External egg structure of the Pentatomidae (Hemiptera: Heteroptera) and the search for characters with phylogenetic importance.

The chorionic structure of sixteen pentatomid species is described. Morphological patterns in different taxonomic levels are discussed. In addition, egg characters useful in cladistic analyses are listed, and some of those characters are tested for congruence with two cladistic analyses previously conducted within Pentatomidae. Descriptive studies were conducted with Banasa induta, Capivaccius bufo, Catulona pensa, Chinavia armigera, Chinavia aseada, Chinavia brasicola, Chinavia runaspis, Dichelops furcatus, Euschistus heros, Euschistus riograndensis, Euschistus paranticus, Mormidea cornicollis, Podisus distinctus, Podisus nigrispinus, Serdia apicicornis, and Thoreyella maracaja. The eggs were examined and photographed under light and scanning electron microscopy. Based on literature data, a list of 40 egg characters with potential phylogenetic importance has been compiled. Some of these characters were included in the cladistic analyses of the genus Nezara (six characters) and of the Chinavia obstinata group (five characters). Both analyses were performed in TNT with equal weighting of characters. The eggs of most of the Neotropical species studied were barrel-shaped, chorion translucent and spinose, with aero-micropylar processes that were short and clubbed. The patterns of egg morphology could be identified in Carpocorini, Procleticini, and in the genera Banasa, Chinavia, Euschistus, Mormidea, and Podisus. In the cladistic analyses, the inclusion of egg characters did not affect the topology of the trees shown in the original papers. For the analyses, the egg characters were somewhat informative. At present, a total of 286 Pentatomidae species have their egg stage described. 

Exploring Phylogenetic Relationships of the Mecocephala Group (Hemiptera: Pentatomidae) Using Continuous and Discrete Characters, with Notes on Taxonomy.

The Mecocephala group comprises about 50 species, restricted to the Neotropics and with the highest species richness in Neotropical South America. Several species use rice as host plants and their identification is facilitated by the presence of exaggerated head proportions and a unique male genitalic morphology. The taxonomy of the group has been extensively explored, but inferring its monophyly and especially its internal phylogenetic relationships has been challenging. Here, we inferred the phylogenetic relationships for the group assembling the most complete taxonomic sampling to date, analyzing discrete and continuous morphological characters through equal and implied weighted parsimony analyses. The monophyly of the group was recovered, but internal relationships varied slightly according to the dataset tested. Thus, we propose internal arrangements for the group and provide a formal description of the Mecocephala group, diagnoses for each genus, a dichotomous key to identify its genera, and illustrations of the morphological characters and type species.

Antiteuchus tripterus (Fabricius, 1787) (Heteroptera: Pentatomidae): First Record of This Native Stink Bug Damaging the Threatened Araucaria angustifolia in Brazil.

This is the first report of Antiteuchus tripterus (Fabricius, 1787) (Heteroptera: Pentatomidae) damaging Araucaria angustifolia (Bertol.) Kuntze, 1898 (Pinales: Araucariaceae) plants in Brazil. This bug, native to the neotropical region, damaged this plant on fragments of mixed rain forest in Bueno Brandão, Minas Gerais state, Brazil. Araucaria angustifolia, native and with ecological and economic values, was widely used in the lumber market and, therefore, threatened with extinction. Nymphs and adults of A. tripterus were observed in March and April with a population reduction until June 2022. The bioecology and damage caused by A. tripterus on A. angustifolia are not known.

Revision of the genus Alkindus Distant (Hemiptera: Heteroptera: Thyreocoridae: Corimelaeninae).

The neotropical genus Alkindus Distant is revised based on morphological characters (general morphology, including the external scent efferent system and leg structures, and external genital morphology). The male of Alkindus crassicosta Horvath is here described for the first time. Illustrations, an adapted key to species, and a compiled list of plants associated with both species are provided. Distribution records are expanded to include Guatemala and Brazil (Roraima) for Alkindus atratus Distant and Brazil (Santa Catarina) for A. crassicosta.

Contributions to the knowledge of Dichelops Spinola: description of a new species of Dichelops (Diceraeus) and of the male of Dichelops (Prodichelops) divisus (Hemiptera: Heteroptera: Pentatomidae: Pentatominae: Carpocorini).

A new species of Dichelops (Diceraeus) Dallas and the so-far unknown male of Dichelops (Prodichelops) divisus (Walker, 1867) are described, based upon morphological characters. D. (Dice.) caatinguensis sp. nov. is compared to the other four species of Diceraeus. Revised keys to separate the subgenera of Dichelops and the species of Diceraeus are also presented. D. (P.) divisus male is described and compared to the type species. Comparative illustrations of external and internal genitalia of the species are provided.

Diversity of stink bugs (Hemiptera: Heteroptera: Pentatomoidea) in the Brazilian Pampa and the Parque Estadual de Itapuã, Rio Grande do Sul.

Biological surveys represent important contributions to the knowledge of species diversity, conservation, taxonomy, and biogeography. Few surveys of stink bugs and allied groups (Pentatomoidea) were produced in Brazil, especially regarding the Brazilian Pampa, an overlooked biome. Here, a list of Pentatomoidea species of the Brazilian Pampa is presented for the first time, composed of seven families and 152 species. The results of five years of sampling in the Parque Estadual de Itapuã (PEI) are also presented. A total of 693 individuals were sampled, representing 41 species, 29 genera, and five families of Pentatomoidea. Pentatomidae presented a higher richness (28 species), followed by Cydnidae, Scutelleridae, Dinidoridae, and Megarididae. The most abundant species collected in the PEI was Brachystetus geniculatus (Fabricius) sampled in Bromelia balansae Mez (32.76%), followed by Oebalus poecilus (Dallas) found in Persicaria hydropiper (Linnaeus) (11.99%), Pallantia macula (Dallas) collected in B. balansae (10.92%), Dinidor saucius Stål (8.57%) in Smilax cf. campestris Griseb., and Caonabo pseudoscylax (Bergroth) (5.35%) found in Homolepis glutinosa (Sw.) Zuloaga & Soderstrom. This is the first time that a list of species is presented for the Brazilian Pampa and Parque Estadual de Itapuã, providing a baseline to develop further studies with Pentatomoidea in the biome.

Occurrence of the Yellow-Spotted Stink Bug Erthesina fullo (Thunberg) (Hemiptera: Pentatomidae) in Brazil, a Polyphagous Species from Asia.

In recent years, the management of invasive stink bugs (Pentatomidae) has been a challenge in many regions of the world, including the Neotropical, where four non-native species were detected in the last two decades. Species with invasive potential include the yellow-spotted stink bug, Erthesina fullo (Thunberg). It is polyphagous and is known to cause damages to fruit crops in Southeast Asia. Herein, we report for the first time the occurrence of E. fullo in Brazil (Santos, São Paulo). Nymphs and adults have been recorded nearby the Harbor of Santos since November 2020. A fifth instar nymph was collected and reared until adulthood. The insect was found feeding on Inga sp. leaves (Fabaceae), a plant restricted to the Neotropical Region. The detection of multiple specimens in Santos, plus their ability to feed on native and or on cultivated plants, may represent a threat to the Brazilian agribusiness and environment.

Revisiting Stl's thoughts: formalizing the ictericus group in Euschistus (Euschistus) (Hemiptera: Heteroptera: Pentatomidae).

Euschistus Dallas is distributed throughout the Western Hemisphere and is composed of 67 species in three subgenera: Euschistomorphus Jensen-Haarup, Lycipta Stl, and the nominate subgenus. Euschistus (Euschistus) includes several economically important species. Due to the lack of a phylogenetic analysis for the subgenus Euschistus, its groupings are based primarily on male genitalia. In this paper, the following Nearctic taxa that were first grouped together by Stl (1872), namely E. ictericus (Linnaeus), E. variolarius (Palisot de Beauvois), E. servus servus (Say), E. servus euschistoides (Vollenhoven), and E. tristigmus tristigmus (Say), are reviewed and redescribed. Several other similar, and probably related, taxa are also included in this study: E. tristigmus luridus (Dallas), E. inflatus Van Duzee, and E. latimarginatus Zimmer. Morphological patterns pertaining to the internal and external male and female genitalia allowed us to formalize the ictericus group to include the above-mentioned species (except E. tristigmus). The morphological overlap between the subspecies and sympatric areas of distribution have led us to consider E. servus euschistoides as a junior synonym of E. servus servus, and to corroborate the synonymy first proposed by Uhler (1861) of E. tristigmus luridus under E. tristigmus tristigmus.

The eggs and nymphs of predatory stink bugs (Hemiptera: Pentatomidae: Asopinae): what do we know?

The Asopinae are known for their predatory behavior, differing from the phytophagous habits of most pentatomoids, feeding mostly on soft body insects such as larvae of Lepidoptera and Coleoptera. For this reason, asopines have been studied as biological controllers in integrated pest management programs. Notwithstanding their clear relevance, the general knowledge about Asopinae has important gaps, especially regarding immature. Thus, the importance of studying eggs and nymphs of true bugs (Heteroptera) is evident, contributing to understand their classification, biology, and evolution. In this perspective, we conducted a research about immature of predatory stink bugs, highlighting critical features for identification. We present: (1) a literature overview about eggs and nymphs of predatory stink bugs guided by selected categories; (2) images of females laying eggs of ten species and nymphs of thirty-four species, obtained on websites with a citizen science approach; (3) a comparative morphology of immature of six species reared under laboratory conditions, which we examined using light and scanning electron microscopy. We found a remarkable morphological diversity of both eggs and nymphs of Asopinae, revealing key features to establish diagnoses for identification and potential characters to phylogenetics, such as the aero-micropylar processes and chorion scultpturing of the eggs; and the coloration, labium and abdominal plates morphology of nymphs. The results show that little is known about Asopinae immature considering the diversity of the group; however, information obtained by citizen science initiatives, for instance, can improve this knowledge.

Stink bugs (Hemiptera: Heteroptera: Pentatomidae) of Colombia: An annotated checklist of species.

A checklist of 5 subfamilies, 108 genera, and 246 species of stinks bugs (Hemiptera, Pentatomidae) from Colombia is provided. Three genera [Andrallus Bergroth, Placocoris Mayr, and Pseudobebaeus Distant], and eight species [Arocera spectabilis (Drury), Andrallus spinidens (Fabricius), Banasa excavata Thomas, Banasa saileri Thomas, Banasa varians Stl, Chinavia scutellata (Distant), Pelidnocoris haglundi Ruckes, and Pseudobebaeus truncatus (Fallou)] are recorded for the first time from Colombia. Images of dorsal external habitus for all genera that occur in Colombia are offered. This is the first time a comprehensive list of the stink bugs from Colombia is presented, providing a baseline to develop further studies in this group of true bugs.

Taxonomic updates on the Mecocephala group (Hemiptera: Pentatomidae): Redescription of 'Hypatropis complex', with a review of genitalic terminology and new records.

The taxonomic revision of 'Hypatropis complex', composed by Amauromelpia Fernandes Grazia, Hypatropis Bergroth, Luridocimex Grazia, Fernandes Schwertner, Stysiana Grazia, Fernandes Schwertner, and Pedinonotus Fernandes Grazia, updating the genitalic terminology, are presented here. The differential diagnosis of 'Hypatropis complex' and all of its members are provided. Photographs of the type-specimens and genitalia, species distributional map with new records, and a key to genera and species are also provided.

Redescription of Steleocoris Mayr, 1864 and Theloris Thunberg, 1783 (Hemiptera: Pentatomidae), two poorly known African stink bugs.

The knowledge about the monotypic genera Steleocoris Mayr and Theloris Thunberg is restricted to their original descriptions and subsequent nomenclatural acts. These genera are primarily distributed in South Africa and are two of the few genera of Carpocorini (Pentatomidae: Pentatominae) occurring in Africa. Here, we redescribe Steleocoris comma (Thunberg) and Theloris costata (Thunberg) and illustrate the genitalia of both sexes for the first time. Also, an occurrence map is provided based on collection labels from analyzed specimens and records available on GBIF and the web repository iNaturalist.org.

Finding Padaeus bovillus (Hemiptera: Pentatomidae): A Phylogenetic Placement and the Description of Its Sister Species.

The pentatomids (Hemiptera: Heteroptera) are the third most speciose family within the Heteroptera or the true bugs. The family occurs worldwide and comprises around five thousand valid species within 950 genera. Padaeus Stål belongs to a complex of other genera of Carpocorini Mulsant and Rey related to Euschistus Dallas. These genera present similarities in color, size, and shape, and share common features. However, among its four congeneric species, Padaeus bovillus Distant has been highlighted as an outlier by the posterior margins of the bucculae evanescent, while its congeneric species present posterior margins of the bucculae lobed. Thus, herein we redescribe P. bovillus and present a hypothesis regarding its phylogenetic placement within the Carpocorini. Furthermore, a new species similar to P. bovillus is described. Four molecular markers (COI, CytB, 16S, and 28S) plus 86 morphological characters were used to infer the phylogeny under Maximum likelihood and Bayesian inference. For the descriptions, we measured 16 morphometric parameters and dissected the genitalic structures. We also include illustrations of the habitus, internal and external genitalic structures, and provide distribution maps. Mitripus seclusus sp. n. Bianchi, Krein, Rider, and Grazia is recovered as the sister species to Mitripus bovillus comb. n., and both within Mitripus Rolston. Among other shared characters, species of Mitripus have the femora unarmed, they have a macula near the apex of the radial vein, and the mandibular plates tapering apically. Mitripus bovillus comb. n. and Mitripus seclusus sp. n. have the posterior margin of the pygophore projecting as a spine, a unique pattern within the genus. According to our results, Mitripus including M. bovillus comb. n. and Mitripus seclusus sp. n. now includes five species.

Delicate and diverse: A taxonomic monograph with a phylogenetic analysis of the Neotropical genus Ghilianella Spinola (Hemiptera: Reduviidae: Emesinae).

The Neotropical thread-legged bug genus Ghilianella Spinola, 1850 is the most diversified within Metapterini. A taxonomic revision of Ghilianella is presented, in which seventy seven species are recognized as valid, with twenty-one described as new: Ghilianella berengeri sp. nov.; Ghilianella bifurcata sp. nov.; Ghilianella bolivari sp. nov.; Ghilianella caldensis sp. nov.; Ghilianella dilatata sp. nov.; Ghilianella embera sp. nov.; Ghilianella fernandezi sp. nov.; Ghilianella ferruginosa sp. nov.; Ghilianella gilsantanai sp. nov.; Ghilianella goliath sp. nov.; Ghilianella gracilis sp. nov.; Ghilianella huaorani sp. nov.; Ghilianella jaguar sp. nov.; Ghilianella laticauda sp. nov.; Ghilianella maricruzae sp. nov.; Ghilianella quimbaya sp. nov.; Ghilianella scimitarra sp. nov.; Ghilianella tica sp. nov.; Ghilianella urbanoi sp. nov.; Ghilianella ventrimaculata sp. nov.; and Ghilianella weirauchae sp. nov. For the first time a female specimen is described for Ghilianella atriclava Bergroth, 1911, Ghilianella colona McAtee Malloch, 1925 and Ghilianella pachitea McAtee Malloch, 1925. Three new synonyms are recognized: Ghilianella bulbifera Champion, 1898 (=Ghilianella pendula McAtee Malloch, 1925 syn. nov.; Ghilianella inflata Maldonado, 1981 syn. nov.) and Ghilianella strigata McAtee Malloch, 1925 (=Ghilianella fenestrata Maldonado, 1960 syn. nov.). Eleven species are considered nomina dubia and one species nomen nudum. A key to species and digital images of the external morphology and genitalic structures for each species are provided. Additionally, we offer the first phylogenetic hypothesis of relationships within Ghilianella, using cladistic methods. Based on the phylogenetic results we dismiss all subgenera in Ghilianella and discuss the complex evolution of the abdominal expansions.

Taxonomic review of Sibaria Stål, 1872 (Heteroptera: Pentatomidae: Pentatominae: Carpocorini), with description of a new species.

Sibaria Stål is a genus of Pentatomidae occurring exclusively in the Neotropical region. Since its description, little information has been added to the knowledge of the taxon. The hypothesis of monophyly of Sibaria was recently supported by a phylogenetic analysis based on morphological and molecular data. The purpose of this study is to provide a review of Sibaria, redescribe the valid species, and describe a new species. The genus includes four species: S. andicola Breddin, S. armata (Dallas), S. englemani Rolston, and S. amazonica sp. nov.. Diagnoses, macrophotographs (habitus, external and internal genitalia), an identification key, and distribution records of the species are also provided.

Taxonomy of widespread Neotropical species of Podisus Herrich-Schäffer (Hemiptera: Pentatomidae: Asopinae): redescription of P. distinctus (Stål, 1860) and revalidation of P. fuscescens (Dallas, 1851).

Some species of Podisus Herrich-Schäffer have a confuse taxonomic history with several modifications and loss of information over time. The genus has been considered a taxonomic dumping ground in Asopinae, and its current species composition is doubtful. After examining the type specimens of Telepta distincta Stål, Arma submarginata Walker, Arma monospila Walker, Arma fuscescens Dallas, Arma caliginosa Walker, Arma turbida Walker and Podisus mellipes Bergroth, we redescribe and revalidate A. fuscescens as Podisus fuscescens rev. stat., previously considered a junior synonym of Podisus sagitta (Fabricius). The taxonomic status, morphology and distribution of P. fuscescens and P. distinctus are updated and illustrated. Arma caliginosa, A. turbida and Podisus mellipes become synonyms of P. fuscescens. Additionally, Arma submarginata Walker is removed from its synonymy with Podisus ventralis (Dallas) and treated as a synonym of P. distinctus. Podisus signoreti Fallou is removed from its synonymy with P. distinctus and treated as species inquirenda.

New species and new distributional records of Elanela Rolston, 1980 (Heteroptera: Pentatomidae).

Two new species are described, Elanela colombiana sp. nov. from Colombia, with disk of scutellum callused, ivory and abdominal sternite VIII immaculate medially, and Elanela ecuatoriana sp. nov., from Ecuador, distinguished by legs with black spots and the dorsal rim and ventral rim of pygophore with a medial process. Illustrations of habitus and genitalia of each species are provided and the distributional map was updated to include the new species and a new record of E. jordi. Additionally, the identification key to the species of Elanela is updated to include the new species.

Taxonomic modifications in Tylospilus Stål (Hemiptera: Pentatomidae: Asopinae): redescription of T. nigrobinotatus (Berg), description of a new species and an updated identification key for the genus.

The systematics of the New World Asopinae genera is still not completely understood, thus hampering their study in many areas of biological science. Tylospilus Stål is one of the less known genera among New World predatory stink bugs, despite its common occurrence and potential as a biological control agent in crops. Here, we make a contribution to the knowledge of Tylospilus; the species Tylospilus nigrobinotatus (Berg, 1879) is redescribed, and based upon the examination of type specimens, T. armatus Thomas, 1992 is considered to be a junior synonym of the above species. Furthermore, a new species is herein described, and an updated identification key for the species of Tylospilus is provided.

External morphology of the immature stages of Runibia perspicua (Fabricius, 1798) (Hemiptera: Heteroptera: Pentatomidae).

The study of immature stages in Pentatomidae contributes to the knowledge in several areas, mainly systematics, ecology and agriculture. In faunistic inventories, nymphs comprise a significant part of the specimens collected, which often cannot be identified due to lack of knowledge about the immature. Here, we describe the external morphology of eggs and nymphal stages of Runibia perspicua (Fabricius, 1798). The egg is barrel-shaped with the surface of chorium covered by spines, forming polygonal markings interconnected by sheets, similar as observed in some species of Carpocorini. In all the nymphal instars the abdomen is characterized by having rounded reddish spots and an orange/yellow longitudinal macula in the middle of each abdominal tergite, from abdominal segment II to VII. The first instar is the most distinctive, standing out mainly by the head and thorax totally brown. From the second instar the head and thorax acquire pale color with irregular brown spots. The labium presents significant variations in size along the ontogenetic development showing a non-isometric pattern, when compared to the total length of the body. Illustrations and photographs of all the immature stages and the detailed morphology of some structures in SEM are provided.

Revision of Tynacantha Dallas with description of two new species (Hemiptera: Heteroptera: Pentatomidae: Asopinae).

Here, Tynacantha Dallas is revised and two new species are described from Brazil: T. cuprea Brugnera Grazia sp. nov. and T. umeridenigrata Brugnera Grazia sp. nov. The type material is analyzed, figures, illustrations, a distributional map and a key to the species are provided. The color variation found in T. marginata is also discussed.