Exochorion in the tribe Nymphalini (Lepidoptera: Nymphalidae): the genus Hypanartia Hbner, [1821] and comparison with related genera.

We describe and compare the exochorion of six species of Hypanartia (H. paullus, H. bella, H. lethe, H. godmanii, H. dione disjuncta, and H. trimaculata autumna) using specimens collected in the field and deposited in entomological collections. We used a standard staining technique and scanning electron microscopy to visualize and describe the main exochorionic characters, especially the ridges and the differentiation in the micropylar, perimicropylar, and transition zones in the apical region. We included plates with photographs, drawings, and schemes for clearer visualization of these structures, which are tabulated for comparison. For our characterization, we considered the ridges in colonnade as the main exochorionic feature in the Nymphalini. Thus, we made brief comparisons of these features among several species of the tribe, particularly with the genus Antanartia. The exochorionic characters agree with the separation of groups (paullus group and dione group). By examining the characteristics of H. paullus, the basal species of the paullus group, we found that ridges with conspicuous columns (e.g., in one section only) appears to be the plesiomorphic state, while those with elongated shafts and walls only in the intercolumn represent the apomorphic state.

Micropylar and Perimicropylar Regions of the Egg Exochorion in Six Genera of Dismorphiinae (Lepidoptera: Pieridae).

Using scanning electron microscopy (SEM), we examine and compare the micropylar and perimicropylar regions of the egg exochorion in six genera of the subfamily Dismorphiinae (Pieridae): Leptidea Billberg, Enantia Hübner, Pseudopieris Godman Salvin, Lieinix Gray, Dismorphia Hübner, and Moschoneura Butler. We provide the first descriptions of the morphological structures of the chorion in these genera. Leptidea is the most primitive genus, and Dismorphia represents the most diverse. We examined the jethys complex of Enantia to study diversity of characters within the genus. We conclude that the exochorion of Dismorphiinae is the most plesiomorphic in relation to Coliadinae and Pierinae, owing to its simple morphology without a transition zone surrounding the wreath (perimicropylar region). Leptidea (Leptideini) shows the least derived characters of the subfamily, followed by Enantia (Dismorphiini). The latter genus exhibits several typical characters of the tribe that are distinctive from other more specialized genera. Exochorionic similarities among the four species of the jethys complex support their close relationship; E. mazai exhibits the most derived features among the species of the complex. Pseudopieris and Moschoneura exhibit several plesiomorphies, such as a large number of openings (micro-grid), but they also show some derived features, such as three semi-rings in the wreath. Dismorphia and Lieinix exhibit characters in their most derived states, such as a striated micro-grid (most conspicuous in Lieinix) and basal aeropyles (most conspicuous in Dismorphia), a character shared with Moschoneura. The purpose of this work is exploratory and descriptive and would be improved by a larger sample size. This contribution is the third in a series of studies on the chorion of Dismorphiinae eggs.

Toward standards in practices and techniques on ootaxonomy in the Pieridae (Lepidoptera: Papilionoidea).

We present a proposal on the standards used on ootaxonomy practices and techniques in the butterfly family Pieridae Duponchel (Lepidoptera: Papilionoidea) in five stages: 1) getting the specimens, 2) integration into a collection, 3) dissections to recover the exochorion, 4) elaboration of images of it, and 5) the preparation of its description with the necessary diagrams and tabulations. Also, we present the detailed techniques applied in observation and graphic representation, based on the methylene blue staining techniques and those required for the use with the scanning electron microscope (SEM). We compare the result of the standardized techniques with those from macro photography, drawings, and photographs with SEM-all of them found in books with descriptions and images of eggs of the Pieridae. We present a glossary and general aspects of the exochorion in the Pieridae as an Appendix to this article. Standardized techniques show more accurate and extensive character retrieval for systematics. For the scale in which they work, these techniques recovered more information than those present from oviposited eggs, where the exochorionic base is not seen. Also, the descriptions contain detailed data on more structures-which are comparable to each other-than are absent in the references mentioned. We present the recovered characters with the techniques found in the literature as three synthetically supplementary materials.

Chorion exploration in the tribe Anthocharidini (Lepidoptera: Pieridae) and their possible importance in its systematics.

We describe and compare the morphology of the chorion in nine species belonging to five genera of the tribe Anthocharidini (Pieridae: Pierinae), from a sample of 12 females with mature eggs, the bibliographic record of oviposited eggs, and photographs of scanning electron microscopy (SEM). The eggs examined come from Mexico, Spain, Brazil and Costa Rica. Its characterization was made considering the main structural features of the chorion in this tribe; it includes a distinction of structures in the apical zone and differentiation between the chorionic regions. We attached to this work sheets, diagrams, and terminology to understand and clarify the descriptions. Our results agree with the proposal of Anthocharidini as the least derived tribe of the Pierinae, considering that Hebomoia, a specialized genus, is not part of it. Tribes such as Leptosiaini, Elodinini or Nepheroniini also have more chorionic characteristics related to more derived tribes of the Pierinae.

Use of exochorion characters for the systematics of Hamadryas Hübner and Ectima Doubleday (Nymphalidae: Biblidinae: Ageroniini).

We studied the chorionic morphology of six species of Hamadryas, and together with previous studies, we compared our results with previously published phylogenies for the genus. Samples were obtained from 19 females collected between 2013 and 2017 whose abdomens were sectioned and preserved for later dissection. Eggs were extracted from those dissections and used for the descriptions and illustrations of the chorion. The Hamadryas egg is of the globose type; it is quasi-spheroidal and has multiple polygonal grids with differentiation in specific zones/regions, and knolls with macrocells in their summits that arise in the apical third. These characteristics are very different from those found in the majority of Biblidinae and for those reported in the literature for Batesia and Panacea, which belong to the same subtribe as Hamadryas (Ageroniina, now Ageroniini). Chorionic characters support a previously suggested division of the genus (februa, feronia and laodamia groups) and they agree with the phylogenetic proposal based on morphological characters. Our study expands previous morphological work focused on this genus and compiles all the information available to date about the exochorion of Hamadryas, which now includes data for 10 species and that of Ectima thecla thecla, the putative sister group of Hamadryas.

Chorionic sculpture of eggs in the subfamily Dismorphiinae (Lepidoptera: Papilionoidea: Pieridae).

This is the second exploration, comparison, and analysis of the chorion of species (45 sspp.) of the subfamily Dismorphiinae (Pieridae). This study includes nearly 50% of the species of the subfamily, including six of the seven genera in its two subtribes: Leptidea (Leptideini), Enantia, Pseudopieris, Lieinix, Moschoneura, and Dismorphia (Dismorphiini). The material studied originates from more than three dozen localities in six different countries on three continents (America, Asia, and Europe) and two biogeographical regions, the Palearctic and Neotropical, over the last 20 years. We have corrected and added information regarding several morphological aspects of the chorion. The precision of the citriform configuration and the elimination of the meloniform shape in the egg of Dismorphiini were determined with detailed observations on the maturation of the chorion in the ovarioles where each stage appears in a linear sequence. We discerned that the meloniform states correspond to incompletely differentiated or immature eggs. This was confirmed by the study of new samples of Dismorphia amphione, D. eunoe, and D. lewyi. The chorion of Dismorphiinae is basically plesiomorphic with respect to those of Coliadinae and Pierinae because it lacks several typical synapomorphies of these subfamilies, such as the presence of micro-grid and/or perimicropylar and apex differentiation, respectively. The eggs of each Dismorphiinae genus can be diagnosed by a combination of chorionic features, although sometimes by one or more plesiomorphies or apomorphies in each genus, with respect to the form or character states in axes, ribs, and poles in the grid of the three regions of the egg - two polar regions and one equatorial (basal, medial, and apical). Leptidea and Enantia show the most generalized grid pattern; however, two genera retain several plesiomorphies with respect to the undifferentiated axes or a small number of short axes (Pseudopieris), as well as many equidistant ribs (Lieinix). The chorionic grid of Moschoneura, although practically lacking short axes, shows the fewest number of axes in the entire subfamily (eight aligns it with Pseudopieris). The chorionic grid in Dismorphia is highly diverse, as it shows the most derived states; however, it comprises symplesiomorphies or atavisms in two groups of species, which aligns them closer to Lieinix or Pseudopieris, but we do not take them into account in some cases where they are convergences or structural parallelisms. It seems that the combination of the shape and its length:width ratio is correlated with the alar configuration (design, sexual dimorphism, and coloring patterns) and separates three groups of species in Dismorphia, and often correlates with the number of ribs. This also coincides with the Batesian participation in the number of mimetic complexes in which a subgroup of species and their stenoecy are integrated within the primary forests. Finally, two schemes are presented that synthesize and illustrate the changes or progression of the form and chorionic grid in the genera of the subfamily.