrance zone of a small terrestrial cave, some biological, ecological, morphological and cytogenetic data, as well, as the general morphology of their nymphs are presented for the first time. A short taxonomic summary of the species is also provided.
... We amplified one or two regions of 18S rDNA and two or three regions of 28S rDNA (see accession numbers in Table S1). All PCR reactions were done using EmeraldAmp GT PCR Master Mix (TaKaRa, Mountain View, CA, USA) using primers from Forero et al. (2013), Giribet et al. (1996), Menard et al. (2014) and Weirauch and Munro (2009). Each reaction contained 12.5 µL master mix, 9.5 µL water, 0.5 µL of each primer and 2 µL DNA template. ...
Currently comprising only about 430 species, Dipsocoromorpha or minute litter bugs are one of the small infraorders of Heteroptera. They are classified into five morphologically distinct families––Ceratocombidae, Dipsocoridae, Hypsipterygidae, Schizopteridae and Stemmocryptidae––but relationships among and within these families are poorly understood owing to the lack of phylogenetic studies. A phylogenetic hypothesis based on combined molecular and morphological data is important to both evaluate and revise the higher‐level classification and to explore the evolutionary history of morphological features including elytriform (or beetle‐like) forewings, that seem to have evolved multiple times in this group, jumping devices, and the highly modified and diverse male genitalia. We here use a dataset combining Sanger‐derived (~4500 bp; 108 taxa) and Illumina‐generated (~7500 bp; 24 taxa) sequence data with a morphological matrix (159 characters) and taxon sampling that comprises all currently recognized family‐, subfamily‐ and tribal‐level taxa and comprehensive genus‐level sampling to investigate phylogenetic relationships within litter bugs. Our results support the monophyly of Dipsocoromorpha, Schizopteridae and Dipsocoridae, whereas Ceratocombidae and the schizopterine subfamily “Ogeriinae” are polyphyletic and paraphyletic, respectively. A new classification is proposed that recognizes six families, including Trichotonannidae, stat.n., and two subfamilies each within the two larger families Ceratocombidae and Schizopteridae. Ancestral state reconstructions outline the complex evolutionary history of many morphological characters, including 15 independent origins of elytriform forewings, and at least five shifts in the degree of genitalic asymmetry.
... In contrast, P. horrida appears to use its PMG and AMG venoms defensively in a context-dependent manner. All three species belong to the subfamily Reduviinae, but the phylogenetic relationships between P. rhadamanthus, P. biguttatus, and P. horrida remain unclear because the subfamily is likely to be polyphyletic (Hwang & Weirauch, 2012;Weirauch & Munro, 2009). In our experiments, the PMG venom was secreted in response to mild harassment without restraint (allowing the bug to escape from serious confrontation), possibly representing the response to intraspecific conflict. ...
The Heteroptera are a diverse suborder of phytophagous, hematophagous, and zoophagous insects. The shift to zoophagy can be traced back to the transformation of salivary glands into venom glands, but the venom is used not only to kill and digest invertebrate prey but also as a defense strategy, mainly against vertebrates. In this study, we used an integrated transcriptomics and proteomics approach to compare the composition of venoms from the anterior main gland (AMG) and posterior main gland (PMG) of the reduviid bugs Platymeris biguttatus L. and Psytalla horrida Stål. In both species, the AMG and PMG secreted distinct protein mixtures with few interspecific differences. PMG venom consisted mostly of S1 proteases, redulysins, Ptu1‐like peptides, and uncharacterized proteins, whereas AMG venom contained hemolysins and cystatins. There was a remarkable difference in biological activity between the AMG and PMG venoms, with only PMG venom conferring digestive, neurotoxic, hemolytic, antibacterial, and cytotoxic effects. Proteomic analysis of venom samples revealed the context‐dependent use of AMG and PMG venom. Although both species secreted PMG venom alone to overwhelm their prey and facilitate digestion, the deployment of defensive venom was species‐dependent. P. biguttatus almost exclusively used PMG venom for defense, whereas P. horrida secreted PMG venom in response to mild harassment but AMG venom in response to more intense harassment. This intriguing context‐dependent use of defensive venom indicates that future research should focus on species‐dependent differences in venom composition and defense strategies among predatory Heteroptera.
... DNA amplification was carried out specifically for each target according to related publications (Porter and Collins, 1991;Folmer et al., 1994;Monteiro et al., 2003;Weirauch and Munro, 2009, Table 2). Polymerase chain reactions (PCR) were performed using Thermal Cycler T100TM (BIORAD©). ...
Chagas disease, whose etiological agent is the protozoan Trypanosoma cruzi, affects millions of people worldwide and it is mainly transmitted by infected triatomine feces. Triatoma is the most diverse genus and one of the most important from an epidemiological point of view. Species of this genus are grouped into eight complexes and nine subcomplexes. Triatoma sordida subcomplex consists of T. garciabesi, T. guasayana, T. patagonica, and, T. sordida. Given the recent discussions on their phylogenetic status, this study aims to evaluate morphometric and genetics characters that group and distinguish T. garciabesi, T. guasayana, T. patagonica, and T. sordida, as well as to infer the phylogenetic relationships within the group and evaluate the proximity with T. rubrovaria subcomplex. The results corroborate the phylogenetic relationship of T. guasayana and T. patagonica with the T. rubrovaria subcomplex. Molecular data confirm the proximity of T. jurbergi, T. matogrossensis, T. vandae as T. garciabesi, and T. sordida. Together, genetic variability was shown between T. sordida populations.
... Four were mitochondrial genes -cytochrome b (Cyt b), cytochrome c oxidase subunit 2 (COII), the large subunit ribosomal RNA (16S-rRNA), small subunit ribosomal RNA (12S-rRNA)-and two were nuclear genes e18S ribosomal RNA (18S-rRNA), internal transcribed spacer 2 (ITS2). Table S2 presents the sequence of the primers used and their origin (Barnabé et al., 2020;Gomez-Palacio et al., 2013;Justi et al., 2014;Kocher et al., 1989;Marcilla et al., 2001;Monteiro et al., 2013;Patterson and Gaunt, 2010;Weirauch and Munro, 2009). The standardized PCR procedure was as follows: buffer, 1×; dNTPs, 50 μM; primers, 0.2 μM each; Taq, (1 IU); 2 μl DNA solution (ca. ...
... We reconstructed an almost fully resolved phylogeny for the Triatomini, recovering various relationships that were obtained in previously published estimates of phylogenies of the group (Hypša et al., 2002;Dorn et al., 2009;Weirauch and Munro, 2009;Espinoza et al., 2013;Monteiro et al., 2013;Ibarra-Cerdeña et al., 2014;Dorn et al., 2016;Justi et al., 2016), as well as novel relationships. Our phylogenetic analyses with the amino acid data set also reconstructed topologies that were generally similar to those obtained with the nucleotide matrices. ...
Triatominae is a subfamily of blood-sucking reduviid hemipterans of public health importance primarily in tropical and sub-tropical regions of the Americas, whose members possess various morphological adaptations closely associated to hematophagy. Despite their medical importance, the systematics of the subfamily is far from resolved, particularly within the tribe Triatomini. Here we employed mitochondrial genome DNA sequences to reconstruct the phylogenetic relationships among 19 species of the North-Central American (NCA) clade of Triatomini and to estimate the times of origin and diversification of its main clades. Twenty-nine mitogenomes were examined for representative specimens of 25 species, including the outgroup. Phylogenetic informativeness estimated for each protein-coding gene showed that cox1, cox2 and atp6 were the most informative markers, whereas atp8 and nad4 had high saturation levels. Phylogenetic analyses excluding the latter two protein-coding genes recovered an almost fully resolved topology. The NCA clade apparently originated shortly after emergence of an initial land bridge of the Panama Isthmus, ca. 15.05–20.05 Mya. An Asian/pantropical subclade with Linshcosteus costalis, Triatoma rubrofasciata and T. migrans was nested within the NCA clade, from which it diverged ca. 12.42–17.3Mya. Uncorrected cox1 and 13 protein-coding gene distances suggest the existence of additional species within the dimidiata complex. In contrast, T. phyllosoma, T. mazzottii and T. longipennis, from the phyllosoma complex, have considerably low cox1 and 13 PCG distances among them, suggesting mitochondrial introgression or conspecificity. Our study yielded a robust phylogeny for the group, which could be tested with further phylogenetic hypotheses based on nuclear genome-wide markers.
... In studies in which specimens of the Cavernicolini and Borbodelini tribes were included, these tribes were grouped with the Rhodniini and not with the Triatomini, although all tribes preserved their monophyletic status. However, it should be noted that some of these studies only included one or two species of the Rhodniini tribe, given that their focus was not on the Rhodniini tribe but on the subfamily Triatominae and other Reduviidae [120][121][122][123][124][125]. The presence of two groupings within the pictipes group (Table 2) has led some authors to propose that the genus Rhodnius should be composed of clades or lineages. ...
The Triatominae subfamily includes 151 extant and three fossil species. Several species can transmit the protozoan parasite Trypanosoma cruzi, the causative agent of Chagas disease, significantly impacting public health in Latin American countries. The Triatominae can be classified into five tribes, of which the Rhodniini is very important because of its large vector capacity and wide geographical distribution. The Rhodniini tribe comprises 23 (without R. taquarussuensis) species and although several studies have addressed their taxonomy using morphological, morphometric, cytogenetic, and molecular techniques, their evolutionary relationships remain unclear, resulting in inconsistencies at the classification level. Conflicting hypotheses have been proposed regarding the origin, diversification, and identification of these species in Latin America, muddying our understanding of their dispersion and current geographic distribution. Clarifying these factors can help for the design of vector control strategies. The aim of this review is to depict the different approaches used for taxonomy of the Rhodniini and to shed light on their evolution and biogeography.
... More compelling support comes from molecular evidence. In studies focused on general subfamily-level relationships in Reduviidae, Weirauch & Munro (2009) and Hwang & Weirauch (2012), using a single species of Rhaphidosoma, found the group to be paraphyletic with respect to Harpactorini. In a phylogeny taxonomically focused on Harpactorinae, Zhang & Weirauch (2014) corroborated these results with three 'Rhaphidosomini' (although with only moderate support), with these taxa appearing sister to Coranus Curtis, 1833, a widely distributed Old World genus of morphologically antithetical (=short, robust) assassins. ...
Knowledge of the biodiversity of a region can have wide-ranging implications for other disciplines and enterprises, including public health, agriculture and pest management, and conservation. However, in order to properly assess a region’s biodiversity, an accurate and up-to-date taxonomy is needed.While description of new taxa plays a sizeable role in how biodiversity changes over time, the re-assessment of previously described taxa and the recognition of synonyms are of equal importance. In this service, taxonomic study of ‘Rhaphidosomatini’ and its variously associated taxa was undertaken. ‘Rhaphidosomatini’ are a group of harpactorine assassin bugs centred around three predominantly sub-Saharan genera. However, its composition has remained obfuscated, as nine other small (mostly monotypic) genera have been subsequently associated with the group, either explicitly or through implied relationship. Its basic history, culminating in the rejection of ‘Rhaphidosomatini’ as a valid taxonomic group, is here discussed. Also, during this investigation, both a new species and a new generic synonym were discovered, necessitating the following taxonomic changes. Hoffmannocoris sinuatus sp. nov. is herein described from Grahamstown, South Africa. Harrisocoris africanus Miller, 1959 is here considered a junior synonym of Igora repraesentatus Hesse, 1925 syn. nov.; this taxonomic change relegates the genus Harrisocoris Miller, 1959 to a junior synonym of Igora Hesse, 1925 syn. nov. Finally, a single new country record for Rhaphidosoma collarti Schouteden, 1952 from Uganda is reported.
... Primers for 18S and 28S rRNA correspond to those described by Menard et al. (2013); we have sequenced two parts of the 18S rRNA and four parts of 28S rRNA. We used primers from Jung & Lee (2012) and Menard et al. (2013) for COI and from Weirauch & Munro (2009) for 16S. The primer sequences and annealing temperatures are provided in Table 4. ...
Rhinomirini (Insecta: Heteroptera: Miridae: Cylapinae) is a heterogeneous tribe of uncertain systematic position. We provide a phylogenetic analysis of this tribe based on morphological characters and molecular markers. Monophyly of the tribe was not supported by our findings. The Rhinocylapus complex and Rhinomiriella complex are transferred to Fulviini. Rhinomirini are restricted to the Rhinomiris complex. The record of Rhinomirini in Australia is considered dubious. The genera Punctifulvius and Yamatofulvus are transferred to the Rhinocylapus complex. The Rhinocylapus complex is recorded from Australia for the first time, with four new species: Mycetocylapus alexeyi sp. nov., Punctifulvius austellus sp. nov., Punctifulvius aquilonius sp. nov. and Rhinocylapoides valentinae sp. nov. The Philippine species Mycetocylapus major is synonymized with Mycetocylapus minor. The Australian species Rhinomiriella tuberculata is redescribed, and females of this species are described for the first time. Rhinomiriella aidani sp. nov. is described as new to science. The female genitalia of Proamblia elongata, Rhinocylapidius velocipidoides and Rhinomiris conspersus and the male genitalia of Rhinomiris conspersus and Rhinomiridius aethiopicus are illustrated and discussed. Sexual dimorphism in the Rhinocylapus complex and wing modifications in Cylapinae are discussed.
... In addition to the coloration pattern, cuticular extensions of insects, such as hair, have been studied and classifications have been proposed for use as an identification tool, including for taxonomy. Despite several morphological studies in triatomine insects, knowledge regarding the distribution and morphological types of these structures in the subfamilies remains scarce and fragmented 7,14 . ...
Introduction:
Oral transmission of acute Chagas disease is an emerging public health concern. This study aimed to detect insect fragments in experimentally contaminated food, by comparing triatomines with other insects.
Methods:
Food samples were experimentally contaminated with insects, processed to recover their fragments by light filth, and analyzed by microscopy and Polymerase Chain Reaction (PCR).
Results:
Morphological differences between coleopteran and triatomine insects were observed in microscopic images. PCR was efficient in amplifying Triatominae DNA in the experimentally contaminated food.
Conclusions:
This methodology could be utilized by food analysts to identify possible insect contamination in food samples.
... Peiratines also are distinctive among assassin bugs in possessing asymmetrical external male genitalia (e.g., Villiers 1948, Schuh & Slater 1995, Coscarón 1997. The monophyly of this group has been supported in recent molecular phylogenies, although only a few peiratine species were included in each of the studies (e.g., Weirauch & Munro 2009, Hwang & Weirauch 2012, Zhang & Weirauch 2014. The subfamily contains several highly speciose genera (i.e., Ectomocoris Mayr, 1865: 99 spp.; Lestomerus Amyot & Audinet-Serville, 1843: 33 spp.; Peirates Audinet-Serville, 1831: 38 spp.; Sirthenea Spinola, 1837: 41 spp.), one of which, Sirthenea, is distributed throughout the New and Old World (e.g., Willemse 1985, Chłond & Bugaj-Nawrocka 2015, Chłond 2018, Chłond et al. 2019. ...
Herein are erected several new taxa and taxonomic changes based on a study of the Old World genera of Peiratinae: Sphodrembas gen. nov. is erected for a single new species, Sphodrembas fumipennis sp. nov. from Kenya and Tanzania. Ectomocoris caccabatus sp. nov. is newly described and represents the first record of the genus from the Korean Peninsula. Chaulioleistes gen. nov. is erected for Peirates singularis Stål, 1874, resulting in Chaulioleistes singularis (Stål, 1874) comb. nov. Isdegardes Distant, 1909, described in Acanthaspidinae (=Reduviinae), is considered a junior synonym of the peiratine genus Phalantus Stål, 1863 syn. nov., resulting in Phalantus melanocephalus (Distant, 1909) comb. nov. for the type species of Isdegardes. Lestomerus wroughtoni Bergroth, 1915 is transferred to Brachysandalus Stål, 1866, resulting in Brachysandalus wroughtoni (Bergroth, 1915) comb. nov. The previously proposed synonymy of Polychitonocoris formosus Miller, 1940 with Calistocoris caesareus Reuter, 1881 is corroborated. All of the above taxonomic acts are supplemented with a key to the species of Calistocoris, a key to the species of Pachysandalus, and a key to the peiratine genera of the Old World. Also reported are thirteen new country records for eleven other species: Brachysandalus bicolor (Villiers, 1948) (Kenya, Tanzania); Ectomocoris bimaculatus Schouteden, 1909 (Ghana); Ectomocoris fenestratus (Klug, 1830) (Ghana); Ectomocoris maculicrus (Fairmaire, 1858) (Uganda, Zimbabwe); Lestomerus affinis (Audinet-Serville, 1831) (Malaysia); Lestomerus basilewskyi Villiers, 1962 (Kenya); Lestomerus bicolor (Villiers, 1948) (Kenya); Lestomerus dubius Villiers, 1948 (Cameroon); Pachysandalus schoutedeni Villiers, 1962 (Sudan); Peirates collarti Schouteden, 1931 (Cameroon); Peirates quadrinotatus (Fabricius, 1798) (Malaysia); and Phalantus africanus Stål, 1874 (Ghana).
... The cycling conditions were: 94°C for 3 min; 94°C for 30 s; variable annealing temperature for 30 s, 72°C for 30 s, 35 cycles; 72°C for 7 min. Some primers had been previously designed Justi et al., 2014;Kocher et al., 1989;Marcilla et al., 2001;Monteiro et al., 2013;Weirauch and Munro, 2009) whereas others were designed by the researchers of this study, either on the basis of the two entire mitochondrial genomes deposited in GenBank (T. dimidiata, AF301594 and T. infestans KY640305) or on the basis of our first sequencing results (Table S2). ...
Limited genetic data are currently available for three vectors of Chagas disease in Ecuador, Panstrongylus howardi, P. chinai, and P. rufotuberculatus. Previously regarded as mainly sylvatic, these species have been poorly studied. Recently, they have been more frequently reported in domiciles and peridomiciles and are now considered true secondary vectors of Chagas disease in a country where an estimated 200,000 people are infected by Trypanosoma cruzi, a causative agent of this disease. In order to fill this gap, we obtained DNA for sequencing from 53 insects belonging to these three species and mainly sampled from the two Ecuadorian provinces of Loja and Manabí. We used six mitochondrial loci (COI, COII, ND4, CytB, 16S, and 12S) and two nuclear ones (ITS2 and 18S). We interpreted the phylogenetic trees built with single and concatenated data through maximum likelihood, Bayesian Markov chain Monte Carlo, and maximum parsimony methods. We provide evidence that P. chinai and P. howardi are indeed two supported species closely related and derived from a common ancestor. Additionally, the phylogenetic position of P. rufotuberculatus was confirmed as being distant from P. chinai and P. howardi and clustered with Triatoma dimidiata, a species belonging to the Northern American Triatoma clade.
... Ambrose [4] prepared the checklist 14 subfamilies with 144 genera and 464 species from Indian faunal limits. Mitochondrial genes are utilized to study the phylogeny [5] and phylogeography [6] and the revision of insect phylogenetics [7,8] particularly, Reduviidae [9,10] . Up to date, complete sequences of mitochondrial genes of assassin bugs viz., Brontostoma colossus from subfamily Ectrichodiinae, Oncocephalus breviscutum from subfamily Stenopodainae, Peirates arcuatus and Sirthenea flavipes from subfamily Peiratinae, Triatoma dimidiata (Latreille) from subfamily Triatominae, Valentia hoffmanni Stål from subfamily Salyavatinae, Agriosphodrus dohrni (Signoret) a representative of Harpactorinae were sequenced [8] . ...
The present study is on the phylogenetic analysis of mitochondrial COI gene of four species of Rhynocoris Kolenati, viz., Rhynocoris fuscipes (Fabricius), Rhynocoris kumarii Ambrose and Livingstone, Rhynocoris longifrons (Stål) and Rhynocoris marginatus (Fabricius), four ecotypes of R. kumarii viz., Kalthuruthy (RK-KAZ), Maruthuvazhmalai (RK-MAR), Muppandal (RK-MUP), Theni (RK-THE) and three morphs of R. marginatus viz. Niger (RM-NIG), Nigrosanguineous (RM-NSAN), and Niger banded morphs of R. marginatus (RM-NB). The nucleotide sequences of Rhynocoris species, ecotypes and morphs revealed intrageneric diagnostic characteristics as well as interspecific genetic variations among the four Rhynocoris and interecotypic and intermorphic peculiarities. The understating of such characteristics and variation in the phylogenetic reconstruction of family Reduviidae is discussed. Introduction Reduviidae is a diverse group of mostly predatory insects with currently close to 7000 species and subspecies in 913 genera and 25 subfamilies described worldwide [1]. Reduviids are abundant, occur worldwide and are voracious predators. Hence, they are referred to as "assassin bugs". After Maldonado's [1] world checklist on assassin bugs many Indian species have been described and redescribed and considerable changes have been incorporated at species, generic, tribe and subfamily levels [2, 3]. Ambrose [4] prepared the checklist 14 subfamilies with 144 genera and 464 species from Indian faunal limits. Mitochondrial genes are utilized to study the phylogeny [5] and phylogeography [6] and the revision of insect phylogenetics [7, 8] particularly, Reduviidae [9, 10]. Up to date, complete sequences of mitochondrial genes of assassin bugs viz., Brontostoma colossus from subfamily Ectrichodiinae, Oncocephalus breviscutum from subfamily Stenopodainae, Peirates arcuatus and Sirthenea flavipes from subfamily Peiratinae, Triatoma dimidiata (Latreille) from subfamily Triatominae, Valentia hoffmanni Stål from subfamily Salyavatinae, Agriosphodrus dohrni (Signoret) a representative of Harpactorinae were sequenced [8]. The most abundant harpactorine assassin bugs, Rhynocoris in Oriental region are the promising biological control agents [2, 11]. Since they occur as diverse morphs and ecotypes accurate identification of these biocontrol potential reduviids is imperative for their utilization in Insect Pest Management programmes. Ambrose et al. [12] analysed the phylogenetics of intrageneric and intraspecific variations of fifteen species of Rhynocoris and four ecotypes of R. kumarii and three morphs of R. marginatus of Indian and Non Indian origin and the role of geographical isolation on speciation using three genes viz., 16S, Cyt b and COI genes. Their analysis of 16S gene revealed the affinity between Indian species R. fuscipes and R. segmentarius of South Africa. The Cyt b revealed affinity between the non-Indian R. fuscipes with Indian R. fuscipes and the four Indian species of Rhynocoris having two affinity clusters between viz., R. kumarii and R.
... Most genera have only modest diversity (average number of species per genus ≈ 5.6), whereas only a few contain greater than 30 species (i.e., Cleptria Stål, 1855: 41 Formerly, the millipede assassins were the sole members of the subfamily Ectrichodiinae, yet recent studies have expanded the taxonomy of the group. Prior molecularly inferred phylogenies supported the monophyly of the millipede assassins, along with a close relationship to the triangle-headed bugs of the subfamily Tribelocephalinae (e.g., Weirauch & Munro 2009;Hwang & Weirauch 2012;Zhang et al. 2016). However, Forthman and Weirauch (2017) recently, in a more comprehensive analysis, demonstrated the polyphyly of Ectrichodiinae sensu stricto and the subsequent need to include the triangleheaded bugs within Ectrichodiinae, wherein they proposed a revised classification. ...
Malingeus tumidus gen. et sp. nov., a new millipede assassin from Queensland, and Yaramayahus rufescens gen. et sp. nov., a new millipede assassin from New South Wales, are herein described from the tribe Ectrichodiini (Ectrichodiinae). Additionally, Caloundranius formosus Miller, 1957, syn. nov. is here considered a junior synonym of Mendola puellula Breddin, 1900, which relegates the genus Caloundranius Miller, 1957, syn. nov. to a junior synonym of Mendola Breddin, 1900. The previously proposed status of Tasmanocoris evansi Miller, 1952, as a junior synonym of Nebriscus pupus Bergroth, 1895, is corroborated. Brisbanocoris fuscipennis Miller, 1957, syn. nov. is here considered a junior synonym of Lyramna vigil Breddin, 1900, which relegates the genus Brisbanocoris Miller, 1957, syn. nov. to a junior synonym of Lyramna Breddin, 1900. Finally, an updated generic key to the ectrichodiine genera of Australia is presented.
Predator-prey relationships are a driving force when examining community ecology. One question in predator-prey interactions is how a predator may respond to novel prey phenotypes, a form of frequency dependent prey selection. A predator may selectively target or avoid the novel prey, a behavior referred to as anti-apostatic or apostatic selection respectively. We examined apostatic prey selection by observing feeding behavior in two species of assassin bugs (Hemiptera: Reduviidae), the white-spotted assassin bug (Platymeris biguttatus) and the red-spotted assassin bug (Platymeris rhadamanthus) which were fed unmodified and modified (novel) phenotypes of Turkistan cockroaches, Blatta lateralis (Blattodea: Blattidae). The two species of assassin bugs represent either generalist (P. biguttatus) or specific feeders (P. rhadamanthus). We hypothesized that the generalist feeder would engage in anti-apostatic feeding while the specific feeder would engage in apostatic feeding. Our results indicated that there was no difference in feeding behavior between the two species, however an overall trend of apostatic feeding was observed in response to two of the four novel prey phenotypes. Factors beyond prey phenotype may also be influencing feeding choice in these insects. While the two predator species did not exhibit different feeding behaviors, the observation of apostatic feeding in this genus suggest an overall trend of utilizing specific prey images in feeding.
Urostylididae, a phytophagous heteropteran family that feeds on the sap of various trees and shrubs, comprises 8 genera and 173 species. Its phylogeny has received little attention, and no studies have revealed its generic monophyly or relationships. We present the first molecular phylogeny of Urostylididae based on complete mitogenomes and nuclear ribosomal genes from almost all genera and representative species, using maximum likelihood analysis and Bayesian inference. All phylogenetic results showed overall consistent topological relationships, indicating polyphyly of the three most speciose genera, Urolabida, Urochela, and Urostylis. Among the three monotypic genera, Chelurotropella formed a stable relationship with Urochela wui in all trees, Cobbenicoris was a stable sister group of Urostylis fici but with position variation among trees, and Urochellus formed a sister group with Urostylis cuneata or species of two or more genera in different trees. The smaller genus Tessaromerus was always sister to Urostylis tricarinata, but their positions varied among trees. Thus, the currently recognized genera of Urostylididae are unreliable. Furthermore, our phylogenetic results suggested some topological incongruence among the trees produced with different analytical methods and data sets, mainly among deep internal nodes, with short branches and low support values. Given the quartet‐based evaluation system and branch patterns, Urostylididae underwent rapid radiation resulting in incomplete lineage sorting and introgression in our data sets, making our phylogenetic analyses more sensitive to the data or method used. Moreover, the rapid radiation might have caused morphological homoplasy of diagnostic characters of genera, leading to taxonomic confusion for Urostylididae. Therefore, a thorough taxonomic revision of this family is needed.
In the present study, Collartida eowilsoni sp. nov. is described from Guadalcanal (Solomon Islands), which extends the distribution of the genus and the tribe Collartidini as a whole to the Oceanian biogeographic realm. The highly aberrant morphology of the new species required a reassessment of the diagnostic characters and generic limits of Collartida. To evaluate the systematic position of C. eowilsoni sp. nov. a morphology-based cladistic analysis was performed, for which homologies of cephalic armature were reinterpreted. Phylogenetic relationships of the new species to its congeners and among other members of the tribe are discussed.
Мақалада «Алтынемел» мемлекеттік ұлттық табиғи паркінің территориясындағы 2020-2021 жылдардағы зерттеу нәтижелері бойынша пайдалы жартылай қаттықанаттыларына фаунистикалық-экологиялық шолу жасалған. Зиянды насекомдар мен кенелердің санын табиғи реттеуде маңызы бар зоофагтар мен зоофитофагтар фаунасы берілген. Зерттеу нәтижесінде құрлық жартылай қаттықанаттыларының 4 тұқымдасының 67 түрі анықталды. Зерттеу нәтижелері бойынша Аңшы қандалалар тұқымдасы (Nabidae: 3 туыс - Prostemma (1 түр), Nabis (12 түр), Himacerus (1 түр) мен Ұсақ жыртқыштар тұқымдасынан (Аnthocoridae: 3 туыс - Anthocoris (6 түр), Temnostethus (1 түр), Orius (7 түр) – 14 түрден, Жыртқыштар тұқымдасынан (Reduviidae: 7 туыс - Empicoris (1 түр), Holotricius (1 түр), Reduvius (1 түр), Oncocephalus (1 түр), Coranus (3 түр), Rhynocoris (2 түр), Vachiria (1 түр) – 10 түр, Жай көзшесіздер тұқымдасынан (Miridae: 11 туыс - Deraeocoris (5 түр), Malacocoris (1 түр), Pilophorus (2 түр), Atractotomus (1 түр), Campylomma (2 түр), Myrmecoris (1 түр), Phytocoris (5 түр), Globiceps (2 түр), Psallus (7 түр), Cyllecoridea (1 түр), Blepharidopterus (2 түр) – 29 түр белгілі болды. Бұлардың ішінде қоректік байланысы жағынан 38 түр – зоофагтар (Nabidae, Аnthocoridae, Reduviidae тұқымдас өкілдері), ал 29 түр - зоофитофагтар (Miridae тұқымдас өкілдері) болып табылады. Жыртқыш қандалалар жаз бойы барлық жерлерде кездеседі, олар табиғаттағы әртүрлі буынаяқтылардың жұмыртқалары, дернәсілдері және ересек дараларымен қоректенеді. Демек, зиянды жәндіктермен және кенелермен қоректеніп, олардың санын биологиялық жолмен реттейді, сөйтіп табиғатқа көп пайда келтіреді.
The concept of the previously monogeneric subfamily Pseudocetherinae (Hemiptera: Heteroptera: Reduviidae) is revised and expanded. We here transfer Gerbelius Distant, 1903, Kayanocoris Miller, 1954, Microvarus Jeannel, 1917, Paragerbelius Miller, 1958, and Voconia Stål, 1866 from Reduviinae to Pseudocetherinae and treat Kayanocoris, Microvarus, Paragerbelius, and Pseudocethera Villiers, 1963 as junior synonyms of Voconia, resulting in new combinations for Voconia conradti (Jeannel, 1917) comb. nov., V. ifana (Villiers, 1963) comb. nov., V. monodi (Villiers, 1963) comb. nov., V. motoensis (Schouteden, 1929) comb. nov., V. ornata (Distant, 1903) comb. nov., V. schoutedeni (Villiers, 1964) comb. nov., V. typica (Miller, 1958) comb. nov., and V. wegneri (Miller, 1954) comb. nov. We also describe 23 new species of Voconia: V. bakeri sp. nov., V. bracata sp. nov., V. brachycephala sp. nov., V. chrysoptera sp. nov., V. coronata sp. nov., V. decorata sp. nov., V. dolichocephala sp. nov., V. fasciata sp. nov., V. grandioculata sp. nov., V. hemera sp. nov., V. isosceles sp. nov., V. laosensis sp. nov., V. lasiosoma sp. nov., V. lirophleps sp. nov., V. loki sp. nov., V. mexicana sp. nov., V. minima sp. nov., V. nyx sp. nov., V. smithae sp. nov., V. tridens sp. nov., V. trinidadensis sp. nov., V. tuberculata sp. nov., and V. vittata sp. nov. Lectotypes are designated for Gerbelius confluens Distant, 1903, G. typicus Distant, 1903, V. conradti comb. nov., V. ornata comb. nov., and V. pallidipes Stål, 1866. A revised diagnosis and description of Pseudocetherinae are provided along with photographs of the species and of the male genitalia of 13 pseudocetherine and five closely related reduviine species. An identification key to the two genera of Pseudocetherinae as well as a key to species of Voconia are presented. A phylogenetic hypothesis is proposed for the relationships of Pseudocetherinae using parsimony analyses of 77 morphological characters.
The Neotropical harpactorine assassin bug genus Acanthischium Amyot & Serville, 1843 (Hemiptera: Reduviidae: Harpactorini) includes four valid species: Acanthischium maculatum Amyot & Serville, 1843 (with ten color varieties proposed by Stål [1872]), Acanthischium haglundi Stål, 1866, Acanthischium superbum Haviland, 1931, and Acanthischium invium Elkins, 1961. A taxonomic revision and a morphological phylogenetic analysis were carried out to test species limits and to propose a hypothesis of evolutionary relationships. As a result of the phylogenetic analysis, Acanthischium is characterized on its external morphology by the elongate postocular region, posterior margin of the pronotum with spines, ventral surface of protrochanter with apical and basal protuberances, profemur wider than mesofemur, and protibia strongly curved. After the taxonomic revision we propose nine valid species, taking into consideration characters of external morphology, male and female genitalia, and color patterns: Acanthischium dimidiatum Stål, 1859 stat. rev.; Acanthischium flaviceps Stål, 1872 stat. rev.; Acanthischium haglundi Stål, 1866; Acanthischium kuna sp. nov. from Panama and Colombia; Acanthischium maculatum Amyot & Serville, 1843; Acanthischium meloae sp. nov. from Suriname; Acanthischium nigrum Stål, 1872 stat. rev.; Acanthischium semiflavum Stål,1872 stat. rev.; and Acanthischium superbum Haviland, 1931. We further synonymize A. invium Elkins, 1961 with A. superbum. All but one of the species-A. flaviceps-were included in the phylogenetic analysis, in which Acanthischium was found to be monophyletic, with the following set of relationships: (A. haglundi + (A. nigrum + (A. dimidiatum (A. maculatum, A. semiflavum, A. kuna, A. meloae, A. superbum)))). The resulting phylogenetic pattern indicate a body color transition from drab to more vivid ones, such as orange or red. The species exhibiting red and orange colorations are found in a clade, suggesting an underlying biological phenomenon, such as wasp mimetism.
Bedbugs (Cimex lectularius and C. hemipterus) have reemerged as a major public health problem around the world. Their bites cause various skin lesions as well as discomfort and anxiety. Their role as potential vectors of various infectious agents is discussed. Accordingly, all suspected cases of bedbug infestations need to be documented thoroughly, with an unequivocal identification of the arthropods involved, if any are present. Although morphological identification is easily and quickly performed by entomologists or professionals, it can be challenging otherwise. Also, distinguishing Cimex lectularius and C. hemipterus requires entomological expertise. MALDI-TOF mass spectrometry has been recently presented as an additional tool for arthropod identification. In this study, we assess the use of MALDI-TOF MS for the identification of laboratory and wild strains of C. lectularius and C. hemipterus. Several body parts of laboratory reared C. lectularius specimens were used to develop a MALDI-TOF MS protocol for bedbug identification, which was later validated using five other laboratory and wild populations of C. hemipterus and C. lectularius. A total of 167C. lectularius and C. hemipterus bedbug specimens (98 laboratory specimens and 69 wild specimens) were submitted to MALDI-TOF MS analysis. 143/167 (85.63%) provided high quality MS spectra. The in-lab database was then upgraded with a total of 20 reference spectra from all bedbug populations and the rest of the MS spectra (123 bedbugs) were blind tested. All specimens were properly identified to the species level using MALDI-TOF MS and 86,25% (69/80) were aptly identified according to their origin with LSVs ranging from 1.867 to 2.861. MALDI-TOF MS appears as a reliable additional tool for the identification of these two anthropophilic species.
A new plant bug genus and species, Andrewia bochkovi gen. et sp. nov., of the Phylinae is described from the Biak Island, Papua Province of Indonesia. The described species has been characterised molecularly based on COI, 16S and 28S markers, and its phylogenetic position has been tested using model‐based (RAxML and Bayesian) phylogenetic analyses. Photographs of habitus, male and female genitalia, illustrations of male genitalia, scanning electron micrographs of morphological structures and host information are also provided. The systematic position of the new genus within Phylinae is discussed. Andrewia gen. nov. forms a separate clade sister to all other phylines except for the tribe Hallodapini.
Chenicoris dilatatus gen. n. & sp. n. from southern China is described and placed in the largest assassin bug subfamily Harpactorinae (Hemiptera: Heteroptera: Reduviidae). The new species is similar to members of Isyndus Stål, 1859 in general habitus, but can be distinguished from the latter by the unarmed anterior pronotal lobe, the laterally dilated lobe-like abdominal segment V and several features of its genitalia.
Assassin bugs (Hemiptera: Heteroptera: Reduviidae) of Poland: systematic review, distribution, key to species]. This paper summarises the distribution of eleven species of the family Reduviidae in Poland basing on literature data and new records. Coranus aethiops is for the first time reported from Poland. Moreover, the key for identification of species occurring in Poland is also provided. High quality graphics have been prepared for each species, and in some cases, photos from nature were also included.
The subfamily Bactrodinae is revised. Its sole genus includes five species: Bactrodes femoratus (Fabricius), B. biannulatus Stål, B. spinulosus Stål, B. multiannulatus Berg, and B. misionensis, new species. A cladistic analysis; descriptions of the subfamily, the genus, and its species; and a key are included, as well as a map with geographical distributions.
A checklist of the 137 current valid species placed in the subfamily Triatominae and their geographical distribution is presented. Confirmed and doubtful synonyms are given, as are nomenclatural and taxonomic comments.
The monophyly of the 7 infraorders of Heteroptera and history of higher group concepts and interrelationships within the Heteroptera are briefly reviewed. Data from 31 morphological characters are combined with 669 bases of 18S nuclear rDNA for 29 taxa, including several outgroups to the Heteroptera, to produce a phylogeny based on the total available evidence. The molecular data alone and in conjunction with morphological data indicate that: the Homoptera are probably not monophyletic; the Auchenorrhyncha are the sister group of Coleorrhyncha + Heteroptera ; the Enicocephalomorpha are the sister group of remaining Heteroptera; the Dipsocoromorpha are the sister group of remaining Heteroptera; the Gerromorpha are the sister group of remaining Heteroptera; the Nepomorpha are the sister group of remaining Heteroptera; the Leptopodomorpha are the sister group of the Cimicomorpha + Pentatomomorpha. The molecular evidence corroborates the morphologically based theory of a sister group relationship between Aradoidea and trichophoran Pentatomomorpha. This scheme deviates from that previously published by Schuh, in which the Leptopodomorpha were treated as the sister group of the Nepomorpha.
Scanning electron microscopy observations are presented for freshly hatched nymphs and adults of Zelus longipes L., an hemipteran species of the family Reduviidae (assassin bugs). The emphasis is on the structure and distribution of pores and different types of hairs covering the tibiae of the forelegs in relation to a viscous substance associated with them. The substance is instrumental in catching prey. In both developmental stages of the animal, the legs are abundantly covered with two major types of hairs. There are the so-called sundew hairs characterized by spines on their upper part. The legs also carry smooth peg-like setae. In addition, the adult animals show a few smooth, needle-shaped hairs. At this developmental stage, the surface of the legs has developed ring-like invaginations. The animals catch prey using their raised forelegs that are covered with a layer of sticky substance. It is assumed that the sundew hairs in combination with the peg-like setae play a role in mechanically stabilizing the film of stick)' substance covering the legs in both developmental stages. The ring-like invaginations found only in the adult animals are interpreted as the external openings of integumental glands, responsible for the production of the substance. The openings are missing in freshly hatched nymphs, and it is assumed that they use the sticky substance deposited by the female at the bases of the laid eggs to set up their sticky traps.
The program MRBAYES performs Bayesian inference of
phylogeny using a variant of Markov chain Monte Carlo.
Availability: MRBAYES, including the source code, documentation,
sample data files, and an executable, is available at http://brahms.biology.rochester.edu/software.html.
Contact: johnh{at}brahms.biology.rochester.edu
Despite recent advances achieved by application of high-performance computing methods and novel algorithmic techniques to maximum likelihood (ML)-based inference programs, the major computational bottleneck still consists in the computation of bootstrap support values. Conducting a probably insufficient number of 100 bootstrap (BS) analyses with current ML programs on large datasets-either with respect to the number of taxa or base pairs-can easily require a month of run time. Therefore, we have developed, implemented, and thoroughly tested rapid bootstrap heuristics in RAxML (Randomized Axelerated Maximum Likelihood) that are more than an order of magnitude faster than current algorithms. These new heuristics can contribute to resolving the computational bottleneck and improve current methodology in phylogenetic analyses. Computational experiments to assess the performance and relative accuracy of these heuristics were conducted on 22 diverse DNA and AA (amino acid), single gene as well as multigene, real-world alignments containing 125 up to 7764 sequences. The standard BS (SBS) and rapid BS (RBS) values drawn on the best-scoring ML tree are highly correlated and show almost identical average support values. The weighted RF (Robinson-Foulds) distance between SBS- and RBS-based consensus trees was smaller than 6% in all cases (average 4%). More importantly, RBS inferences are between 8 and 20 times faster (average 14.73) than SBS analyses with RAxML and between 18 and 495 times faster than BS analyses with competing programs, such as PHYML or GARLI. Moreover, this performance improvement increases with alignment size. Finally, we have set up two freely accessible Web servers for this significantly improved version of RAxML that provide access to the 200-CPU cluster of the Vital-IT unit at the Swiss Institute of Bioinformatics and the 128-CPU cluster of the CIPRES project at the San Diego Supercomputer Center. These Web servers offer the possibility to conduct large-scale phylogenetic inferences to a large part of the community that does not have access to, or the expertise to use, high-performance computing resources.
The evolutionary history and times of divergence of triatomine bug lineages are estimated from molecular clocks inferred from nucleotide sequences of the small subunit SSU (18S) and the second internal transcribed spacer (ITS-2) of the nuclear ribosomal DNA of these reduviids. The 18S rDNA molecular clock rate in Triatominae, and Prosorrhynchan Hemiptera in general, appears to be of 1.8% per 100 million years (my). The ITS-2 molecular clock rate in Triatominae is estimated to be around 0.4-1% per 1 my, indicating that ITS-2 evolves 23-55 times faster than 18S rDNA. Inferred chronological data about the evolution of Triatominae fit well with current hypotheses on their evolutionary histories, but suggest reconsideration of the current taxonomy of North American species complexes.
In this paper we summarize the systematics of the 130 currently recognized species of Triatominae and the key features of their evolutionary background. There is increasing evidence that the subfamily has polyphyletic origins, with the various tribes and species groups probably arising from different reduviid lineages in relatively recent times.
MrBayes 3 performs Bayesian phylogenetic analysis combining information from different data partitions or subsets evolving under different stochastic evolutionary models. This allows the user to analyze heterogeneous data sets consisting of different data types—e.g. morphological, nucleotide, and protein—and to explore a wide variety of structured models mixing partition-unique and shared parameters. The program employs MPI to parallelize Metropolis coupling on Macintosh or UNIX clusters.
Availability: http://morphbank.ebc.uu.se/mrbayes
Contact: fredrik.ronquist@ebc.uu.se
*
To whom correspondence should be addressed.
The accuracy of multiple sequence alignment program MAFFT has been improved. The new version (5.3) of MAFFT offers new iterative
refinement options, H-INS-i, F-INS-i and G-INS-i, in which pairwise alignment information are incorporated into objective
function. These new options of MAFFT showed higher accuracy than currently available methods including TCoffee version 2 and
CLUSTAL W in benchmark tests consisting of alignments of >50 sequences. Like the previously available options, the new options
of MAFFT can handle hundreds of sequences on a standard desktop computer. We also examined the effect of the number of homologues
included in an alignment. For a multiple alignment consisting of ∼8 sequences with low similarity, the accuracy was improved
(2–10 percentage points) when the sequences were aligned together with dozens of their close homologues (E-value < 10−5–10−20) collected from a database. Such improvement was generally observed for most methods, but remarkably large for the new options
of MAFFT proposed here. Thus, we made a Ruby script, mafftE.rb, which aligns the input sequences together with their close
homologues collected from SwissProt using NCBI-BLAST.
Unlabelled:
RAxML-VI-HPC (randomized axelerated maximum likelihood for high performance computing) is a sequential and parallel program for inference of large phylogenies with maximum likelihood (ML). Low-level technical optimizations, a modification of the search algorithm, and the use of the GTR+CAT approximation as replacement for GTR+Gamma yield a program that is between 2.7 and 52 times faster than the previous version of RAxML. A large-scale performance comparison with GARLI, PHYML, IQPNNI and MrBayes on real data containing 1000 up to 6722 taxa shows that RAxML requires at least 5.6 times less main memory and yields better trees in similar times than the best competing program (GARLI) on datasets up to 2500 taxa. On datasets > or =4000 taxa it also runs 2-3 times faster than GARLI. RAxML has been parallelized with MPI to conduct parallel multiple bootstraps and inferences on distinct starting trees. The program has been used to compute ML trees on two of the largest alignments to date containing 25,057 (1463 bp) and 2182 (51,089 bp) taxa, respectively.
Availability:
icwww.epfl.ch/~stamatak
The accuracy and scalability of multiple sequence alignment (MSA) of DNAs and proteins have long been and are still important
issues in bioinformatics. To rapidly construct a reasonable MSA, we developed the initial version of the MAFFT program in
2002. MSA software is now facing greater challenges in both scalability and accuracy than those of 5 years ago. As increasing
amounts of sequence data are being generated by large-scale sequencing projects, scalability is now critical in many situations.
The requirement of accuracy has also entered a new stage since the discovery of functional noncoding RNAs (ncRNAs); the secondary
structure should be considered for constructing a high-quality alignment of distantly related ncRNAs. To deal with these problems,
in 2007, we updated MAFFT to Version 6 with two new techniques: the PartTree algorithm and the Four-way consistency objective
function. The former improved the scalability of progressive alignment and the latter improved the accuracy of ncRNA alignment.
We review these and other techniques that MAFFT uses and suggest possible future directions of MSA software as a basis of
comparative analyses. MAFFT is available at http://align.bmr.kyushu-u.ac.jp/mafft/software/.
Coscarón, Maria Del Carmen, Melo, Maria Cecilia (2003): Revision of the subfamily Bactrodinae (Heteroptera, Reduviidae), with a phylogenetic analysis of Bactrodes. Zootaxa 304: 1-15, DOI: 10.5281/zenodo.156879
Xenocaucus China and Usinger 1949, is fixed as type of the new tribe Xenocaucini of Tribelocephalinae. Abelocephala, new genus, and A. thai, new species, is described from Thailand. Key to the tribes and genera in Tribelocephalinae is given. Tomolus Stål is redescribed, the holotype is fixed, and a paratype of Tomolus costalis redescribed.
Certain Holoptilini (Reduviidae: Holoptilinae) have long been believed to attract and paralyze ants through secretions released from a sternal structure on the abdomen termed a "trichome." In this study, the trichome for adults of Ptilocnemus lemur is examined in detail, the first time in any Holoptilini. The trichome comprises a median elevation on abdominal sternite 2, a transverse groove, a median projection on sternite 3, and four groups of setae and hair-like projections. The hair-like projections (hairs of group 3 and 4) are shown to be associated with a layer of glandular units, which may be the source of ant-attracting or paralyzing secretions. Apart from these glandular units, a second type of glands is present on the sternum. These so far undocumented glandular units are arranged in distinct areas on the pregenitalic sternites, and are conspicuous by their large pores. Nymphs of P. lemur do not possess a trichome, but the glandular areas on the pregenitalic segments are already present.
Current methods of Chagas disease control rely mainly on elimination of the domestic vector populations. Domestication of these insects has clearly been a recent event in evolutionary terms, associated with a series of genetic and phenetic changes, and this paper reviews current knowledge about the sequence of events leading to the domestication of species of Rhodnius that are important vectors of Chagas disease in the Andean pact and Central American countries. Available evidence suggests that species of Rhodnius have radiated from an ancestral source in the Amazon region, giving three main adaptive lines: southwards into the cerrados of Brazil, northwards into the llanos of Venezuela, and northwestwards through the Andean Cordillera into the Magdalena valley of Colombia. There has also been specialisation within the Amazon forest itself. The form of radiative adaptation is predicted from morphological and biogeographic characters, and subsequently supported by a series of morphometric and genetic markers including mtDNA sequence analysis.
In the cocoa farms the members of family Reduviidae are distributed in a variety of habitats associated with particular food, defense and breeding requirements for various species. In general the ectrichodiines are exclusively litter dwellers and reduviines occur only under the loose bark of primary forest trees. The members of subfamily Harpactorinae are associated with herbs, shrubs and the foliage of cocoa trees. A group of harpactorines comprising Authenta, Margasus, Nagusta and Odontogonus has been found to be completely arboreal. The food and feeding behavior of the cocoa farm Reduviidae were explored and data on the insects preyed upon by the Reduviinae obtained for the first time. In one locality at Tafo, one species of this subfamily utilized 24 different species of ants belonging to four subfamilies. As a preliminary to feeding, the reduviids were generally observed to exhibit grooming behavior which at times was repeated once the bug had finished feeding. Coloration, mimicry and scent glands were considered to contribute greatly to the primary defense of the Reduviidae found in the cocoa farms. The majority of Reduviidae living in cocoa canopy or concealed situations such as leaf litter tended to be dark-colored, usually matching their surroundings. The reduviids at shrub level are conspicuously colored and their adults exhibit a behavior pattern highly suggestive of certain wasps. Excepting the Emesinae and Tribelocephalinae, where the scent glands are completely absent, repugnatorial glands producing generally a pungent secretion were found in all the reduviidae occurring in cocoa farms. A unique system of glands situated at the base of dorsoabdominal hairs was discovered for the first time. This system produces secretions which assist in sticking camouflage material to the back of reduviine larvae. Early instars of some species of Harpactorinae were observed to mimic smaller species of ants such as Pheidole and Anochetus. Courtship, mating and oviposition in Reduviidae were studied and the significance of the precopulatory riding position, parental care and laying of eggs singly or in groups is discussed.
— We studied sequence variation in 16S rDNA in 204 individuals from 37 populations of the land snail Candidula unifasciata (Poiret 1801) across the core species range in France, Switzerland, and Germany. Phylogeographic, nested clade, and coalescence analyses were used to elucidate the species evolutionary history. The study revealed the presence of two major evolutionary lineages that evolved in separate refuges in southeast France as result of previous fragmentation during the Pleistocene. Applying a recent extension of the nested clade analysis (Templeton 2001), we inferred that range expansions along river valleys in independent corridors to the north led eventually to a secondary contact zone of the major clades around the Geneva Basin. There is evidence supporting the idea that the formation of the secondary contact zone and the colonization of Germany might be postglacial events. The phylogeographic history inferred for C. unifasciata differs from general biogeographic patterns of postglacial colonization previously identified for other taxa, and it might represent a common model for species with restricted dispersal.
A phylogenetic analysis for the Cimicomorpha was conducted using 92 taxa, including eight outgroups and six species of Thaumastocoridae. Density of taxon sampling allows for tests of relationships at the family level for most taxa, whereas in the Miridae denser sampling allows for doing so on the tribal level. This level of sampling also corresponds with the availability of testable published hypotheses of relationships. Morphological data for 73 characters are coded for all taxa. Approximately 3500 base pairs of DNA were sequenced for the following gene regions for 83 taxa: 16S rDNA, 18S rDNA, 28S rDNA and COI. Results are presented for analysis of morphological data, individual molecular partitions, combined molecular data, combined morphological and molecular data for 83 taxa and combined morphological and molecular data for 92 taxa. Analyses of morphological data were performed using the parsimony programs NONA and PIWE: molecular and combined data were analysed using direct optimization with the program POY. Major conclusions of the present study include recognition of the following monophyletic groups: The Geocorisae is a monophyletic group. The monophyly of the Cimicomorpha - including Thaumastocoridae - is not supported in most analyses. The Reduviidae is monophyletic, with the Phymatinae Complex being the sister-group of the remaining subfamilies. The circumscription of the Cimiciformes is altered from the prior conception of Schuh and ˇ Stys to also include the Joppeicidae, Microphysidae and Velocipedidae, as well as the recently described family Curaliidae; the monophyly of the Cimiciformes is supported in most analyses; the Cimiciformes is treated as the sister-group of the Miroidea in most analyses. The monophyly of the Cimicoidea, including Curaliidae, is supported in all analyses including molecular data, whereas Curaliidae is treated as a more basal cimiciform in all other analyses. The monophyly and placement of the Thaumastocoridae is ambiguous across the range of analyses, and the monophyly of the Miroidea sensu Schuh and ˇ Stys receives limited support in the combined analyses of morphology þ molecular data. The Tingidae and Miridae are each monophyletic and together almost invariably form a monophyletic group. Within the Miridae, several inclusive monophyletic groups at the subfamily/tribal level are more or less consistently recognized when molecular data are included; however, the interrelationships of the subfamilies vary substantially across the range of analyses. Of the individual molecular partitions, only 18S rDNA shows significant congruence with combined analyses of morphological, combined molecular or combined morphological and molecular data. Scenarios are discussed for the evolution of the metathoracic scent- efferent system and the origin of the fossula spongiosa.
Evidences of the identity of the veins of the cubitalanal field of the hemipterous wing are given. In the fore wing of auchenorrhynchous Homoptera these veins are: Cm (anterior to the clavus) ; Cu2 (closely associated with the claval furrow) ; Pcu, 1st A, and 2d A (in the clavus). In Heteroptera Cu2 is absent, and the clavus contains only Pcu and 1st A. There is no anal lobe in either suborder, the clavus representing the postcubital region of the fore wing. In the hind wing of Auchenorrhyncha the veins of the cubital-anal field are the same as in the fore wing; a cubital furrow lies just behind Cu2, and the anal fold lies just behind 1st A. In Heteroptera this region of the hind wing is similar, except that Cu2 is absent, and the cubital furrow often is bifid and may contain secondary veins between its branches. In both Auchenorrhyncha and Heteroptera the anal lobe of the hind wing usually is well developed, and the jugal lohe is always absent. In the Cimicomorpha the hind-wing venation of the Miridae, Nabidae, Microphysidae, Anthocoridae, Joppeicidae, Thaumastocoridae, and Velocipedidae is of the same basic type, characterized particularly by the absence of Sc and by having a Y- or V-shaped pattern of Pcu and 1st A. The tingids resemble this group, but have a distinct Sc. The nabids and microphysids seem exceptional in sometimes having R and M separated distally, as in the Pentatomomorpha. The reduviid venation is distinct from that of the other Cimicomorpha, and is best characterized by a narrow postcubital sector containing subparallel Pcu and 1st A veins. Venation is useful in distinguishing subfamilies of the Nabidae. Anthocoridae, and Reduviidae; in the Reduviidae it serves in characterizing three major groups of subfamilies.
Representatives of the 8 nominal subfamilies comprising the harpactoroid complex have been studied and their various characters analyzed in terms of phylogenetic significance. These groups usually have in common 13 characters or character complexes most of which are apomorphic and, therefore, indicative of a close relationship between groups. The characters are absent or uncommon in other reduviid subfamilies. Analysis of the characters distinguishing the various harpactoroid groups indicates that, except for the Bactrodinae, these characters are relatively few and minor. Since most of the harpactoroid groups have in common many apomorphic characters and each has rather few autapomorphic characters, it is concluded that these groups should comprise a redefined subfamily Harpactorinae. Accordingly, the former Harpactorinae, as well as the Raphidosominae, Tegeinae, Apiomerinae, Diaspidinae, and Ectinoderinae, should be reduced to tribal status. The Phonolibinae are not sufficiently different from the tegeines to justify separate group treatment and should, therefore, be placed in the tribe Tegini. The Bactrodinae lack many of the harpactoroid characters and have several synapomorphic characters. Therefore, it is concluded that they should remain as a subfamily.
A multiple sequence alignment program, MAFFT, has been developed. The CPU time is drastically reduced as compared with existing methods. MAFFT includes two novel techniques. (i) Homo logous regions are rapidly identified by the fast Fourier transform (FFT), in which an amino acid sequence is converted to a sequence composed of volume and polarity values of each amino acid residue. (ii) We propose a simplified scoring system that performs well for reducing CPU time and increasing the accuracy of alignments even for sequences having large insertions or extensions as well as distantly related sequences of similar length. Two different heuristics, the progressive method (FFT-NS-2) and the iterative refinement method (FFT-NS-i), are implemented in MAFFT. The performances of FFT-NS-2 and FFT-NS-i were compared with other methods by computer simulations and benchmark tests; the CPU time of FFT-NS-2 is drastically reduced as compared with CLUSTALW with comparable accuracy. FFT-NS-i is over 100 times faster than T-COFFEE, when the number of input sequences exceeds 60, without sacrificing the accuracy.
The classification of the Reduvioid bugs has long remained in an unsettled state. Stål's masterful classification (1872, 1873) is still our best and only reference for subfamilies and genera and Reuter's general classification of the Heteroptera (1910) remains as the standard for higher groups. Evidence from the male genitalia (Singh-Pruthi, 1925) and from the mandibular levers (Ekblom, 1929) has since shown clearly that the semi-aquatic Mesoveliidae and Hebridae belong elsewhere. Despite a truly remarkable diversity, the group as thus restricted is apparently monophyletic.
Certain Holoptilini (Reduviidae: Holoptilinae) have long been believed to attract and paralyze ants through secretions released from a sternal structure on the abdomen termed a “trichome.” In this study, the trichome for adults of Ptilocnemus lemur is examined in detail, the first time in any Holoptilini. The trichome comprises a median elevation on abdominal sternite 2, a transverse groove, a median projection on sternite 3, and four groups of setae and hair-like projections. The hair-like projections (hairs of group 3 and 4) are shown to be associated with a layer of glandular units, which may be the source of ant-attracting or paralyzing secretions. Apart from these glandular units, a second type of glands is present on the sternum. These so far undocumented glandular units are arranged in distinct areas on the pregenitalic sternites, and are conspicuous by their large pores. Nymphs of P. lemur do not possess a trichome, but the glandular areas on the pregenitalic segments are already present.
Scanning electron microscopy observations are presented for freshly hatched nymphs and adults of Zelus longipes L., an hemipteran species of the family Reduviidae (assassin bugs). The emphasis is on the structure and distribution of pores and different types of hairs covering the tibiae of the forelegs in relation to a viscous substance associated with them. The substance is instrumental in catching prey. In both developmental stages of the animal, the legs are abundantly covered with two major types of hairs. There are the so-called sundew hairs characterized by spines on their upper part. The legs also carry smooth peg-like setae. In addition, the adult animals show a few smooth, needle-shaped hairs. At this developmental stage, the surface of the legs has developed ring-like invaginations. The animals catch prey using their raised forelegs that are covered with a layer of sticky substance. It is assumed that the sundew hairs in combination with the peg-like setae play a role in mechanically stabilizing the film of sticky substance covering the legs in both developmental stages. The ring-like invaginations found only in the adult animals are interpreted as the external openings of integumental glands, responsible for the production of the substance. The openings are missing in freshly hatched nymphs, and it is assumed that they use the sticky substance deposited by the female at the bases of the laid eggs to set up their sticky traps.
Abstract With more than 6600 species worldwide, Reduviidae (Insecta: Heteroptera), or assassin bugs, form the second largest and one of the most diverse groups of true bugs. The poor condition of the higher-level classification of Reduviidae is reflected by the facts that different authors recognize between 21 and 32 subfamily-level names and that Reduviidae were never subjected to a rigorous cladistic analysis using an exemplar approach. In the present study, a cladistic analysis of higher-level taxa of Reduviidae based on 162 morphological characters and 75 ingroup and outgroup species is presented. Twenty-one subfamily-level taxa of Reduviidae were examined, accounting for 28 tribes. In addition to characters previously used for diagnosis in Reduviidae, information on recently published character complexes is used in the present analysis, supplemented with new character information gathered specifically for this project. Reduviidae are supported as a monophyletic group with Pachynomidae as their sister taxon. The major results of this study are the support of a sistergroup relationship of Hammacerinae with the remaining Reduviidae, the monophyly of the Phymatine Complex, the relatively basal position of Harpactorinae within Reduviidae as well as a novel hypothesis on the relationships within this group, and the sistergroup relationship of Ectrichodiinae + Tribelocephalinae and their placement in a clade that also contains Emesinae, Saicinae, and Visayanocorinae. The analysis further supports a clade formed by paraphyletic Salyavatinae + Sphaeridopinae, renders Vesciinae non-monophyletic, and demonstrates the polyphyly of Reduviinae. Pseudocetherinae are shown to group with some Reduviinae. Triatominae are supported as a monophyletic group and are nested among additional Reduviinae and Stenopodainae.
Inference of phylogenetic trees comprising hundreds or even thousands of organisms based on the maximum likelihood method is computationally intensive. We present simple heuristics which yield accurate trees for synthetic as well as real data and significantly reduce execution time. Those heuristics have been implemented in a sequential, parallel, and distributed program called RAxML-II, which is freely available as open source code. We compare the performance of the sequential program with PHYML and MrBayes which—to the best of our knowledge—are currently the fastest and most accurate programs for phylogenetic tree inference based on statistical methods. Experiments are conducted using 50 synthetic 100 taxon alignments as well as nine real-world alignments comprising 101 up to 1000 sequences. RAxML-II outperforms MrBayes for real-world data both in terms of speed and final likelihood values. Furthermore, for real data RAxML-II requires less time (a factor of 2–8) than PHYML to reach PHYML's final likelihood values and yields better final trees due to its more exhaustive search strategy. For synthetic data MrBayes is slightly more accurate than RAxML-II and PHYML but significantly slower. The non-deterministic parallel program shows good speedup values and has been used to infer a 10 000-taxon tree comprising organisms from the domains Eukarya, Bacteria, and Archaea. Copyright © 2005 John Wiley & Sons, Ltd.
Morphometric analysis of Old World species of Linshcosteus and Triatoma showed the two genera to be well separated. However, although the analysis showed clear distinction between the five species of Linshcosteus, there was considerable overlap between the eight Old World species of Triatoma, including specimens of T. rubrofasciata from Old and New World populations. The results are interpreted as supporting the idea of a New World origin for the Old World species of Triatoma, with an independent Old World origin for Linshcosteus.
The most representative sample of molecular data, especially 16S and 12S rDNAs, is used to study the phylogeny and evolution of 57 species of three tribes, Rhodniini, Linshcosteini, and Triatomini, of the subfamily Triatominae. For the first time both New World and Old World species are brought together in a single phylogenetic analysis. Maximum-parsimony and distance estimation place both the Asiatic representatives, Linshcosteus and Triatoma rubrofasciata, as sister groups. The Linshcosteus-T. rubrofasciata clade nests firmly within Triatomini, in most analyses branching as a basalmost lineage, thus supporting a monophyletic origin of Triatominae. A paraphyly of "Triatoma" with respect to Linshcosteus, Dipetalogaster, Eratyrus, and Panstrongylus and the paraphyly of "Rhodnius" with respect to Psammolestes is observed in most of the analyses. Reinterpretation of triatomine biogeography points to the origin of Triatominae in northern areas of South America, in Central America, or in the southern region of North America. A few taxonomic changes are proposed: (1) reinclusion of Linshcosteus in Triatomini, (2) inclusion of Psammolestes in Rhodnius, (3) elevation of the "T. flavida complex" to the full genus Nesotriatoma (including N. flavida, N. bruneri, and N. obscura), (4) inclusion of the "T. spinolai complex" in Mepraia (including M. spinolai, M. gajardoi, M. eratyrusiformis, and M. breyeri), and (5) inclusion of "T." dimidiata in Meccus (M. dimidiatus).
In 2002, we developed and released a rapid multiple sequence alignment program MAFFT that was designed to handle a huge (up to approximately 5,000 sequences) and long data (approximately 2,000 aa or approximately 5,000 nt) in a reasonable time on a standard desktop PC. As for the accuracy, however, the previous versions (v.4 and lower) of MAFFT were outperformed by ProbCons and TCoffee v.2, both of which were released in 2004, in several benchmark tests. Here we report a recent extension of MAFFT that aims to improve the accuracy with as little cost of calculation time as possible. The extended version of MAFFT (v.5) has new iterative refinement options, G-INS-i and L-INS-i (collectively denoted as [GL]-INS-i in this report). These options use a new objective function combining the weighted sum-of-pairs (WSP) score and a score similar to COFFEE derived from all pairwise alignments. We discuss the improvement in accuracy brought by this extension, mainly using two benchmark tests released very recently, BAliBASE v.3 (for protein alignments) and BRAliBASE (for RNA alignments). According to BAliBASE v.3, the overall average accuracy of L-INS-i was higher than those of other methods successively released in 2004, although the difference among the most accurate methods (ProbCons, TCoffee v.2 and new options of MAFFT) was small. The advantage in accuracy of [GL]-INS-i became greater for the alignments consisting of approximately 50-100 sequences. By utilizing this feature of MAFFT, we also examined another possible approach to improve the accuracy by incorporating homolog information collected from database. The [GL]-INS-i options are applicable to aligning up to approximately 200 sequences, although not applicable to thousands of sequences because of time and space complexities.
Biological notes on the hemipteron Ptilocerus ochraceus Tijdschrift voor entomologie 54 denBeinen vonZelus (Diplocodus) MAFFT: a novel method for rapid multiple sequence alignment based on fast Fourier transform
- E G V Jacobson
- Zoological
- K Katoh
- K Misawa
- K Kuma
- T Miyata
Jacobson, E., 1911. Biological notes on the hemipteron Ptilocerus ochraceus. Tijdschrift voor entomologie 54, 175–179. denBeinen vonZelus (Diplocodus) Heteroptera Maynard, (Coleorrhyncha (Eds.), to G.V.Zoological the subfamilyBactrodinae MededelingenLandbouwhogeschool Katoh, K., Misawa, K., Kuma, K., Miyata, T., 2002. MAFFT: a novel method for rapid multiple sequence alignment based on fast Fourier transform. Nucleic Acids Res. 30, 3059–3066
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Die Fangdrusen an den Beinen von Zelus (Diplocodus) leucogrammus (Perty 1834) (Heteroptera, Reduviidae)
- R Barth
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leucogrammus (Perty 1834) (Heteroptera, Reduviidae). Zool. Jahrb. (Anat.) 73,
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