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.

Efficient capture of natural history data reveals prey conservatism of cryptic termite predators.

Stenophagy, specialization of a clade on a narrow range of taxa, has not been well studied in speciose clades of predators, principally due to the difficulty of obtaining adequate natural history data. The pantropical Salyavatinae (Hemiptera: Reduviidae; 17 genera, 107 species) contains members with enigmatic morphology and specialized behavior for feeding on termites. All Salyavatinae are suspected specialist termite predators; however, existing observations are limited to seven species. Prior analyses indicate that Salyavatinae may be paraphyletic with respect to another subfamily, Sphaeridopinae, also hypothesized to feed on termites. A molecular phylogeny of these putative termite assassins is here constructed using seven loci from 28 species in nine genera and is used in a dating analysis to shed light on the timing of Neotropical colonization by this primarily Old World clade. DNA extracted from gut contents of 50 individuals was assayed using PCR with prey-specific primers.Molecular assays, along with recent photographs and observations, provide substantial evidence that this clade feeds specifically upon termites, documenting 28 new individual associations. Our phylogeny supports a sister group relationship of the Neotropical genus Salyavata with Sphaeridopinae. Termite association data combined with our phylogeny provide evidence of previously unknown prey conservatism among clades of one of the most diverse groups of specialist termite predators.

Phylogenetics and biogeography of the endemic Madagascan millipede assassin bugs (Hemiptera: Reduviidae: Ectrichodiinae).

For at least the past 80my, Madagascar, a major biodiversity hotspot, has been isolated from all other landmasses. This long-term isolation, along with geologic and climatic factors within Madagascar and throughout the Indian Ocean, has undoubtedly influenced the evolution of the island's biota. However, few systematic analyses incorporating modern divergence dating and biogeographic analyses have focused on Madagascan insects. The diverse Madagascan millipede assassin bugs (Heteroptera: Reduviidae: Ectrichodiinae) offer an opportunity to contribute to a limited body of insect-related research that explores Madagascar's historical biogeography. A molecular dataset (COI mtDNA and 18S, 28S D2 and D3-D5 rDNAs) for 56 taxa (39 ingroup) and a combined morphological (145 characters) and molecular dataset for 110 taxa (93 ingroup) are analyzed with maximum likelihood (ML) and parsimony approaches. Based on the molecular ML phylogeny, divergence times were estimated using fossil and secondary calibrations and biogeographic analyses performed using DIVA, DEC, and DEC+j models to determine the role and patterns of vicariance and dispersal in the origin of Madagascan Ectrichodiinae. Results indicate that Ectrichodiinae in Madagascar do not form a monophyletic group, different clades are closely related to Afrotropical and Oriental lineages, and have colonized the island via transoceanic dispersal at least twice from the Oriental region and once from the Afrotropical region in the last ∼68my. Additionally, the DEC+j and DIVA models infer a single out-of-Madagascar dispersal event to the Afrotropical region. Oceanic and geologic factors that may have facilitated dispersal between these three regions are discussed. Results of the combined analyses are used to explore character support for Madagascan taxa and inform taxonomic diagnoses. Our results are congruent with the small but growing body of biogeographic research supporting Cenozoic transoceanic dispersal for Madagascan invertebrates to and from Oriental and Afrotropical regions.

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.

Triatominae species of Suriname (Heteroptera: Reduviidae) and their role as vectors of Chagas disease.

Nine species of Triatominae, representing three tribes and five genera, are currently known in Suriname. An annotated list of the species based on the collections of the Bureau of Public Health (Suriname), the National Zoological Collection Suriname and the National History Museum Leiden (the Netherlands) is provided. Additionally, the results of several years of opportunistic collection in two domestic environments are presented. The most common species are Rhodnius pictipes Stål, 1972, Rhodnius robustus Larrouse, 1972 and Panstrongylus geniculatus (Latreille, 1811). The significance of the species as vectors of Chagas disease in Suriname is discussed.

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.

Myrmicella, a new genus of Harpactorinae (Hemiptera: Heteroptera: Reduviidae) from Madagascar.

Myrmicella verticospinosa gen. et sp. nov., is described, based on four specimens (two males and two females) collected in south-west Madagascar (Zombitse-Vohibasia National Park and Isalo National Park). Three specimens have been collected by sifting the leaf litter, one female was collected using yellow pan traps. Genitalia of both sexes are described and illustrated.

Chromosomes and their meiotic behavior in twelve species of the subfamily Harpactorinae (Hemiptera: Heteroptera: Reduviidae) from north India.

The chromosome complements and male meiosis of 12 species belonging to the subfamily Harpactorinae from north India have been described for the first time. All the species show twelve pairs of autosomes and X multiplicity with 9 species having X1X2X3Y and 3 species having X1X2Y sex mechanism. The general course of meiosis in all the presently studied species is fairly uniform. Autosomes show a high degree of decondensation and sex chromosomes are condensed during the diffuse stage. Single terminal chiasma per bivalent is seen in all except Sycanus croceovittatus Dohrn, which has two terminal chiasmata in one or two bivalents. A regular arrangement of chromosomes at metaphase I has been observed in 7 species, but in the rest, no definite pattern is recorded. In Euagoras erythrocephala Livingstone and Ravichandran, more than one type of arrangement is seen. At metaphase II, autosomes form a ring in the center of which lies the pseudomultivalent in all the species excepting Rhynocoris costalis (Stål). In Sycanus croceovittatus, a proper pseudotetravalent is lacking. Cytogenetic trends within the subfamily have been discussed in the light of newly added cytogenetic data of 12 species, which may later be used in tracing the evolution of chromosomes in Harpactorinae.

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.