Molecular phylogenetics and biogeography of the ambush bugs (Hemiptera: Reduviidae: Phymatinae).

The ambush bugs (Heteroptera: Reduviidae: Phymatinae) are a diverse clade of predators known for their cryptic hunting behavior and morphologically diverse raptorial forelegs. Despite their striking appearance, role as pollinator predators, and intriguing biogeographic distribution, phylogenetic relationships within Phymatinae are largely unknown and the evolutionary history of the subfamily has remained in the dark. We here utilize the most extensive molecular phylogeny of ambush bugs to date, generated from a 3328 base pair molecular dataset, to refine our understanding of phymatine relationships, estimate dates of divergence (BEAST 2), and uncover historical biogeographic patterns (S-DIVA and DEC). This taxon set (39 species of Phymatinae and six outgroups) allowed reevaluation of the proposed sister group of Phymatinae and tribal-level relationships within the group, and for the first time proposes species-level relationships within Phymata Latreille, the largest genus of ambush bugs (∼109spp.). Available evidence suggests that Phymata originated in the Neotropical region, with subsequent dispersals to the Nearctic and Palearctic regions. This study provides a framework for future research investigating the evolutionary history of ambush bugs, as well as ecological and microevolutionary investigations.

Origin, evolution and function of the hemipteran perimicrovillar membrane with emphasis on Reduviidae that transmit Chagas disease.

The peritrophic matrix is a chitin-protein structure that envelops the food bolus in the midgut of the majority of insects, but is absent in some groups which have, instead, an unusual extra-cellular lipoprotein membrane named the perimicrovillar membrane. The presence of the perimicrovillar membrane (PMM) allows these insects to exploit restricted ecological niches during all life stages. It is found only in some members of the superorder Paraneoptera and many of these species are of medical and economic importance. In this review we present an overview of the midgut and the digestive system of insects with an emphasis on the order Paraneoptera and differences found across phylogenetic groups. We discuss the importance of the PMM in Hemiptera and the apparent conservation of this structure among hemipteran groups, suggesting that the basic mechanism of PMM production is the same for different hemipteran species. We propose that the PMM is intimately involved in the interaction with parasites and as such should be a target for biological and chemical control of hemipteran insects of economic and medical importance.

A new species of the genus Sphedanolestes Stål 1866 (Hemiptera: Reduviidae: Harpactorinae) from China, with a key to Chinese species.

A new harpactorine bug, Sphedanolestes zhengi sp. nov., is described and illustrated based on specimens collected from southwestern China. A key to 17 Chinese species of the genus, including the new species, is provided. The bionomics of the new species is briefly noted.

Revision of the genus Narvesus Stål, 1859 (Hemiptera: Heteroptera: Reduviidae: Stenopodainae) and a new record for Argentina.

This is a revision of Narvesus Stål, a Neotropical genus with two species: Narvesus carolinensis Stål 1859 and Narvesus minor Barber 1930. Both known species are redescribed and illustrated, including data on male genitalia and new distributional records. We compared with the genus Diaditus Stål.

Description of a new species of the genus Henricohahnia Breddin (Hemiptera: Heteroptera: Reduviidae: Harpactorinae) from China, with a key to Chinese species.

Henricohahnia parva sp. nov. is described and illustrated based on the specimens collected from Yunnan Province of China. It represents the smallest-sized species of the genus Henricohahnia Breddin so far known. A key to the eight Chinese species of the genus is provided.

Flexitibia, a new genus of Harpactorinae (Hemiptera: Heteroptera: Reduviidae), with a discussion on the functional morphology of forelegs of the related genera.

Flexitibia, a new genus, in the division Euagorasaria of the assassin bug subfamily Harpactorinae from Yunnan Province of China is described. The type species, Flexitibia orientalis sp. nov., is described and illustrated. A key to the closely related genera is provided. The type specimens are kept in the Entomological Museum of China Agricultural University, Beijing.

A new genus, Neoschidium (Hemiptera: Reduviidae: Emesinae), with a redescription of the type genus, Neoschidium phasma (Distant) [Ghilianella phasma Distant and Schidium phasma (Distant)], recorded for the first time from India.

A new genus, Neoschidium was erected with the type genus, Neoschidium phasma (Distant). It was earlier described under Ghilianella Spinola 1850 as G. phasma Distant and later under Schidium Bergroth 1916 as Schidium phasma (Distant) by Bergroth (1916). Because it exhibits characters not only of Ghilianella and Schidium but also intermediate specific characters that are not found in both the genera, the type genus Neoschidium phasma (Distant) is redescribed with additional taxonomic details, morphometrics, and illustrations. It is also recorded for the first time from India.

A review of the taxonomic position of Vesbius seyrigi Villiers, 1970, with a description of two new species of Peyrierocoris Chlond & Junkiert, 2010 (Hemiptera: Heteroptera: Reduviidae: Harpactorinae).

Peyrierocoris seyrigi (Villiers, 1970) is redescribed and its taxonomic position is revised to accommodate Vesbius seyrigi Villiers, 1970, erroneously described under the generic name Vesbius Stål, 1865. Redescription and drawings of the dorsal habitus and characters of the male and female genitalia, as well as a drawing of the ovarian egg of P. seyrigi (Villiers, 1970), are given. Two new species, P. gorczycai sp. nov. and P. herczeki sp. nov., are described and illustrated with 21 figures.

Intrageneric phylogenetics based on mitochondrial DNA variation among fifteen harpactorine assassin bugs with four ecotypes and three morphs (Hemiptera: Reduviidae: Harpactorinae).

Available mitochondrial DNA sequences viz., 16S, Cyt b, Cyt c oxidase subunit - I, and Cyt c subunit-like - I gene of Rhynocoris (Kolenati) species were subjected to phylogenetic analysis to understand the intrageneric and intraspecific variations and the role of geographical isolation on speciation; using CLUSTAL W in MEGA version 5.1. This analysis includes fifteen species and four ecotypes of R. kumarii Ambrose and Livingstone and three morphs of R. marginatus (Fabricius) from four countries viz., Canada, China, Korea, and South Africa. The pairwise genetic distances were calculated and phylograms were constructed using Maximum Likelihood, Maximum Parsimony, and Neighbor-Joining methods. These preliminary analyses not only demarcated the fifteen species of Rhynocoris, the four ecotypes of R. kumarii, and the three morphs of R. marginatus, but also revealed phylogenetic relationships and the role of geographical isolation and polymorphism on speciation.

Phylogeny and character evolution in the bee-assassins (Insecta: Heteroptera: Reduviidae).

Apiomerus, the charismatic bee-assassins (>108 spp.), belong to the New World resin bugs in the harpactorine tribe Apiomerini (12 extant genera) that is characterized by a novel predation strategy, resin trap predation. Apiomerini also exhibit striking genitalic diversity that has shaped subgeneric classifications within the genus Apiomerus and females of some species of Apiomerus are known to engage in unique maternal care behaviors. The lack of a phylogenetic framework currently hinders evolutionary interpretations of genitalic morphology and maternal care. We here present a molecular phylogeny based on 4, 477 bp of six ribosomal and protein coding genes and 95 terminal taxa using parsimony and maximum likelihood approaches as a way of addressing these shortcomings. Apiomerini are monophyletic, with Heniartes being the sistergroup to all remaining taxa that form the monophyletic Manicocoris (Calliclopius, Manicocoris, Micrauchenus, and Ponerobia) and Apiomerus (Agriocoris, Apiomerus, and Sphodrolestes) clades. Previously proposed subgeneric groups are polyphyletic, but several proposed species groups are recovered as monophyletic. Ancestral state reconstruction of the metatibial comb indicates that this structure evolved in the ancestor of all Apiomerini where it was present in males and in females; it became strongly sexually dimorphic (better developed in females than in males) in the Apiomerus clade (Apiomerus + Agriocoris + Sphodrolestes). Genitalic features reveal a pattern of homoplasy, but frequently are nonetheless useful to diagnose supraspecific groups within Apiomerus. The complex genitalia found within Apiomerus are derived for that clade. We conclude that, using the metatibial comb as a proxy, maternal care is relatively common in the tribe Apiomerini and propose that it likely evolved at the base of the Apiomerus clade if not at the base of Apiomerini.

Revalidation and redescription of Triatoma brasiliensis macromelasoma Galvão, 1956 and an identification key for the Triatoma brasiliensis complex (Hemiptera: Reduviidae: Triatominae).

Triatoma brasiliensis macromelasoma is revalidated based on the results of previous multidisciplinary studies on the Triatoma brasiliensis complex, consisting of crossing experiments and morphological, biological, ecological and molecular analyses. These taxonomic tools showed the closest relationship between T. b. macromelasoma and Triatoma brasiliensis brasiliensis. T. b. macromelasoma is redescribed based on specimens collected in the type locality and specimens from a F1 colony. The complex now comprises T. b. brasiliensis, T. b. macromelasoma, Triatoma melanica, Triatoma juazeirensis and Triatoma sherlocki. An identification key for all members of the complex is presented. This detailed comparative study of the morphological features of T. b. macromelasoma and the remaining members of the complex corroborates results from multidisciplinary analyses, suggesting that the subspecific status is applicable. This subspecies can be distinguished by the following combination of features: a pronotum with 1+1 narrow brownish-yellow stripes on the submedian carinae, not attaining its apex, hemelytra with membrane cells darkened on the central portion and legs with an incomplete brownish-yellow ring on the apical half of the femora. Because the T. brasiliensis complex is of distinct epidemiological importance throughout its geographic distribution, a precise identification of its five members is important for monitoring and controlling actions against Chagas disease transmission.

A new synonymy of Graptocleptes bicolor (Burmeister), with taxonomical notes (Hemiptera: Heteroptera: Reduviidae: Harpactorinae: Harpactorini).

Hiranetis coleopteroides (Walker, 1873) is here found to be conspecific with Graptocleptes bicolor (Burmeister, 1838). Graptocleptes bicolor is redescribed and the male genitalia characters are illustrated for the first time. Intraspecific morphological, color and male genitalia variability are discussed. Furthermore, the species is recorded from Paraguay for the first time.

Two new species of Oncerotrachelus Stål (Hemiptera: Heteroptera: Reduviidae: Saicinae), from Amazonas, Brazil, with taxonomical notes.

Oncerotrachelus amazonensis sp. nov. and O. paraconformis sp. nov. from Amazonas, Brazil, are described. Oncerotrachelus conformis Uhler is recorded for the first time from Brazil and a description of male genitalia of this species is provided.

A new genus and species of ambush bugs from South Africa (Hemiptera: Heteroptera: Reduviidae: Phymatinae).

Doesburgella gen. nov. and its type species D. dilatata sp.nov (Hemiptera, Heteroptera: Reduviidac, Phymatinae; Macrocephalini) are described from South Africa.

Berengeria Gil-Santana & Coletto-Silva, a junior synonym of Ectrichodiella Fracker & Bruner, with new records and taxonomic notes on Ectrichodiinae from Brazil, and with keys to Ectrichodiinae and Reduviinae genera of the New World (Hemiptera: Heteroptera: Reduviidae).

Berengeria Gil-Santana & Coletto-Silva, 2005 is considered a junior synonym of Ectrichodiella Fracker & Bruner, 1924. Ectrichodiella minima (Valdés, 1910) and E. rafaeli (Gil-Santana & Coletto-Silva, 2005), new. comb. are redescribed. Taxonomic notes on Brontostoma alboannulatum (Stål, 1860), B. discus (Burmeister, 1835), B. nanus Carpintero, 1980, B. rubrovenosum (Stål, 1860), and B. trux (Stål, 1859) are given. Brontostoma diringshofeni Gil-Santana & Baena, 2009, B. nanus, and Racelda robusta Bérenger & Gil-Santana, 2005 are recorded from Brazil for the first time. Keys to Ectrichodiinae and Reduviinae genera of the New World are presented.

Description and Ecology of A New Cavernicolous, Arachnophilous Thread- legged Bug (Hemiptera: Reduviidae: Emesini) from Kartchner Caverns, Cochise County, Arizona.

A new cavernicolous, arachnophilous thread-legged bug (Phasmatocoris labyrinthicus sp. nov.; Reduviidae: Emesini) is described from Kartchner Caverns, a limestone cavern in Kartchner Caverns State Park near Benson, Arizona, USA. Cavernicolous emesines are recorded from caves in many parts of the world and are distributed across several genera, but are generally uncommon. P. labyrinthicus shows no obvious troglomorphy but ecological evidence suggests it is, at minimum, a cave-limited troglophile. The species seems to be low-humidity intolerant, due to its occurrence in a cave within a desert region, effectively confines the population to the cave, and the species may thus actually be troglobitic by default. Arachnophily in emesines is more common, including in Phasmatocoris Breddin, but has been previously documented in only a single cavernicolous species, Bagauda cavernicola Paiva, reported from India, Malaysia and Sri Lanka. However, unlike P. labyrinthicus, B. cavernicola is apparently not morphologically adapted for its arachnophilous association. P. labyrinthicus is the only known troglophilic emesine that is also a morphologically adapted and behaviorally functional arachnophile. The only other known cavernicolous Phasmatocoris (P. xavieri Gil-Santana, Alves, Barrett and Costa) is recorded from a sandstone cave in Brazil. P. xavieri exhibits morphological features indicative of a potentially arachnophilous habit, but its ecology has not been studied. Adults of P. labyrinthicus share characteristics with the species Phasmatocoris praecellens Bergroth, P. minor McAtee and Malloch, P. xavieri, P. spectrum Breddin, and P. rapax McAtee and Malloch. Phasmatocoris is primarily a Neotropical genus and the discovery of P. labyrinthicus represents a significant range extension for the genus, being the first Nearctic species identified, with its geographically nearest relative an undescribed species from Mazatlan, Mexico, over 1,000 km to the south.

Outside the pattern: Evolution of the genital asymmetry in Saicinae (Hemiptera: Heteroptera: Reduviidae).

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.

Gross morphology and histology of head and salivary apparatus of the predatory bug, Rhynocoris marginatus.

Rhynocoris marginatus Fabricius (Hemiptera: Reduviidae) is an important biological control agent against more than 25 insect pests in India. For a better understanding of the feeding adaptation of this bug, the gross morphology and histology of its head and salivary apparatus were studied using both a light microscope and scanning electron microscope. The head is more or less elongate, mobile, and immersed into the eyes. R. marginatus has a three-segmented curved rostrum; the middle segment is longer than the other two segments. The terminal rostral segment bears spines and trichobothria externally. Stylet bundles bear two pairs of maxillary and mandible stylets in the curved rostrum with serrations. The stylets help to penetrate into the tissue and directly pump the toxic venomous saliva deep into the prey. The principal gland is bi-lobed (anterior lobe and posterior lobe), whereas the accessory gland is uni-lobed, exhibiting distinct functional and histological differences. These glands receive tracheal and nerve supply. Mononucleated, binucleated, trinucleated and polynucleated cells are distributed both in anterior and posterior lobes of the principal gland. The cytoplasm has collecting vacuoles with secretions. Therefore, this predator is highly equipped with well-developed mouthparts that are attached to the salivary apparatus.

Biological, ecological, morphological and cytogenetic analyses, with taxonomic notes of emZelurus/em emochripennis/em (Stål, 1854) (Hemiptera: Heteroptera: Reduviidae: Reduviinae).

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.

Proguithera caspersi/, a new cavernicolous thread-legged bug species (Hemiptera: Heteroptera: Reduviidae: Emesinae) from Hainan, China.

The thread-legged bug genus Proguithera Wygodzinsky, 1966 is recorded from China for the first time, and a new species, P. caspersi sp. nov., is described and illustrated. The new species is cavernicolous and was collected on the walls of a cave together with another emesine species Myiophanes tipulina Reuter, 1881. The type species of the genus, P. drescheri (Wygodzinsky, 1966), is re-diagnosed based on the examination of the holotype. An updated key to species of the Guithera-Lutevula genus group is provided.