Sticky predators: a comparative study of sticky glands in harpactorine assassin bugs (Insecta: Hemiptera: Reduviidae)

Zhang, G. and Weirauch, C. 2011. Sticky predators: a comparative study of sticky glands in harpactorine assassin bugs (Insecta: Hemiptera: Reduviidae). —Acta Zoologica (Stockholm) 00 : 1–10. For more than 50 years, specialized dermal glands that secrete sticky substances and specialized setae have been known from the legs of New World assassin bugs in the genus Zelus Fabricius (Reduviidae: Harpactorinae). The gland secretions and specialized ‘sundew setae’ are involved in enhancing predation success. We here refer to this predation strategy as ‘sticky trap predation’ and the specialized dermal glands as ‘sticky glands’. To determine how widespread sticky trap predation is among Reduviidae, we investigated taxonomic distribution of sticky glands and sundew setae using compound light microscopical and scanning electron microscopical techniques and sampling 67 species of Reduviidae that represent 50 genera of Harpactorini. We found sticky glands in 12 genera of Harpactorini and thus show that sticky trap predation is much more widespread than previously suspected. The sticky glands vary in shape, size and density, but are always located in a dorsolateral position on the fore tibia. Sundew setae are present in all taxa with sticky glands with the exception of Heza that instead possesses unique lamellate setae. The sticky trap predation taxa are restricted to the New World, suggesting a New World origin of this unique predation strategy.     

Molecular phylogeny of Harpactorinae and Bactrodinae uncovers complex evolution of sticky trap predation in assassin bugs (Heteroptera: Reduviidae)

Sticky trap predation, the use of adhesive substances to trap and capture prey, is an intriguing yet poorly studied predatory strategy. Unique among known sticky trap predators, assassin bugs (Reduviidae) have evolved both exogenous and endogenous sticky trap predatory mechanisms: some trap their prey with sticky plant resins, some scavenge insects entrapped by sticky plant trichomes and others self‐produce sticky secretions. The evolution of these different strategies in assassin bugs is poorly understood due to the lack of comprehensive phylogenies. We reconstruct a phylogeny of Reduviidae (141 taxa; > 5000 bp) focusing on the Harpactorinae and Bactrodinae that engage in sticky trap predation. Ancestral state reconstruction, and temporal and geographical divergence analyses show that sticky trap predation techniques in assassin bugs evolved at least seven times independently since the late Cretaceous: use of sticky plant trichomes evolved as many as four times, resin‐use twice independently and once as a transition from trichome use, and ‘self‐stickiness’ once. Exogenous and endogenous sticky traps first appeared in the Neotropics, with the two exogenous mechanisms (resin and trichome use) subsequently evolving independently in the Old World. This study illustrates, for the first time, the complex evolutionary pattern of sticky trap predation within assassin bugs.     

Molecular phylogeny of the assassin bugs (Hemiptera: Reduviidae), based on mitochondrial and nuclear ribosomal genes

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.     

Evolutionary History of Assassin Bugs (Insecta: Hemiptera: Reduviidae): Insights from Divergence Dating and Ancestral State Reconstruction

Assassin bugs are one of the most successful clades of predatory animals based on their species numbers (∼6,800 spp.) and wide distribution in terrestrial ecosystems. Various novel prey capture strategies and remarkable prey specializations contribute to their appeal as a model to study evolutionary pathways involved in predation. Here, we reconstruct the most comprehensive reduviid phylogeny (178 taxa, 18 subfamilies) to date based on molecular data (5 markers). This phylogeny tests current hypotheses on reduviid relationships emphasizing the polyphyletic Reduviinae and the blood-feeding, disease-vectoring Triatominae, and allows us, for the first time in assassin bugs, to reconstruct ancestral states of prey associations and microhabitats. Using a fossil-calibrated molecular tree, we estimated divergence times for key events in the evolutionary history of Reduviidae. Our results indicate that the polyphyletic Reduviinae fall into 11–14 separate clades. Triatominae are paraphyletic with respect to the reduviine genus Opisthacidius in the maximum likelihood analyses; this result is in contrast to prior hypotheses that found Triatominae to be monophyletic or polyphyletic and may be due to the more comprehensive taxon and character sampling in this study. The evolution of blood-feeding may thus have occurred once or twice independently among predatory assassin bugs. All prey specialists evolved from generalist ancestors, with multiple evolutionary origins of termite and ant specializations. A bark-associated life style on tree trunks is ancestral for most of the lineages of Higher Reduviidae; living on foliage has evolved at least six times independently. Reduviidae originated in the Middle Jurassic (178 Ma), but significant lineage diversification only began in the Late Cretaceous (97 Ma). The integration of molecular phylogenetics with fossil and life history data as presented in this paper provides insights into the evolutionary history of reduviids and clears the way for in-depth evolutionary hypothesis testing in one of the most speciose clades of predators.     

Functional diversity and trade‐offs in divergent antipredator morphologies in herbivorous insects

Predator–prey interactions may be responsible for enormous morphological diversity in prey species. We performed predation experiments with morphological manipulations (ablation) to investigate the defensive function of dorsal spines and explanate margins in Cassidinae leaf beetles against three types of predators: assassin bugs (stinger), crab spiders (biter), and tree frogs (swallower). There was mixed support for the importance of primary defense mechanisms (i.e., preventing detection or identification). Intact spined prey possessing dorsal spines were more likely to be attacked by assassin bugs and tree frogs, while intact armored prey possessing explanate margins were likely to avoid attack by assassin bugs. In support of the secondary defense mechanisms (i.e., preventing subjugation), dorsal spines had a significant physical defensive function against tree frogs, and explanate margins protected against assassin bugs and crab spiders. Our results suggest a trade‐off between primary and secondary defenses. Dorsal spines improved the secondary defense but weakened the primary defense against tree frogs. We also detected a trade‐off in which dorsal spines and explanate margins improved secondary defenses against mutually exclusive predator types. Adaptation to different predatory regimes and functional trade‐offs may mediate the diversification of external morphological defenses in Cassidinae leaf beetles.     

On the evolution of raptorial legs – an insect example (Hemiptera: Reduviidae: Phymatinae)

The presence of chelate and subchelate fore legs in Phymatinae (Hemiptera: Reduviidae), or ambush bugs, provides a unique opportunity to study the evolution of different types of raptorial legs in a closely related group of arthropods. Themonocorini have simple, possibly raptorial legs, Phymatini and Macrocephalini distinct subchelate fore legs, and the charismatic Carcinocorini are the only insects with a chelate fore leg apart from female dryinid Chysidoidea (Hymenoptera). Relationships between the four phymatine tribes are here analyzed in a cladistic framework thus permitting testable hypotheses on the evolution of raptorial legs. The presented analysis of phymatine tribal level relationships is based on a dataset comprising 11 species of Phymatinae and 54 non‐phymatine Reduviidae and Heteroptera. The molecular data set consists of ～3500 MAFFT aligned bases of 16S, 28S D2–D3, and 18S ribosomal genes. Parsimony and maximum likelihood analyses resulted in identical topologies for the ingroup with the relationships Themonocorini + (Phymatini + (Carcinocorini + Macrocephalini)) receiving high support values. Eleven morphological characters, eight of them derived from fore leg morphology, were optimized on the parsimony analysis. These optimizations indicate that the ancestral ambush bug had a simple raptorial leg; that size reduction of the tarsus, enlargement of the femur, curvature of the fore tibia, armature of tibia and femur with rows of tiny tubercles that allow for gripping of a prey insect, and the large process on the ventral surface of the femur arose in the common ancestor of Carcinocorini + Macrocephalini + Phymatini. The chelate leg in Carcinocorini is likely derived from a subchelate precursor similar to the one seen in recent Macrocephalini and may have evolved through elongation of the ventral, proximal portion of the fore femur and modification of the median process to form part of the digitus fixus. © The Willi Hennig Society 2010.     

Geological Changes of the Americas and their Influence on the Diversification of the Neotropical Kissing Bugs (Hemiptera: Reduviidae: Triatominae)

Background

The family Reduviidae (Hemiptera: Heteroptera), or assassin bugs, is among the most diverse families of the true bugs, with more than 6,000 species. The subfamily Triatominae (kissing bugs) is noteworthy not simply because it is the only subfamily of the Reduviidae whose members feed on vertebrate blood but particularly because all 147 known members of the subfamily are potential Chagas disease vectors. Due to the epidemiological relevance of these species and the lack of an efficient treatment and vaccine for Chagas disease, it is more common to find evolutionary studies focusing on the most relevant vectors than it is to find studies aiming to understand the evolution of the group as a whole. We present the first comprehensive phylogenetic study aiming to understand the events that led to the diversification of the Triatominae.

Methodology/Principal Findings

We gathered the most diverse samples of Reduviidae and Triatominae (a total of 229 Reduviidae samples, including 70 Triatominae species) and reconstructed a robust dated phylogeny with several fossil (Reduviidae and Triatominae) calibrations. Based on this information, the possible role of geological events in several of the major cladogenetic events within Triatominae was tested for the first time. We were able to not only correlate the geological changes in the Neotropics with Triatominae evolution but also add to an old discussion: Triatominae monophyly vs. paraphyly.

Conclusions/Significance

We found that most of the diversification events observed within the Rhodniini and Triatomini tribes are closely linked to the climatic and geological changes caused by the Andean uplift in South America and that variations in sea levels in North America also played a role in the diversification of the species of Triatoma in that region.     

Millipede assassins and allies (Heteroptera: Reduviidae: Ectrichodiinae,Tribelocephalinae): total evidence phylogeny,revised classification and evolution of sexual dimorphism

Evolution of sexual dimorphism in animals has long been of interest to scientists, but relatively few studies have reconstructed evolutionary patterns of extreme sexual dimorphism at a phylogenetic scale, especially in insects. Millipede assassin bugs (Heteroptera: Reduviidae: Ectrichodiinae; 736 spp.) and their sister taxon, Tribelocephalinae (150 spp.), exhibit sexual dimorphism that ranges from limited to extreme, a phenomenon apparently modulated by female morphology. Here, we reconstruct the first phylogeny for the subfamilies Ectrichodiinae and Tribelocephalinae with comprehensive generic representation (152 taxa in 72 genera) using morphological and molecular data (six gene regions). The combined phylogenetic results indicate that Tribelocephalinae are paraphyletic with respect to Ectrichodiinae, and that Ectrichodiinae themselves are polyphyletic. Based on these results, we synonymize Tribelocephalinae with Ectrichodiinae syn.n. , describe three new tribes (Ectrichodiini trib.n. , Tribelocodiini trib.n. , and Abelocephalini trib.n. ) and two new subtribes (Opistoplatyina subtrib.n. and Tribelocephalina subtrib.n. ), and revise Tribelocephalini sensu n. Ancestral state reconstruction of sexual dimorphism reconstructed limited sexual dimorphism in the ancestor of Ectrichodiinae sensu n. with at least seven evolutionary transitions to extreme sexual dimorphism within the clade. This published work has been registered in ZooBank, http://zoobank.org/urn:lsid:zoobank.org:pub:C810E20F‐D66A‐461F‐A0E6‐AB1073EA3E3C .     

Cladistic analysis of Reduviidae (Heteroptera: Cimicomorpha) based on morphological characters

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.     

Entrapped carrion increases indirect plant resistance and intra‐guild predation on a sticky tarweed

Many plants employ indirect defenses against herbivores; often plants provide a shelter or nutritional resource to predators, increasing predator abundance, and lessening herbivory to the plant. Often, predators on the same plant represent different life stages and different species. In these situations intraguild predation (IGP) may occur and may decrease the efficacy of that defense. Recently, several sticky plants have been found to increase indirect defense by provisioning predatory insects with entrapped insects (hereafter: carrion). We conducted observational studies and feeding trials with herbivores and predators on two sticky, insect‐entrapping asters, Hemizonia congesta and Madia elegans, to construct food webs for these species and determine the prevalence of IGP in these carrion‐provisioning systems. In both systems, intraguild predation was the most common interaction observed. To determine whether IGP was driven by resource abundance, whether it reduced efficacy of this indirect defense and whether stickiness or predator attraction was induced by damage, we performed field manipulations on H. congesta. Carrion supplementation led to an increase in predator abundance and IGP. IGP was asymmetric within the predator guild: assassin bugs and spiders preyed on small stilt bugs but not vice versa. Despite increased IGP, carrion provisions decreased the abundance of the two most common herbivores (a weevil and a mealybug). Overall seed set was driven by plant size, but number of seeds produced per fruit significantly increased with increasing carrion, likely because of the reduction in the density of a seed‐feeding weevil. Observationally and experimentally, we found that carrion‐mediated indirect defense of tarweeds led to much intraguild predation, though predators effectively reduced herbivore abundance despite the increase in IGP.     

Araneophagic behavior of Gardena brevicollis Stål (Heteroptera: Reduviidae): Foraging on pre‐dispersal spiderlings in the spider's web

Araneophagic behavior of an Emesinae assassin bug, Gardena brevicollis Stål (Hemiptera: Reduviidae) was observed in the field. The bug invaded a web of Acusilas coccineus Simon (Araneae: Araneidae) spiderlings and hunted five juvenile spiders. The bug showed stalking tactics to prey on spiders, whereas luring tactics, which have been reported in other Emesinae bugs, were not observed. To the best of my knowledge, the present study is the first report of araneophagy in the genus Gardena and the first report of araneophagic behavior in East Asian assassin bugs.     

The evolutionary position of Lestoniidae revealed by molecular autapomorphies in the secondary structure of rRNA besides phylogenetic reconstruction (Insecta: Hemiptera: Heteroptera)

Pentatomoidea (stink bugs and their relatives) is the third largest superfamily in Heteroptera, or the true bugs. The phylogenetic relationships among the families within Pentatomoidea remain controversial. The family Lestoniidae is morphologically highly specialized, currently including only two species endemic to Australia. Previous researchers have suggested a close relationship of Lestoniidae to either Plataspidae or Acanthosomatidae, based on morphological characters. In this study, phylogenetic tree reconstruction revealed that Lestoniidae and Acanthosomatidae form a monophyletic clade. In addition, in comparisons of the secondary structures of 18S and 28S rRNAs representing 15 families of Pentatomoidea, four length‐variable regions in 18S and 28S rRNAs that can serve as autapomorphies for the clade Lestoniidae + Acanthosomatidae were recognized. Among them, E in 18S rRNA and D3‐1 and D5‐1 in 28S rRNA are unique in length in Lestoniidae and Acanthosomatidae. Based on the new molecular evidence and morphological evidence published by previous authors, Lestoniidae is suggested to be a highly specialized group derived from a common ancestor with Acanthosomatidae.     

Phylogenetic comparative analysis supports aposematic colouration–body size association in millipede assassins (Hemiptera: Reduviidae: Ectrichodiinae)

The diversity of colour patterns and its importance in interactions with the environment make colouration in animals an intriguing research focus. Aposematic colouration is positively correlated with body size in certain groups of animals, suggesting that warning colours are more effective or that crypsis is harder to achieve in larger animals. Surprisingly, this relationship has not been recovered in studies investigating insects, which may have been confounded by a focus on aposematic taxa that are also gregarious. Millipede assassin bugs (Hemiptera: Reduviidae: Ectrichodiinae) comprise species with cryptic and aposematic colour patterns across a range of body sizes, are typically solitary as adults and are thus an excellent model for investigating a possible association between colouration and body size. Here, we use a comprehensive phylogeny for Ectrichodiinae, ancestral state reconstruction of colouration, and phylogenetic comparative methods to test for a colouration–body size association. The ancestor of Ectrichodiinae is reconstructed as cryptically coloured, with multiple subsequent transitions between aposematic and cryptic colouration. Aposematic colouration is positively associated with male body length and supports the hypothesis that selection on Ectrichodiinae body size may influence evolutionary transitions between aposematic and cryptic colouration or alternatively that selection for aposematic colouration influences body size evolution.     

Untangling the assassin's web: Phylogeny and classification of the spider-associated Emesine complex (Hemiptera: Reduviidae)

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.     

Identifying the predator complex of <Emphasis Type="Italic">Homalodisca vitripennis</Emphasis> (Hemiptera: Cicadellidae): a comparative study of the efficacy of an ELISA and PCR gut content assay

A growing number of ecologists are using molecular gut content assays to qualitatively measure predation. The two most popular gut content assays are immunoassays employing pest-specific monoclonal antibodies (mAb) and polymerase chain reaction (PCR) assays employing pest-specific DNA. Here, we present results from the first study to simultaneously use both methods to identify predators of the glassy winged sharpshooter (GWSS), Homalodisca vitripennis (Germar) (Hemiptera: Cicadellidae). A total of 1,229 arthropod predators, representing 30 taxa, were collected from urban landscapes in central California and assayed first by means of enzyme-linked immunosorbent assay (ELISA) using a GWSS egg-specific mAb and then by PCR using a GWSS-specific DNA marker that amplifies a 197-base pair fragment of its cytochrome oxidase gene (subunit I). The gut content analyses revealed that GWSS remains were present in 15.5% of the predators examined, with 18% of the spiders and 11% of the insect predators testing positive. Common spider predators included members of the Salticidae, Clubionidae, Anyphaenidae, Miturgidae, and Corinnidae families. Common insect predators included lacewings (Neuroptera: Chrysopidae), praying mantis (Mantodea: Mantidae), ants (Hymenoptera: Formicidae), assassin bugs (Hemiptera: Reduviidae), and damsel bugs (Hemiptera: Nabidae). Comparison of the two assays indicated that they were not equally effective at detecting GWSS remains in predator guts. The advantages of combining the attributes of both types of assays to more precisely assess field predation and the pros and cons of each assay for mass-screening predators are discussed.     

Living in the danger zone: Exposure to predators and the evolution of spines and body armor in mammals

Mammals have independently evolved a wide variety of morphological adaptations for use in avoiding death by predation, including spines, quills, dermal plates, and noxious sprays. Although these traits appear to protect their bearer from predatory attack, it is less obvious why some species evolved them and others have not. We investigated the ecological correlates favoring the evolution of specialized defenses in mammals, focusing on conspicuousness to predators due to body size and openness of habitat. We scored species for the degree to which they are protected by spines, quills, dermal plating, and sprays and used phylogenetic comparative analyses to study the morphological and ecological factors that may favor their evolution. We show that medium‐sized insectivorous mammals (～800 g to 9 kg) that live in open habitats are more likely to possess one of these defensive traits to reduce predation. Smaller species (<200 g) and those in closed habitats can typically rely on crypsis to avoid predators, and larger species (>10 kg) are less susceptible to predation by most small‐ to medium‐sized predators. We discuss how diet, metabolic rate, and defensive strategy evolve in concert to allow species to exploit this ecomorphological “danger zone” niche.     

Evolution of wing polymorphism and genital asymmetry in the thread‐legged bugs of the tribe Metapterini Stål (Hemiptera,Reduviidae, Emesinae) based on morphological characters

Wing polymorphism and asymmetric male genitalia are intriguing morphological phenomena occurring in insects. Among Emesinae, or thread‐legged bugs, the tribe Metapterini Stål exhibits these two interesting morphological attributes. Nonetheless, evolutionary interpretations of these phenomena cannot be put forward because phylogenetic hypotheses for Emesinae are lacking. Thread‐legged bugs are easily recognized among assassin bugs due to their elongated and seemingly delicate body. The tribe Metapterini has 28 genera and approximately 280 described species. The only available phylogenetic hypothesis among Emesinae tribes was proposed by Wygodzinsky (1966), and it hypothesized Deliastini Villiers as the sister group of Metapterini, although this hypothesis has never been tested with cladistic approaches. Recent analyses using character sets of genitalia and prolegs suggest that Metapterini might not be monophyletic. In order to test these ideas, we compiled a morphological dataset of 138 characters that includes external morphological characters, detailed features of prolegs and genitalia of both sexes for Metapterini, which were analysed cladistically including 55 terminals, comprising 24 genera (85.7% of the generic diversity), 43 species of Metapterini and 12 outgroups. Metapterini was recovered as paraphyletic by the inclusion of Bergemesa Wygodzinsky, Palacus Dohrn and Stalemesa Wygodzinsky, all currently assigned to Deliastini. Gardena Dohrn (Emesini) was recovered as the sister group of Metapterini + Deliastini as suggested by Wygodzinsky (1966). Based on these results, we synonymize Deliastini syn. n. with Metapterini sensu n. and propose two new genera: Bacata Castro‐Huertas & Forero gen. n. , for three Andean species previously placed in Liaghinella Wygodzinsky, and Valkyriella Castro‐Huertas & Forero gen. n. for Ghilianella borgmeieri Wygodzinsky. Ancestral state reconstruction of wing polymorphism indicates that males and females were fully winged in the ancestor of Metapterini sensu n. with two independent evolutionary transitions to the apterous and brachypterous conditions. The analysis of the symmetry of the male genitalia shows an ancestor with symmetric male genitalia and two independent emergences of asymmetrical male genitalia in Metapterini.     

A survey of the tribe Ectinoderini (Hemiptera: Reduviidae: Harpactorinae) of China,with the description of a new Ectinoderus species

The two resin assassin bugs belonging to the tribe Ectinoderini (Hemiptera: Reduviidae: Harpactorinae) occurring in China are documented: Amulius malayus Stål, 1867 is redescribed and Ectinoderus sinicus Chen & Cai, sp. nov. is described as new. The discovery of the new species extends the distribution of the genus and the tribe northwards from tropical Southeast Asia to southern mainland China. A key to the two ectinoderine genera and species from China is provided to facilitate their identification. DNA barcodes and bionomic data are provided for the species concerned, and their conservation is briefly discussed.http://zoobank.org/urn:lsid:zoobank.org:pub:A7F7402E-E9A8-4FAD-B72A-4B1C9B2A7276.