Description of seven new species of Edessa Fabricius, 1803 from Central America (Heteroptera, Pentatomidae, Edessinae).

Not much is known about the Central American stink bug fauna of the Edessinae subfamily. A recent survey of Costa Rican species from this subfamily listed 65 species, of which 47 were new additions. More recently, a description of a group of species of Edessa added more three species to the fauna of Costa Rica. This work is part of a series of articles aiming to review Edessinae from Central America. In this article seven new species are described from Panama and Costa Rica: Edessa bituberculata sp. n., E. bivenulata sp. n., E. enargocelida sp. n., E. fuscolimbata sp. n., E. fuscopunctata sp. n., E. helvoalata sp. n. and E. holochlorata sp. n.

Diagnose of the nominal subgenus of Edessa and descriptionof the E. sexdens group (Heteroptera, Pentatomidae, Edessinae).

The subgenus Edessa, the largest and to date the most confusing taxon of Edessa is diagnosed here, comprising 79 species listed in this work. The subgenus is characterized by the humeral angles conical, somewhat developed, and projected, with apex rounded or somewhat acuminated, but never acute, shyny and impunctate or with a few punctures. The E. sexdens group, composed of 22 species, is proposed here to the subgenus Edessa. This group can be characterized by the anterior arms of the metasternal process with excavated apex; connexival segments with a pair of dark spots; ventral side of the abdomen raised medially; rounded shallow excavation near each spiracle; area between dorsal rim of the pygophore and posterolateral angles grooved; superior process of the genital cup laminar and continued ventrally by a carina, oblique to the parameres; parameres with three lobes; area between dorsal rim of the pygophore and posterolateral angles grooved. The species of the E. sexdens group are distributed in the Neotropical region, from Mexico to Argentina. Descriptions, measurements, and illustrations are provided. An identification key and distributional map are presented. The species Edessa suturata Dallas, 1851 is considered junior synonym of E. alces Erichson, 1848; E. inclyta Walker, 1868, E. fuscidorsata Distant, 1881, and E. gnu Breddin, 1905 are considered junior synonyms of E. electa Walker, 1868; E. olivacea Stål, 1862 and E. ventralis Walker, 1868 are considered junior synonym of E. phoenicopus Dallas, 1851; E. nigromarginata Distant, 1881 is considered junior synonym of E. sexdens Fabricius, 1803; E. subandina Breddin, 1994 is considered a valid species; Edessa dentata (Dallas, 1851) and Edessa excellens (Walker, 1868) are considered junior synonym of E. (E.) urus Erichson, 1848.

Seventeen new Species of Hypoxys Amyot amp; Serville, 1843 (Heteroptera: Pentatomidae: Edessinae).

Hypoxys was recently raised from a subgenus of Edessa to a genus. In the same work Hypoxys was divided in four groups of species: H. quadridens; H. triangularis; H. oxyacanthus and H. balteatus, gathering 17 species transferred from Edessa. The H. balteatus group was recently revised and 15 species were added to this originally monotypic group. Several new species await description in other groups. The H. quadridens group is composed by very similar species; therefore, their main diagnostic features are found in the male genitalia. In this work we describe 17 new species belonging to H. quadridens group: H. angulatus sp. n., H. bitumidus sp. n., H. clavatum sp. n., H. crassimarginatum sp. n., H. cristatus sp. n., H. dorsocarinatus sp. n., H. excavatus sp. n., H. flexuosus sp. n., H. nigrosinuatus sp. n., H. omegoides sp. n., H. pentagonalis sp. n., H. rugulosus sp. n., H. runcinatus sp. n., H. similis sp. n., H. trilobulatus sp. n., H. unguiculatus sp. n., H. ventrotuberculatus sp. n.. Diagnosis, photographs of the external genitalia, body in dorsal and ventral view, and distribution map are provided, including a key to the species of the group.

Three new species of emEurystethus/em from Brazil (Hemiptera: Heteroptera: Pentatomidae).

The Heteroptera (true bugs) and its fourth-largest family Pentatomidae (stink bugs) are worldwide distributed groups with higher diversity in the tropics. The pentatomid subfamily Discocephalinae is almost restricted to the Neotropical Region, and it is divided into the tribes Discocephalini and Ochlerini. Eurystethus Mayr, 1864 is placed within Discocephalini and comprises 18 species, 16 out of them classified into two subgenera. Here we describe Eurystethus jo sp. nov., Eurystethus multipunctatus sp. nov., and Eurystethus rufodorsatus sp. nov., all occurring in Brazil, compare them to other species and provide a distribution map.

A new species group to emEdessa/em, the emE. ovina/em group, with description of a new species (Heteroptera: Pentatomidae: Edessinae) from Brazil.

The E. ovina group of species is proposed here to include Edessa ovina Dallas, 1851 from Trinidad and Tobago, Colombia, Paraguay, Brazil, Argentina (new records) and Guyana; E. impura Bergroth, 1891 from Brazil and Argentina (new record); E. sahlbergii Stål, 1872 restricted to Brazil; and E. graziae sp. nov. from Brazil and Argentina. The E. ovina group can be identified by the apex of the second pair of wings narrowing distally and by a tumid area on the ventral surface of the pygophore (male genitalia) projected posteriorly. Descriptions, measurements, and illustrations of the metasternal process, external and internal genitalia of male and female are provided. A map with the distribution of the species is presented. The holotype of Edessa ovina is designated here. Edessa argali Breddin, 1903 is considered a junior synonym of E. impura. Females of E. ovina and E. impura, and male of E. sahlbergii are described for the first time.

Termitaphididae, a family of true bugs (Heteroptera) new to Brazil, with the redescription of Termitaradus trinidadensis.

The family Termitaphididae and the species Termitaradus trinidadensis (Morrison) are recorded for the first time from Brazil. This species had been found only in Trinidad and Tobago and is redescribed based on specimens collected in Heterotermes tenuis (Hagen) termite nests from forested and open areas in the Brazilian Amazon. Scanning electron micrographs of integumental ornamentation, illustrations of male and female terminalia, and pictures of live specimens are provided for the first time.

Voragocoris weirauchae sp. n. (Heteroptera: Schizopteridae: Schizopterinae), a further minute litter bug species from Brazil.

The genus Voragocoris Weirauch, 2012 is composed of Voragocoris schuhi Weirauch, 2012 and Voragocoris amrishi Makhan, 2013, previously recorded from Peru and Suriname, respectively. In this paper, a new species, Voragocoris weirauchae sp.n., is described based on specimens collected in the Brazilian Amazon, representing the first record of the genus from Brazil. We provide diagnosis, description, photographs of habitus, and scanning electron micrographs of the diagnostic features. A key to species based on males of Voragocoris is also presented.

Revision of Hypoxys Amyot amp; Serville, 1843 stat. rest. (Heteroptera Pentatomidae).

Hypoxys is one of four subgenera of Edessa (Fabricius, 1787) together with Aceratodes (De Geer, 1773), Dorypleura (Lepeletier Serville, 1825) and Edessa (Fabricius, 1787). This taxon has a very confusing taxonomy due to incorrect use of the names available, imprecise descriptions, and large number of very similar species. To illustrate the confusion, Hypoxys potentially includes a large number of species of Edessa, but actually contains only the type species, according to the most recent catalogue. The name E. quadridens Fabricius, 1803, type species of Hypoxys, was used for more than a century as a label to identify a large group of similar species. In this work we continue revising Edessa elevating Hypoxys to genus based on the following morphological characters: proximal part of costal margin black; evaporatorium with a deep notch on lateral margin; anterior arms of metasternal process acuminate; pygophore longer than wide and subrectangular or barrel-shaped in dorsal view; dorsal rim of pygophore well-projected, almost reaching posterolateral angles and finishing in a small lobe; and posterolateral angles of pygophore not developed. Hypoxys is being divided in four species groups to include 17 species transferred from Edessa. H. quadridens group comprises: Hypoxys quadridens (Fabricius, 1803) (type species), H. boerneri (Breddin, 1904) comb. nov., H. claricolor (Breddin, 1907) comb. nov., H. dolosus (Breddin, 1907) comb. nov., H. eburatulus (Breddin, 1907) comb. nov., H. necopinatus (Breddin, 1907) comb. nov., H. offuscatus (Breddin, 1907) comb. nov. and H. trabeculus (Breddin, 1907) comb. nov. H. triangularis group comprises: H. triangularis (Dallas, 1851) comb. nov. (type species), H. capito (Breddin, 1904) comb. nov. and H. subrastratus (Bergroth, 1891) comb. nov. H. oxyacanthus group comprises: H. oxyacanthus (Breddin, 1904) comb. nov. (type species), H. brachyacanthus (Breddin, 1904) comb. nov., H. infulatus (Breddin, 1904) comb. nov., H. leptacanthus (Breddin, 1904) comb. nov. and H. tragelaphus (Breddin, 1903) comb. nov. H. balteatus group comprises only H. balteatus (Walker, 1868) comb. nov. These groups of species will be useful to organize the new species of Hypoxys that will be described in upcoming papers. Descriptions, measurements, photos of genitalia of both sexes; photos in dorsal and ventral views of the species; and distribution maps are provided. Males of H. capito, H. claricolor, H. eburatulus, H. infulatus, and H. subrastratus are described for the first time. Female of H. necopinatus is described for the first time. Edessa jugalis is considered a junior synonym of H. quadridens; E. rimata a junior synonym of H. offuscatus; E. scabriventris and E. leprosula junior synonyms of H. triangularis; E. pachyacantha a junior synonym of H. tragelaphus; E. orba a junior synonym of E. oxyacanthus. A key to the species of Hypoxys is also provided.

Two new species of Hoplonannus (Heteroptera: Schizopteridae: Schizopterinae) from the Brazilian Amazon.

The genus Hoplonannus McAtee Malloch, 1925 is composed of H. brunneus, H. paenebrunneus, H. craneae, and H. australis, all previously recorded from Central and South America. In this paper, we describe Hoplonannus bifidus sp.n. and Hoplonannus robustus sp.n. based on several specimens collected in the Brazilian Amazon, representing the first record of the genus from northern Brazil. We provide diagnoses, photographs of habitus, scanning electron micrographs of male genitalia, and illustrations of female genitalia. A key to species based on males of Hoplonannus is also presented.

A new species of Hypsohapsis (Heteroptera: Schizopteridae: Hypselosomatinae) from the Brazilian Amazon.

The genus Hypsohapsis Hoey-Chamberlain Weirauch was erected based on only one male specimen from southern Brazil (São Paulo). Hypsohapsis amazonica sp.n., described herein from the Brazilian Amazon, is based on a male and two females collected in pristine tropical forest. The female of Hypsohapsis is described for the first time. In addition, an updated key to the genera of Hypselosomatinae is provided.

Stink bugs (Hemiptera: Pentatomidae) from Brazilian Amazon: checklist and new records.

A list of the stink bugs (Pentatomidae) species of the Brazilian Amazon is provided for the first time, based on literature review and identification of the specimens of Pentatomidae deposited in the entomological collection of the Museu Paraense Emílio Goeldi (MPEG). Amazonian data base presented here has 324 species of Pentatomidae belonging to 102 genera, ten tribes and five subfamilies: Asopinae, Cyrtocorinae, Discocephalinae, Edessinae and Pentatominae. The Pentatominae has the largest number of species registered to Brazilian Amazon (144 species). The MPEG entomological collection has 122 species distributed in 53 genera, nine tribes and five subfamilies. Forty species from eight tribes and four subfamilies are recorded for the first time to Brazilian Amazon, including species from Podisus Herrich-Schäffer (Asopinae); Adoxoplatys Breddin, Macropygium Spinola, Neoadoxoplatys Kormilev, Ochlerus Spinola and Platycarenus Fieber (Discocephalinae); Edessa Fabricius (Edessinae); Arocera Spinola, Arvelius Spinola, Banasa Stål, Dichelops Spinola, Euschistus Dallas, Mormidea Amyot Serville, Oebalus Stål, Pellaea Stål, Piezodorus Fieber, Proxys Spinola and Thyanta Stål (Pentatominae). Three of these species-Podisus maculiventris (Say, 1831) Neoadoxoplatys saileri Kormilev, 1956 and Edessa laticornis Stål, 1872-are new records to South America. The number of species of Pentatomidae to Brazilian Amazon represents approximately 50% of the species registered to Brazil. Such list of species is important to better understand the distributional range and diversity of the species of Pentatomidae in the Amazon region.

Revision of Pygoda Amyot Serville, 1843 stat. rest. (Heteroptera: Pentatomidae: Edessinae) with description of four new species.

Edessa is the largest genus of Edessinae and comprises five subgenera: Edessa, Aceratodes, Dorypleura, Hypoxys and Pygoda. Systematics of Edessa is very confusing due to several factors, among them, large number of species and vague diagnosis to the subgenera. In this work we continue revising Edessa elevating Pygoda to genus level based in morphological characters: pronotal angles short and acute. Abdominal segment VII developed posteriorly and at least 1.5 longer than wide at the base. Males with pygophore large in relation to body size when compared with other edessines. Phallus large, constricted distally and showing a remarkable ventral development. Females with capsula seminalis large in comparison with other edessines. Distal part of the ductus receptaculi sclerotized. Pars intermedialis with sclerotized part bigger than membranous part. This genus comprises nine species: the type species Pygoda polyta, four species originally described in Edessa-E. irrorata, E. thoracica, E. expolita and E. civilis, and four new species-P. amianta, P. variegata, P. poecila and P. ramosa. Diagnosis, illustration of the internal genitalia from males and females, photographs of the external genitalia from both sexes, body in dorsal and ventral view, and distribution map are provided. A key to the species of Pygoda is also provided. The lectotypes of Edessa irrorata, Pentatoma polyta, Edessa nervosa, Edessa civilis and Edessa thoracica are here designated.

A new genus and new species of Edessinae (Hemiptera: Heteroptera: Pentatomidae).

The genus Plagaedessa gen. n. is proposed to accommodate four species: Edessa celsa Distant and three new species: P. distanti sp. n., P. nigrovittata sp. n. and P. maranhensis sp. n. The new genus is characterized by the antennae black with the antennomer I showing a yellow spot on ventral surface, legs yellow with femora and tibiae showing two or three longitudinal black stripes, gonapophysis 9 with a sclerotized area bearing a protuberance close to gonocoxites 9 and pygophore with black posterolateral angles strongly developed. Diagnosis, illustrations of the metasternal process and external and internal genitalia, photographs of the dorsal and ventral view, and a distribution map are provided. A key to the genera of Edessinae and subgenera of Edessa, and a key to the species of Plagaedessa are also provided. Male and female genitalia of Plagaedessa celsa n. comb. are described for the first time. Distribution of P. celsa is extended to Costa Rica and Mexico. The lectotype of Edessa celsa is here designated.

Reevaluation of the type species and redescription of five species of Edessa (Heteroptera: Pentatomidae: Edessinae).

In accordance with the rules in the International Code of Zoological Nomenclature, the type species for the genus Edessa Fabricius, 1803 is now recognized to be Cimex antilope Fabricius, 1798 rather than the previously recognized Cimex cervus Fabricius, 1787. Edessa antilope is redescribed, as well as the following four species that have in the past been related or compared to E. antilope: E. arabs (Linnaeus, 1758) from French Guiana, Costa Rica and Panama (new record), E. helix Erichson, 1848 from Costa Rica, Panama, Venezuela, Suriname (new record), Guyana, Brazil (new record), Bolivia (new record) and Argentina (new record), E. ibex Breddin, 1903 from Costa Rica, Panama (new record), Ecuador, Brazil (new record), Peru and Bolivia (new record), and E. taurina Stål, 1862 from Mexico, Guatemala, Belize (new record), Honduras (new record). The distribution of Edessa antilope is expanded to Venezuela. The female of E. antilope and the male of E. taurina are described for the first time. Edessa antilope is removed from the synonymy of E. arabs and reinstated as valid species; additionally, E. costalis Stål, 1872 is removed from the synonym of E. helix and is placed as a junior synonym of E. antilope. Edessa saiga Breddin, 1903 is considered a junior synonym of E. ibex. Lectotypes are designated for all species. A key is provided for the identification of the species.

Description of a new group of species of Edessa (Hemiptera: Pentatomidae: Edessinae).

The new group of species viridula is proposed to five new species: Edessa viridula sp. n., Edessa exigusternata sp. n., Edessa angusticlada sp. n., Edessa rugulata sp. n. and Edessa eucnema sp. n. based on 58 specimens examined. Descriptions of the group and species are provided. Illustrations of the species along with metasternal process and male and female genitalia are provided; an identification key and distributional map are presented.

Chromosomal evolutionary dynamics of four multigene families in Coreidae and Pentatomidae (Heteroptera) true bugs.

Previous chromosome mapping of multigene families in Pentatomomorpha (Heteroptera) insects, which was restricted to the major rDNA, revealed remarkable conservation of number of clusters and chromosomal positions. Aiming to understand the chromosomal organization and evolutionary patterns of multigene families in karyotypes of Heteroptera, we performed a chromosomal mapping using four distinct multigene families in representatives of Coreidae (ten species) and Pentatomidae (five species). A single pair of the major rDNA cluster (18S rDNA probe) and a single pair of the minor rDNA cluster (5S rDNA probe), both terminally located were primarily observed, being, in most species, located in distinct chromosomes. However, some alternative patterns were also observed. In species in which the U2 snDNA and H4 gene clusters were mapped, they were mainly located in one autosomal pair each, wherein the H4 gene cluster was located in different positions. Our data suggest that the karyotype diversity reported in Coreidae is not reflected in the distribution diversity of multigene families. This contrasts with the data for Pentatomidae, with a conserved gross karyotype but a discrete diversity in the location of the clusters of multigene families, indicating genome dynamics for these markers. The findings are discussed to shed light on the possible causes for the conservation or variation observed and to assist in understanding the chromosomal evolutionary trends in the group.

New evidences supporting trophobiosis between populations of Edessa rufomarginata (Heteroptera: Pentatomidae) and Camponotus (Hymenoptera: Formicidae) ants

ABSTRACT Despite its important effect on the maintenance of tritrophic interactions among plants, insect herbivores, and ants, there is still a paucity of natural history and basic biology information involving trophobiosis among Heteroptera stink bugs. Here, based on previous observations of a new trophobiotic interaction between Edessa rufomarginata (De Geer, 1773) and Camponotus rufipes (Fabricius, 1775) ants, we describe the chemical profile of the honeydew obtained by Gas Chromatography–Mass Spectrometry. There were mainly three different sugars (trehalose, glucose, and sorbose) within our samples. The extrafloral nectaries of Caryocar brasiliense Camb., the host plant of E. rufomarginata, attracts a wide assemblage of Cerrado ants with varying aggressiveness toward herbivores. Therefore, this facultative trophobiotic interaction may allow the survival of the stink bug while feeding on the risky, highly ant-visited plant. Given the rarity of trophobiotic interactions between Pentatomidae species and ants and considering a zoological perspective within this family, here we discuss the ecological and evolutionary routes that may allow the rise of these interactions.

Analysis of the karyotype structure in Ricolla quadrispinosa (Linneus, 1767): inferences about the chromosomal evolution of the tribes of Harpactorinae (Heteroptera, Reduviidae).

The subfamily Harpactorinae is composed of six tribes. Phylogenetic studies bring together some of Harpactorinae tribes, but by and large the data on evolutionary relationships of the subfamily are scarce. Chromosome studies are of great importance for understanding the systematics of different groups of insects. For Harpactorinae, these studies are restricted to some subfamilies and involved only conventional chromosome analysis. This work analyzed cytogenetically Ricolla quadrispinosa (Linneus, 1767). The chromosome number was determined as 2n = 24 + X1X2Y in males. In metaphase II the autosomal chromosomes were organized in a ring with the pseudo-trivalent of sex chromosomes in its center. After C-banding followed by staining with DAPI, AT-rich blocks in autosomes were observed and the negatively heteropycnotic sex chromosomes. The data obtained, together with existing data for other species of the group, indicated that different chromosomal rearrangements are involved in the evolution of the species. In addition, a proposal of karyotype evolution for the subfamily, based on existing phylogenetic studies for the group is presented.

New species of Edessa Fabricius, 1803 (Hemiptera: Pentatomidae) from Costa Rica.

The edessines from Costa Rica are little known; only 18 species have been registered or described from this country so far. Mainly based in a large sample from Instituto Nacional de Biodiverdidad (INBio), Costa Rica, we decided to update the information concerning Edessinae from Costa Rica. We present a list of species from Costa Rica raising the number of known species from Costa Rica to 65. We are also describing nine new species: Edessa bella Fernandes & Silva, E. bruneolineata Fernandes & Correia, E. curvata Fernandes & Nunes, E. lewisi Fernandes & Silva, E. nigroangulata Fernandes & Silva, E. osae Fernandes & Nunes, E. oxcarti Fernandes & Correia, E. pallidoangulata Fernandes & Nunes and E. puravida Fernandes & Correia. Species were described, illustrated and photographed. Distribution maps for the species are also provided.

Paraedessa, a new genus of Edessinae (Hemiptera: Heteroptera: Pentatomidae).

The new genus Paraedessa is proposed, to include four species previously placed in Edessa. The new genus is characterized by males with the pygophore showing a lateral expansion of proctiger, and females with gonocoxites 8 reduced; gonapophyses 8 exposed and sclerotized. The species are very similar and their identification is possible only by comparing genitalia. New combinations proposed are Paraedessa stolida (Linnaeus) from Suriname and Brazil; P. heymonsi (Breddin) from Peru and Bolivia; P verhoeffi (Breddin) from Venezuela, Colombia, Peru and Brazil and P paravinula (Barber) from Puerto Rico. New species proposed are P. albomaculata from Suriname; P. cyanodorsata from Costa Rica, Panama and Venezuela; P. ecuadoriensis from Colombia and Ecuador; P. silvicola from French Guiana; P. subretangulata from Nicaragua, Costa Rica, Colombia and Venezuela. The species were illustrated and photographed. Descriptions were made using mostly male and female genitalia external morphological characters. A key and distribution map for the species are also provided.