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External morphology of the egg and the first and fifth instars of Cyrtocoris egeris Packauskas & Schaefer (Hemiptera: Heteroptera: Pentatomidae: Cyrtocorinae)
Abstract
Cyrtocorinae is an uncommon, small, and exclusively Neotropical group in Pentatomidae, whose immatures are poorly understood. In this paper, the egg and first and fifth instars of Cyrtocoris egeris Packauskas & Schaefer are studied with the scanning electron microscopy (SEM). Specimens were collected in Maquiné, Rio Grande do Sul, Brazil. Immatures were analyzed with light stereomicroscope and SEM. The egg's chorion surface is predominantly smooth alternated with granulated areas, and an average of 54 clubbed aero-micropylar processes are arranged in three irregular rows. In the first and fifth instars, organization of the external dorso-abdominal scent efferent system suggests an analogy to the metatho-racic external scent efferent system of the adult, because of structures similar to the evaporatorium, evaporatory channel, and auricle peritreme; the last structure is absent in the first instar. Abdominal sterna III–VII have 1+1 (first instar) and 2+2 trichobothria (fifth instar). Ultrastructural observations allowed recognition of unique characters at the egg stage and conserved features at the nymphal stage in C. egeris.
... Investigation of the immature stages of D. silphoides has also become very important for the same reason. The study of eggs indicated that the white one is a healthy egg that acquires red spots before hatching, while black eggs indicate (Peredo, 2002;Williams et al., 2005;Bianchi et al., 2011) and in Scutellera perplexa (Westwood), ...
... The stridulatory projections detected on the abdominal sterna in nymphs and adults have also been described in the scutellerid subfamilies Hoteinae, Odontotarsinae and Pachycorinae (Carapezza, 2009;Bianchi et al., 2011). The stridulitrum is important for the production of vibrational signals that allow for communication especially among females and early nymphs, as is evident in Pachycoris klugii and P. stalii both of which exhibit maternal care (Peredo, 2002;Williams et al., 2005). ...
... A reticulated pattern is probably more effective for the evaporation of a secretion (Vilímová and Kutalová, 2012). Different reticulated sculpturing is detected on the mesial and lateral portions of the evaporatorium of DAG 2,3, and this was also observed in Galeacius martini Schouteden (Scutelleridae) by Bianchi et al. (2011). DAG 1 secretes non-toxic products to induce aggregation behavior, while DAGs 2 and 3 secrete alarm and defence pheromones (Calam and Scott, 1969). ...
Polychromatism in adult Deroplax silphoides (heteroptera: scutelleridae) and a morphological examination of the immature stages, Saudi Journal of Biological Sciences (2022), doi: https://doi. Polychromatismin adult Deroplax silphoides (Heteroptera: Scutelleridae) and a morphological examination of the immature stages Abstract Polychromatism is a very common phenomenon in several insect orders. Adults of Deroplax silphoides show color variation in their bodies. In such cases, the identification of the immature stages is challenging and misleading. The field collection of adults and immature stages was conducted on the campus of Ain Shams University during the active period of the insect in 2019. Nineteen pronotal and scutellar macular patterns in addition to fifteen patterns of abdominal sternal variation in the color variation and size of the stridulatory area in D. silphoides adult were reported for the first time. In addition, the variation was observed in the shape, size and placement of eggs masses on Dodonaea viscosa commonly called 'sticky hop bush'. The main morphological features and an identification key for the five nymphal instars of this species are also provided for the first time through this work. The proportions of antennal and rostral segments and the degree of scutellum and wing pad development provide excellent characters for distinguishing the five nymphal instars. This work forms a basis for further investigation of such diversity at the molecular level and provides an insight into this phenomenon in the fascinating insect world.
... The most comprehensive phylogenetic hypothesis in terms of taxa coverage (Grazia et al., 2008) places Cyrtocorinae as sister-group to all remaining pentatomids, while Pentatominae, Edessinae, Discocephalinae, and Asopinae are in a polytomy. Since the morphology of pentatomid nymphal SES is highly variable (e.g., Bianchi et al., 2011b;Biasotto et al., 2013;Bottega et al., 2015), and a robust phylogenetic hypothesis for the family is lacking, we investigate whether there is a correspondence between SES morphology and classification within Pentatomidae. In this study, we compare the external scent efferent system of selected species representing different subfamilies of Pentatomidae, also including samples of other pentatomoideans, in order to assess the morphological diversity and bringing light to future phylogenetic studies in different taxonomic levels of Pentatomoidea. ...
... Observation and evaluation of SES were made in resting position and in ostiolar plan following Bottega et al. (2015). Terminology follows Kment and Vilímová (2010a) (as proposed by Bianchi et al., 2011b) and Vilímová and Kutalová (2012). Description of the external scent efferent system of the DAGs, as SES throughout the text, follows Bottega et al. (2015). ...
... Kment and Vilímová (2010a) defined six basic peritremal shapes for the metathoracic scent efferent system but highlighted that, in some cases, transitional kinds between basic shapes could be found. Bianchi et al. (2011b) proposed using the definitions by Kment and Vilímová (2010a) for analogous structures of SES. Despite variations observed, only the spout peritreme seems to occur in Pentatomoidea. ...
... The study of the immature stages is highly valuable in insect phylogeny and cladistic analysis, as well to the recognition and control of pest species (Brailovsky et al. 1992;Costa et al. 2006). Eggs and/or nymphs of 146 species of the Pentatomidae have already been studied, although eggs of only 57 pentatomid species have been investigated by scanning electron microscopy (Matesco et al. 2009;Campos et al. 2010;Candan & Suludere 2010;Lopes & Cervantes 2010;Bianchi et al. 2011;Bundy & McPherson 2011). ...
... Head width measures allow the differentiation of instars (Matesco et al. 2007), and are presented in a separate table (Table 2). Terminology follows Matesco et al. (2009) for eggs, Matesco et al. (2006Matesco et al. ( , 2007 for nymphs, and Kment and Vilímova (2010) adapted by Bianchi et al. (2011) for abdominal external scent efferent system. ...
The study of immature stages is important in phylogeny, as well to the recognition and control of pest species.Glyphe-pomis Berg is included in the tribe Carpocorini.This paper describes the morphology of eggs and nymphs of Glyphepomis spinosa Campos & Grazia.Adult specimens were collected on rice plants in Formoso do Araguaia, Tocantins, Brazil, and they were reared in the laboratory.The immatures obtained were analyzed with a light stereomicroscope and scanning electron microscope.Eggs and nymphs were measured and illustrated and/or photographed.Eggs are barrel-shaped, light green; operculum round and convex; chorion translucent, bright and granulated; aero-micropylar processes clubbed, 68 on average.All instars have oval body and obtuse tylus, surpassing jugae.In the 1st instar, head, thorax and abdominal plates are black, and abdomen is reddish brown; punctures are absent.From 2nd instar on, head and thorax are grayish with dark punctures, and the abdomen is whitish with dark-red punctures.In relation to other Carpocorini species that occur on rice, the eggs of G.spinosa resemble those of Oebalus poecilus (Dallas), a major rice pest in South America.However, nymphs of both species can be easily differentiated by their general color: in O.poecilus all instars have a dark head and thorax and red abdomen.
... Although there are many studies on the morphology and surface structure of eggs of various Heteroptera species, many taxonomic groups still need research [5][6][7][8][9][10][11][12][13][14][15][16][17][18][19][20][21][22][23][24]. ...
In this study, spermatheca and eggs of Pyrrhocoris apterus (Heteroptera: Pyrrhocoridae were examined by light microscope and SEM. The eggs have ~1.30 mm length and 0.94 mm width. Newly laid eggs turn from white to yellow as soon as embryonic differentiation. P. apterus egg surface shows polygonal (usually hexagonal and pentagonal) shapes. The eggs have pipe shaped 6-7 micropylar projections. The egg-burster which is a dark T-shaped or triangular pattern, explosive becomes visible when the embryo is well developed. The nimph, which has completed its embryonic development, comes out of the egg with peristaltic movements. The spermatheca of P. apterus consists of a bulb, a pump, a flange of pump, and duct. The morphology of the egg and spermatheca is useful for classification, because they show a great diversity among species and genera.
... The terminology used for the sensilla follows Altner and Prillinger (1980), Zacharuk (1980), Brozek and Chłond (2010) and Shields (2010), and for SES it follows Kment and Vilímov a (2010a, b) (as proposed by Bianchi et al., 2011), Vilímov a and Kutalov a (2012) and Bianchi et al. (2016). ...
Heteropterans communicate chemically through thoracic or abdominal glandular complexes. The dorso-abdominal scent glands (DAGs) are externalized by cuticular specialized structures forming the external scent efferent system (SES). Different groups in Heteroptera present other glands, such as the Asopinae (Pentatomidae), whose males can have ventral abdominal pheromonal glands externalized by cuticular modifications, called glandular patches (GPs). Here we describe the SES of DAGs and, for the first time,
the GPs of 22 Asopinae genera. The ostioles of SES 1 vary in shape. SES 2 and SES 3 were restricted to scars, indicating that only the SES 1 remains functional in adults. The GPs are found mostly in segments V and VI. We have not found any difference in the morphological pattern between the SES of species bearing GPs and those lacking it, meaning that the acquisition of GPs is likely related to some sexual behavior, thus not as a complex structure replacing any role of the DAGs. The cuticle of the GPs is microsculptured, bearing many pores surrounding the base of sensilla being the pathway for the secretions to reach the outside of the exoskeleton. The sensilla may be adaptations to increase the contact surface, assisting rapid evaporation of the volatile secretions from the pores.
... There has been an immense contribution of many authors that have enriched the knowledge about morphology, histology, biological functions and components of the scent glands (Remold 1962a(Remold ,1962bCarayon 1971;Aldrich et al.1972;Qadir I, et al., 2015). Recent studies regarding morphology utilize examining electron microscopy that permit more comprehensive portrayals of structures in detail (e.g., Bianchi et al. 2011, Pollo et al. 2012, Gilio-Dias et al. 2013. Thereafter, the process of scrutinizing the exocrine secretions for their associated structures and functions continue till date and. ...
Morphology of the scent (pheromone) glands is the first and basic step in understanding the mechanism of working of this highly evolved and integrated system of communication in insects. Leptocoris augur contains a pair of large and distinct symmetric exocrine dorsal abdominal glands (DAGs) found in the dorsal abdominal segments. The glands are simple, voluminous, sac-like and bright orange that turn brown and downsized with age. They are invested with a large and dense network of trachea and open to the exterior via minute slit shaped openings in the form of ostioles that remain permanently open to the outside. The functional specificity of these glands can be associated with the activities ranging from aggregation to sexual behaviours like courtship and mating. The mating is not restricted to a specific season, being distributed all-round the year; however, the insect prefers low and humid temperature for mating and they can be seen congregating in huge numbers when the hot summers subside and the rainy season arrives. We didn’t observe any considerable variation in size of these glands with respect to the body size in between the instars and the adults. Further, there were no significant difference between males and females in terms of the size of these DAGs.
... Host plant associations include species of Araceae, Euphorbiaceae, Fabaceae, Malvaceae, and Piperaceae ( Brailovsky et al. 1988, Schaefer et al. 2005). The eggs and first-fifth nymphal instars of C. egeris have been recently studied by Bianchi et al. (2011). ...
In this chapter, we provide a taxonomic history for each family and subfamily (if applicable) and, in addition, for the tribes and generic groups within the Pentatomidae, that, perhaps, will give some perception of the taxonomic position of each group. We also discuss the systematic classification within each of these groups and provide general diagnoses and descriptions for all families within the Pentatomoidea and for the subfamilies, tribes, and generic groups within the Pentatomidae. We discuss potentially important characters and point out problems in the existing classification. It is our intention to provide information on the currently recognized or disputed taxa, their defining characters, and their relationships. We also have summarized some of the biological information for each group.
... Terminology for general morphology follows Matesco et al. (2007Matesco et al. ( , 2008 and that for the external structures of the dorsal abdominal scent glands follows Bianchi et al. (2011a) and Vilímová & Kutalová (2012). ...
Nymphs of Euschistus (Mitripus) convergens (Herrich-Schaffer) (Hemiptera: Pentatomidae) are described using light and scanning electron microscopy. In the 1st instar, the integument and the lateral margins of the body are smooth, the dorsal abdominal scent glands bear a rounded peritreme with cuticular valve, and the evaporatorium surface bears pointed projections. From the 2nd to the 5th instars, the integument surface is granulated, the lateral margins of the body are serrated, the dorsal abdominal scent glands bear a spout-shaped peritreme with a cuticular valve, and the evaporatorium surface is reticulated. The overall morphology of E. convergens nymphs resembles the morphology of nymphs of other Euschistus species.
... The associated cuticular structures mostly surround the ostioles. Only seven descriptive papers deal with external DAG structures in the Pentatomoidea: Scutelleridae: Galeacius martini (Bianchi et al., 2011a), Pentatomidae: Cyrtocorinae: Cyrtocoris egeris (Bianchi et al., 2011b;Bottega et al., 2015), Pentatomidae: Pentatominae: Caonabo pseudoscylax (Gilio-Dias et al., 2013), Chinavia obstinata (Bottega et al., 2015), Euschistus (Mitripus) grandis (Biasotto et al., 2013), Glyphepomis spinosa (Pollo et al., 2012), and Acanthosomatidae: Elasmucha ferrugata (Vilímová and Kutalová, 2012). The Pentatomoidea possess a unified pattern of DAGs: three pairs of ostioles, anterior ostioles without peritremes, surrounded only by transversally wrinkled cuticle. ...
The scent efferent system of the nymphal abdominal scent glands of the Rhopalidae: Rhopalinae (Heteroptera) from two tribes, Rhopalini (Rhopalus (Aeschyntelus) maculatus (Fieber, 1837)) and Chorosomini (Chorosoma schillingi (Schilling, 1829)), were described for the first time using SEM. Two unpaired ostioles on medial sclerites are connected by one evaporative area bearing two patterns of microsculpture, tooth-shaped and cone-shaped. Also for the first time, the scent efferent system of dorsal abdominal scent glands of Coreus marginatus (Linnaeus, 1758) (Heteroptera: Coreidae: Coreinae) nymphs was described. The system was compared within the Pentatomoidea, Lygaeoidea, Coreidae, and Rhopalidae; the state in Rhopalidae is intermediate between Lygaeoidea and Coreidae.
Here we present a descriptive analysis of the bibliographic production of the world-renowned heteropterist Dr. Jocélia Grazia and comments on her taxonomic reach based on extracted taxonomic treatments. We analyzed a total of 219 published documents, including scientific papers, scientific notes, and book chapters. Additionally, we applied the Plazi workflow to extract taxonomic treatments, images, tables, treatment citations and materials citations, and references from 75 different documents in accordance with the FAIR (Findability, Accessibility, Interoperability, and Reuse) principles and made them available on the Biodiversity Literature Repository (BLR), hosted on Zenodo, and on TreatmentBank. We found that Dr. Grazia published 200 new names, including species (183) and genera (17), and 1,444 taxonomic treatments in total. From these, 104 and 581, respectively, were extracted after applying the Plazi Workflow. A total of 544 figures, 50 tables, 2,242 references, 2,107 materials citations, and 1,101 treatment citations were also extracted. In order to make her publications properly citable and accessible, we assigned DOIs (Digital Object Identifiers) for all publications that lacked this persistent identifier, including those that were not processed (88 in total), therefore enhancing the open-access share of her publications.
Eggs of 14 species within two subfamilies (Edessinae and Pentatominae, tribes Carpocorini, Nezarini, Pentatomini, Procleticini, and one unplaced species) of neotropical pentatomids were studied with scanning electron microscopy, and their external morphology was characterized in detail (chorion surface, operculum, and aero-micropylar processes). Eggs of these species have similar characteristics to those already described for other pentatomids. Most of them are barrelshaped, except in Edessa meditabunda (with spherical eggs) and Odmalea basalis (whose eggs have flattened lateral faces). A "T"-shaped ruptor ovis is present in all studied species; eclosion line at the operculum rim may be visible or not. Average size of eggs (length x width) ranges from 0.9 ± 0.12 x 0.9 ± 0.05 mm (Euschistus picticornis) to 2.1 ± 0.12 x 1.7 ± 0.10 mm (Chinavia erythrocnemis); the number of the aero-micropylar processes ranges from 10 (C. musiva) to 86 (C. obstinata). Chorion surface has a variable aspect: eggs of Euschistus spp., Loxa deducta, and Pallantia macunaima have the spinose chorion type; C. erythrocnemis, C. longicorialis, C. obstinata, C. pengue, and Grazia tincta have a reticulate chorion; C. musiva, O. basalis, and E. meditabunda have a granulated chorion; and Thyanta humilis has a salebrose chorion type. All the species with described eggs in the Edessinae share the same diagnostic features. In Pentatominae, we did not identify shared characters among the species at subfamily or tribe level. Chorion sculpture pattern and aero-micropylar processes shape, as well as number, are diagnostic features at genus level; however, a few species could be identified in the egg stage.
The 3rd through 5th instars of Cyrtocoris egeris Packauskas & Schaefer are described and illustrated, and the patterns of trichobothria are diagramed. The amount of dorsal abdominal sclerotization and the lateral position of the trichobothria are unusual in the Pentatomoidea.
Banasa maculata sp. nov. is described from a Brazilian Atlantic Forest, including immature stages and aspects of its life history. Adults and nymphs were reared in laboratory and fed on fruits of Miconia sellowiana (Melastomataceae). Eggs and first instars of B. maculata are similar to those of other species of Banasa; however, the color pattern of the abdomen distinguishes B. maculata, particularly first, fourth, and fifth instars. Light and dark morphs were observed for third, fourth, and fifth instars. Head-width measurements overlap only between fourth and fifth instars. Eggs of B. maculata, in S.E.M., show a reticulate pattern with deep cells and irregular rims. The most frequent size of an egg clutch was 12. Average duration of the immature stages (egg to adult) was 37.6 ± 13.24 days. The highest mortality occurred in the fifth instar (45.9%). Banasa maculata belongs to the " cuspidata group " of Banasa because of the presence of an apical projection of each posterolateral angle of the pygophore. Adults of B. maculata resemble B. varians Stål; the new species is recognized by the transversal, irregular, broad macula on pronotum; gonocoxites 8 deeply excavated medially; and the male proctiger subtriangular, narrowing toward the apex.
Eggs of 14 species within two subfamilies (Edessinae and Pentatominae, tribes Carpocorini, Nezarini, Pentatomini, Procleticini, and one unplaced species) of neotropical pentatomids were studied with scanning electron microscopy, and their external morphology was characterized in detail (chorion surface, operculum, and aero-micropylar processes). Eggs of these species have similar characteristics to those already described for other pentatomids. Most of them are barrelshaped, except in Edessa meditabunda (with spherical eggs) and Odmalea basalis (whose eggs have flattened lateral faces). A “T”-shaped ruptor ovis is present in all studied species; eclosion line at the operculum rim may be visible or not. Average size of eggs (length x width) ranges from 0.9 ± 0.12 x 0.9 ± 0.05 mm (Euschistus picticornis) to 2.1 ± 0.12 x 1.7 ± 0.10 mm (Chinavia erythrocnemis); the number of the aero-micropylar processes ranges from 10 (C. musiva) to 86 (C. obstinata). Chorion surface has a variable aspect: eggs of Euschistus spp., Loxa deducta, and Pallantia macunaima have the spinose chorion type; C. erythrocnemis, C. longicorialis, C. obstinata, C. pengue, and Grazia tincta have a reticulate chorion; C. musiva, O. basalis, and E. meditabunda have a granulated chorion; and Thyanta humilis has a salebrose chorion type. All the species with described eggs in the Edessinae share the same diagnostic features. In Pentatominae, we did not identify shared characters among the species at subfamily or tribe level. Chorion sculpture pattern and aero-micropylar processes shape, as well as number, are diagnostic features at genus level; however, a few species could be identified in the egg stage.
Banasa maculata sp. nov. is described from a Brazilian Atlantic Forest, including immature stages and aspects of its life history. Adults and nymphs were reared in laboratory and fed on fruits of Miconia sellowiana (Melastomataceae). Eggs and first instars of B. maculata are similar to those of other species of Banasa; however, the color pattern of the abdomen distinguishes B. maculata, particularly first, fourth, and fifth instars. Light and dark morphs were observed for third, fourth, and fifth instars. Head-width measurements overlap only between fourth and fifth instars. Eggs of B. maculata, in S.E.M., show a reticulate pattern with deep cells and irregular rims. The most frequent size of an egg clutch was 12. Average duration of the immature stages (egg to adult) was 37.6 ± 13.24 days. The highest mortality occurred in the fifth instar (45.9%). Banasa maculata belongs to the “cuspidata group” of Banasa because of the presence of an apical projection of each posterolateral angle of the pygophore. Adults of B. maculata resemble B. varians Stål; the new species is recognized by the transversal, irregular, broad macula on pronotum; gonocoxites 8 deeply excavated medially; and the male proctiger subtriangular, narrowing toward the apex.
The eggs of Apodiphus amygdali (Germar, 1817) were observed with light and scanning electron microscopes. Each female generally laid 14 (13-14) eggs in a mass. The barrel-shaped eggs are average 1.97 mm long and 1.66 mm wide. Eggs were light green color at deposition but changed to yellowish green after embryonic development. The first external evidence of embryonic development is the appearance of two red eye spots opposite each other beneath the operculum. The egg surface is covered with irregular polygonal chorionic pattern seen by scanning electron microscope. There are aeropyles on the corners of polygons. Then, a blackish T-shaped egg burster appears between eye spots and assists hatching. The hatching line is cracked in a circular shape around the operculum by egg burster. They have 18-20 micropylar projections around the hatching line.
The eggshell structures of Eurydema blandum, E. oleraceum, E. rugulosum, E. ventrale, E. spectabile, E. fieberi were investigated by light and scanning electron microscopy. Although the eggs of each species were more or less typical barrel-shaped and had a T-shaped egg burster, some differences in egg morphology were observed, including slight variations in egg shape and size, different patterns of chorionic sculpturing and a specialized opercular area.