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Females of two species of the genus Lasaia Bates, 1868, L. arsis Staudinger, [1887] and L. pseudomeris Clench, 1972 are described from specimens deposited in entomological collections. The identity of these species had been uncertain due to scarcity of data, causing misidentifications and confusion in pairing males and females of the same species. Distribution maps for both species, as well as new records of L. arsis for Mexico are provided.

Se presenta una panorámica de la propuesta actual de Llorente-Bousquets y cols., sobre el empleo de caracteres exocoriónicos en sistemática de Papilionoidea (Insecta: Lepidoptera), con énfasis en sus técnicas de producción de imágenes. Se incluye: 1) breve historia del interés taxonómico de Llorente-Bousquets en estructuras ovocíticas; 2) resúmenes de sus técnicas estandarizadas para el estudio e ilustración del exocorion; 3) discusión sobre el uso de imágenes en la conceptualización, representación y empleo de caracteres exocoriónicos; 4) prospectos a futuro en investigación exocoriónica de Papilionoidea.

The Loxicha Region of Oaxaca, Mexico, has been historically important for the study of Nymphalidae, second in the Papilionoidea for species richness. Describing the diversity patterns of this butterfly clade in Loxicha can improve our understanding of the evolutionary history of the Sierra Madre del Sur, the Mexican Pacific slope, and Mexico in general. Objective: To describe the temporal and spatial patterns of Nymphalid diversity along an elevational gradient (80-2 600 m), and to compare Loxicha’s fauna with other regions in Mexico. Methods: We obtained 28 756 records from 21 sites in the Loxicha Region, representing seven years of sampling. We estimate and analyze the diversity, endemism, and distributional patterns for three elevational levels and five vegetation types. We estimated species composition and similarity with other regions of the Pacific and Atlantic slopes. Results: We identified 189 taxa, including species and subspecies, from 85 genera and ten subfamilies of Nymphalidae. Loxicha contains 46 % of the species in the family recognized for Mexico, including ten endemic species and 56 endemic subspecies. Cloud forest and low elevations were the most diverse habitats for this family. There is a clear divergence between the Atlantic and Pacific faunas, and the Sierra Madre del Sur has two faunal components. High-elevation sites in Oaxaca, and in the neighboring state of Guerrero, have a distinctive fauna, apparently isolated from low-elevation sites, revealing an archipelagic distribution for cloud forest Nymphalidae. Conclusions: The Loxicha Region is one of the richest areas for Nymphalidae in Mexico. Distribution on the Pacific slope is determined by geographical history and ecological conditions, including elevation. Nymphalidae can be used to test hypotheses of biogeographic regionalization in Mexico. Introducción: La Región Loxicha de Oaxaca, México, ha sido históricamente importante para el estudio de Nymphalidae, segunda en riqueza de especies en Papilionoidea. Describir los patrones de diversidad de este taxón de mariposas en Loxicha puede mejorar nuestra comprensión de la historia evolutiva de la Sierra Madre del Sur, la vertiente del Pacífico mexicano y México en general. Objetivo: Describir los patrones temporales y espaciales de la diversidad de Nymphalidae a lo largo de un gradiente altitudinal (80-2 600 m), y comparar la fauna de Loxicha con otras regiones de México. Métodos: Obtuvimos 28 756 registros de 21 sitios de la Región Loxicha, que representan siete años de muestreo. Estimamos y analizamos la diversidad, el endemismo y los patrones de distribución para tres niveles altitudinales y cinco tipos de vegetación. Estimamos la composición de especies y la similitud con otras regiones de las vertientes del Pacífico y Atlántico de México. Resultados: Identificamos 189 taxones, incluyendo especies y subespecies, de 85 géneros y diez subfamilias de Nymphalidae. Loxicha contiene 46 % de las especies de la familia reconocidas para México, incluidas diez especies endémicas y 56 subespecies endémicas. El bosque mesófilo y las elevaciones bajas fueron los hábitats más diversos para esta familia. Existe una clara divergencia entre las faunas del Atlántico y del Pacífico, y la Sierra Madre del Sur tiene dos componentes faunísticos. Los sitios de elevaciones altas en Oaxaca, y en el estado vecino de Guerrero, tienen una fauna distintiva, aparentemente aislada de los sitios de elevaciones bajas, lo que revela una distribución archipelágica para los Nymphalidae del bosque mesófilo. Conclusiones: La Región Loxicha es una de las zonas más diversas para Nymphalidae en México. La distribución en la vertiente del Pacífico está determinada por la historia geográfica y las condiciones ecológicas, incluida la elevación. Nymphalidae puede usarse para probar hipótesis de regionalización biogeográfica en México.

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.

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.

We present a synthesis of the existing information on the genus Emesis Fabricius in Mexico concerning biogeographical patterns and taxonomical aspects. Emesis is the most diverse genus of Emesidini with 57 species and subspecies, with Mexico as the northern limit of this Neotropical genus. We analyzed 5434 specimens of the Lepidoptera Collection of the MZFC, UNAM and compared them with specimens from collections of Mexico, Central and South America. Taxonomic determination and corroboration were made by analysis of wing patterns and genitalia. Geographic distribution and phenology were obtained from the database MARIPOSA. We present an updated list of Emesis of Mexico, with 17 species and subspecies. For each species, we provide information on phenology, geographic, altitudinal, and vegetation distributions. We discuss taxonomic and undersampling concerns for some species, as well as spatial and temporal patterns with special reference to vegetation types and biogeographic provinces in Mexico.

Se describe y compara el exocorion de tres especies en la tribu Coeini con ejemplares de Mexico: Historis odius dious, Colobura dirce dirce, y Smyrna blomfildia datis. Se caracterizaron los principales rasgos estructurales del exocorion: color, forma, tamano, ‘ornamentacion’, diferenciacion micropilar, perimicropilar, y de la zona de transicion, asi como estructuras ecuatoriales y basales; bajo la tecnica de tincion y el empleo del microscopio electronico de barrido. Se reconocen caracteres microestructurales autapomorficos en el corion de estos generos (calgarys, filas), asi como especializaciones en la base (puentes y domos). Para una comprension mayor de estas estructuras, se incluyen esquemas y un glosario con los terminos adoptados en este trabajo. Se realiza la comparacion de nuestros resultados con descripciones publicadas para estas especies, asi como con algunas filogenias de la subfamilia Nymphalinae.

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.

Se describe e ilustra el exocorion de dos especies del genero Colias: C. harfordii y C. philodice guatemalena bajo la tecnica de tincion con azul de metileno y con el empleo del microscopio electronico de barrido. Se caracterizan a detalle las regiones micropilar, perimicropilar, de transicion, ecuatorial, y basal. Se comparan tambien los diversos aspectos de la reticula en sus diferentes regiones o zonas visualizadas con ambas tecnicas, tanto entre si como con otras seis especies de Colias ya publicadas.

Se describe y compara el exocorion de tres especies de Heliconiinae de Mexico (Euptoieta hegesia meridiania, Agraulis vanillae incarnata, y Laparus doris viridis) con otras especies del sur de Brasil ya publicadas. Se enfatizo la comparacion en los principales caracteres exocorionicos como son la ‘ornamentacion’, la diferenciacion micropilar, perimicropilar y de la zona de transicion, asi como las reticulas ecuatoriales y basales; se empleo la tecnica de tincion y el uso del microscopio electronico de barrido. Se reconocen diferencias estructurales en dos de las tribus de Heliconiinae; en Argynnini es caracteristica su forma conica trunca, la presencia de dos reticulas (supra e infraecuatorial) y tal vez quillas en el apice; Heliconiini presenta formas elipsoidales oblongas y una reticula rectangular, con algunos poligonos penta o hexagonales en la region apical. Se discute de modo escueto la morfologia en Acraeini, las posibles implicaciones de la morfologia corionica en la sistematica de Heliconiinae y los caracteres de algunos generos en especifico. Se incluyen laminas y esquemas para una visualizacion mas clara de las estructuras y zonas descritas en el corion de Argynnini y Heliconiini.

We transfer Apodemia planeca R. de la Maza E. & J. de la Maza E. 2017 to Emesis as Emesis planeca (R. de la Maza E. & J. de la Maza E.) n. comb. based on phylogenetic hypotheses estimated with one mtDNA and two nDNA markers. This is a rare and poorly known metalmark, endemic to the central region of Balsas Basin in Michoacán, Mexico, originally described from material collected 23 years ago. Here we analyze new specimens from the type locality not included in the original description. With this new material, we describe the morphological variation of adults, including male and female genitalia. This variation is then discussed and compared with the original description. Emesis planeca n. comb. is restricted to the Tropical Deciduous Forest and the adults fly only in the dry season. Due to the spatial, temporal, and ecological rareness of Emesis planeca n. comb., considering it has not been collected in more than two decades and that the Tropical Deciduous Forest is one of the most threatened habitats in Mexico; we propose assigning a protection status to this species.

Tribe Emesidini Seraphim, Freitas & Kaminski (Lepidoptera: Riodinidae) is distributed in America from southwest Canada to Brazil and Paraguay, and includes 57 species and 54 subspecies (Callaghan & Lamas 2004; Penz & DeVries 2006; Gallard 2008; Pelham 2008; De la Maza & De la Maza 2017 a,b; Kaminski et al. 2017; Seraphim et al. 2018; Trujano-Ortega et al. 2018; Zhang et al. 2019). The tribe has great taxonomic, morphological and ecological diversity, as well as wide geographic and seasonal variation. This great variation and broad geographic range of some genera entail the need for a taxonomic review (Espeland et al. 2015; Trujano-Ortega et al. 2018).

Introduction: A reliable list of species and the analysis of diversity patterns of hyperdiverse taxa, like butterflies, are fundamental for monitoring and managing biological resources. Oaxaca is one of the most diverse states in Mexico for many groups including Lepidoptera and most of its diversity is unknown. Objective: To estimate and describe the species richness and diversity of Papilionidae and Pieridae along an altitudinal gradient and five vegetation types in the Loxicha Region, Oaxaca, Sierra Madre del Sur. Methods: Sampling effort comprised 222 collecting days during a period of seven years. We estimated the alpha diversity for 17 sites within an elevational gradient from 80 to 2 850 m, with five vegetation types: tropical deciduous forest (TDF), tropical sub-deciduous forest (TSDF), cloud forest (low and middle levels) (CF), oak-pine and cloud forest (high level) (OPCF) and oak-pine forest (OPF). Results: We obtained a list of 69 species (27 Papilionidae and 42 Pieridae), of 34 genera and five subfamilies, from literature records and fieldwork. These species are 60 % of the Pieridae and 48 % of the Papilionidae recorded for the state. The Loxicha Region has 36 % of the Pieridae and 30 % of the Papilionidae of Mexico. Both families present different species richness patterns by vegetation type. Papilionidae is richer in the TDF with 23 estimated species and most of the species of this family (84 %) occur below 500 m. Meanwhile, species richness of Pieridae has non-significant differences among vegetations types, except for OPF which has fewer species than the other types. The elevational gradient was divided into three levels (0-750, 750-1 800, 1 800-2 850 m) showing a reduction of species richness and diversity for both families at higher altitudes. Conclusions: Papilionidae species are more restricted to a vegetation type or elevational level than Pieridae species. Likely reasons are higher vagility (including migrations) and wider ecological tolerance of most Pieridae. Resumen Introducción: Una lista de especies confiable y el análisis de los patrones de diversidad de taxones hiperdiversos, como las mariposas, son fundamentales para el monitoreo y manejo de recursos biológicos. Oaxaca es uno de los estados más diversos de México y mucha de su diversidad aun es desconocida. Objetivo: Estimar y describir la riqueza de especies y diversidad de Papilionidae y Pieridae en un gradiente altitudinal y cinco tipos de vegetación presentes en la Región Loxicha, Oaxaca, en la Sierra Madre del Sur. Métodos: El esfuerzo de muestreo comprendió 222 días a lo largo de siete años. Se estima la diversidad alfa para 17 sitios dentro de un gradiente altitudinal de 880 a 2 850 m con cinco tipos de vegetación: bosque tropical caducifolio, bosque tropical subcaducifolio, bosque mesófilo (nivel bajo y medio), bosque de pino-encino con bosque mesófilo (nivel alto) y bosque de pino-encino. Resultados: Se obtuvo una lista de 69 especies (27 Papilionidae y 42 Pieridae), de 34 géneros y cinco subfamilias, a partir de los registros en la literatura y el trabajo de campo. Estas especies representan el 60 % de los Pieridae y el 48 % de los Papilionidae registrados para el estado. A nivel nacional, la región Loxicha tiene el 36 % de las Pieridae y el 30 % de las Papilionidae del país. Las familias presentan patrones diferentes de riqueza de especies por tipo de vegetación. Papilionidae es más rica en el bosque tropical caducifolio con 23 especies estimadas y la mayoría de las especies de esta familia (84 %) están por debajo de los 500 m Mientras que la riqueza de especies de Pieridae no presenta diferencias significativas entre los tipos de vegetación, excepto por el bosque de pino-encino que tiene menos especies que los demás tipos. El gradiente altitudinal se dividió en tres pisos (0-750, 750-1 800, 1 800-2 850 m) con una reducción de la riqueza de especies y diversidad de ambas familias en altitudes mayores. Conclusiones: Las especies de Papilionidae están más restringidas a un tipo de vegetación o piso altitudinal que las especies de Pieridae. Probablemente por la gran vagilidad, valencia ecológica más amplia y capacidad de migraciones altitudinales de la mayoría de las Pieridae.

While surface microstructures of butterfly wings have been extensively studied for their structural coloration or optical properties within the visible spectrum, their properties in infrared wavelengths with potential ties to thermoregulation are relatively unknown. The midinfrared wavelengths of 7.5 to 14 µm are particularly important for radiative heat transfer in the ambient environment, because of the overlap with the atmospheric transmission window. For instance, a high midinfrared emissivity can facilitate surface cooling, whereas a low midinfrared emissivity can minimize heat loss to surroundings. Here we find that the midinfrared emissivity of butterfly wings from warmer climates such as Archaeoprepona demophoon (Oaxaca, Mexico) and Heliconius sara (Pichincha, Ecuador) is up to 2 times higher than that of butterfly wings from cooler climates such as Celastrina echo (Colorado) and Limenitis arthemis (Florida), using Fourier-transform infrared (FTIR) spectroscopy and infrared thermography. Our optical computations using a unit cell approach reproduce the spectroscopy data and explain how periodic microstructures play a critical role in the midinfrared. The emissivity spectrum governs the temperature of butterfly wings, and we demonstrate that C. echo wings heat up to 8 °C more than A. demophoon wings under the same sunlight in the clear sky of Irvine, CA. Furthermore, our thermal computations show that butterfly wings in their respective habitats can maintain a moderate temperature range through a balance of solar absorption and infrared emission. These findings suggest that the surface microstructures of butterfly wings potentially contribute to thermoregulation and provide an insight into butterflies' survival.

Se describe y compara la macro y la microrreticula exocorionica en tres especies del genero Colias: C. croceus, C. alfacariensis, y C. lesbia mineira; bajo tecnicas de tincion con azul de metileno y con el empleo del microscopio electronico de barrido (MEB). El tamano, la forma, asi como diversos aspectos de la trama en sus diferentes regiones o zonas, se exhibe distinta para cada especie. Las diferencias ocurren en el micropilo, el area perimicropilar, el grosor relativo entre ejes y costillas, el arreglo y numero de los poligonos que componen la microrreticula, entre otros caracteres que se detallan. Todos estos caracteres se ilustran en dibujos y fotografias en el MEB. Los resultados se comparan entre cada especie y con las de trabajos previos.

Se examina y describe la estructura del corion de Leucidia brephos con la tecnica de tincion de azul de metileno y con el empleo de microscopio electronico de barrido (MEB). Se realizo su comparacion morfologica, incluyendo a coriones ilustrados o fotografiados de otros generos de la subfamilia Coliadinae. El corion en Leucidia brephos presenta una macrorreticula bastante generalizada dentro del grupo, ademas de una forma alargada y base con reticulacion poligonal multiple, lo que la hace plesiomorfica. La microrreticula, junto con los ejes y las costillas, conforma una estructura de relieve en tres planos diferentes, bastante particular. Los lados de la microrreticula son de distintos grosores y el numero de poligonos irregulares entre las costillas es reducido, casi siempre menor a 10. Este nuevo conocimiento confirma la presencia de la microrreticula en las Coliadinae, explicando su ausencia en algunas especies como una perdida secundaria y, junto con los datos previamente obtenidos en Dismorphiinae, Coliadinae y Pierinae, nos permiten resumir las tendencias de macrorreticula sobre Pieridae: la tendencia a formar poligonos cuadrangulares, la simplificacion de los ejes y las costillas, y la desaparicion de poligonos irregulares y multiples hacia los polos.

Two new genera of Riodinidae (Insecta: Lepidoptera) are described, Neoapodemia Trujano-Ortega, gen. n. (Neoapodemia nais (W. H. Edwards, 1876), comb. n., N. chisosensis Freeman, 1964, comb. n.) and Plesioarida Trujano-Ortega & García-Vázquez, gen. n. (Plesioarida palmerii palmerii (W. H. Edwards, 1870), comb. n., P. palmerii arizona (Austin, [1989]), comb. n., P. palmerii australis (Austin, [1989]), comb. n., P. hepburni hepburni (Godman & Salvin, 1886), comb. n., P. hepburni remota (Austin, 1991), comb. n., P. murphyi (Austin, [1989]), comb. n., P. hypoglauca hypoglauca (Godman & Salvin, 1878), comb. n., P. hypoglauca wellingi (Ferris, 1985), comb. n., P. walkeri (Godman & Salvin, 1886), comb. n., P. selvatica (De la Maza & De la Maza, 2017), comb. n.). Neoapodemia Trujano-Ortega, gen. n. is distributed in the southwestern USA and northeastern Mexico, while Plesioarida Trujano-Ortega & García-Vázquez, gen. n. is present from the southern USA to Central America. Species of these genera were previously classified as Apodemia C. Felder & R. Felder but molecular and morphological evidence separate them as new taxa. Morphological diagnoses and descriptions are provided for both new genera, including the main distinctive characters from labial palpi, prothoracic legs, wing venation and genitalia, as well as life history traits. A molecular phylogeny of one mitochondrial gene (COI) and two nuclear genes (EF-1a and wg) are also presented of most species of Apodemia, Neoapodemia Trujano-Ortega, gen. n., Plesioarida Trujano-Ortega & García-Vázquez, gen. n., and sequences of specimens from all tribes of Riodinidae. We compare the characters of Apodemia, Neoapodemia Trujano-Ortega, gen. n. and Plesioarida Trujano-Ortega & García-Vázquez, gen. n. and discuss the differences that support the description of these new taxa. This is a contribution to the taxonomy of the Riodinidae of North America of which the generic diversity is greater than previously recognized.

Resumen. Se describe y nombra una subespecie nueva de Prepona brooksiana G. & S., la cual es la mas septentrional de todas las razas geograficas de este taxon; ocurriendo dentro de la vertiente del Golfo de Mexico en la porcion mas boreal del bosque mesofilo de montana. Se describe la variacion poblacional de los caracteres que la diagnostican y se mencionan aspectos de su conducta y habitat. Se determina que las subespecies mesoamericanas del grupo Prepona deiphile realmente corresponden a Prepona brooksiana, cuya distribucion solo ocurre en Mexico y Centroamerica. De esta forma, el grupo deiphile en el genero Prepona Boisduval estaria compuesto de tres especies: P. deiphile Godart y P. xenagoras Hewitson, con distribucion en Sudamerica, y Prepona brooksiana en Mexico y America Central.

Se examina el corion de Ectima thecla thecla y se compara con la morfologia corionica de Hamadryas spp.; se construye un glosario ilustrado de la terminologia empleada en las descripciones corionicas en Ageroniina. El corion de ambos generos se tipifica como globoso y unico para el clado; con estructuras plesiomoficas en Ectima respecto a Hamadryas. Se senala la necesidad de aclarar y detallar las descripciones corionicas de Batesia y Panacea, debido a que publicaciones previas reconocen otros tipos de corion afines a diferentes subgrupos de Biblidinae, cuando tal vez se esperaria un corion globoso para toda la subtribu Ageroniina.

Se examina la estructura corionica de Orophila cardases campaspe; tambien se efectuan comparaciones con el corion de siete especies de la subtribu Callicorina, taxon que aqui se caracteriza. Su corion se tipifica por la presencia de ‘ventanas’ en la base de las crestas, el relieve apical en semicirculos y la ausencia parcial o total de las costillas entre los ejes. Se reconocieron las similitudes del corion entre esta y la subtribu Epiphilina, asi como la posibilidad de reduccion u optimizacion del material que derivaria en las crestas ‘en puente’ de la subtribu Eubagina. Es indispensable una mayor exploracion del corion en otras especies de Callicorina, por su posible empleo como fuente de caracteres y para contrastar la hipotesis de Freitas et al. (2014) sobre las relaciones filogeneticas entre los generos de dicha subtribu.

Numerous animal lineages have expanded and diversified the opsin-based photoreceptors in their eyes underlying color vision behavior. However, the selective pressures giving rise to new photoreceptors and their spectral tuning remain mostly obscure. Previously, we identified a violet receptor (UV2) that is the result of a UV opsin gene duplication specific to Heliconius butterflies. At the same time the violet receptor evolved, Heliconius evolved UV-yellow coloration on their wings, due to the pigment 3-hydroxykynurenine (3-OHK) and the nanostructure architecture of the scale cells. In order to better understand the selective pressures giving rise to the violet receptor, we characterized opsin expression patterns using immunostaining (14 species) and RNA-Seq (18 species), and reconstructed evolutionary histories of visual traits in five major lineages within Heliconius and one species from the genus Eueides. Opsin expression patterns are hyperdiverse within Heliconius. We identified six unique retinal mosaics and three distinct forms of sexual dimorphism based on ommatidial types within the genus Heliconius. Additionally, phylogenetic analysis revealed independent losses of opsin expression, pseudogenization events, and relaxation of selection on UVRh2 in one lineage. Despite this diversity, the newly evolved violet receptor is retained across most species and sexes surveyed. Discriminability modeling of behaviorally preferred 3-OHK yellow wing coloration suggests that the violet receptor may facilitate Heliconius color vision in the context of conspecific recognition. Our observations give insights into the selective pressures underlying the origins of new visual receptors.

Riodinidae are one of the most diverse families of Lepidoptera, mainly in the Neotropical region; however, their biology, ethology, taxonomy, systematics, and biogeography are poorly known. In Mexico, the regional and local distributions of the family are still incomplete. We review the distributional data of the genus Lasaia Bates (Lepidoptera: Riodinidae), based on records from four national and seven international collections. We record five species and seven subspecies in Mexico, with 2722 records, distributed in 314 localities of 24 states. The states with higher species richness are Chiapas, Oaxaca, and Veracruz; also, the genus was recorded in 11 of the 14 biogeographical provinces of Mexico. The tropical semi-deciduous and deciduous forests, below of 1000 m a.s.l., contain most of the diversity of Lasaia. Historical data are crucial for the study of local and regional diversity and ecological patterns at large temporal scales. Data presented here show the morphological and ecological variation of Lasaia over the last 80 years, mostly from the XX century when anthropogenic disturbances were intensified. This kind of studies is the first step in recording the historical distribution of these taxa, which will lead to more complex analyses on distribution range shifts, their causes and consequences.

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 .

In our previous work we studied the ultrastructure of the antennae of pierids through a character analysis. In this study we describe the antennal ultrastructure of three species and six subspecies of Patia focused on these traits at the intrageneric level. Patia is a genus of Dismorphiinae with the highest number of scaleless antennomeres (nudum = 11‒15). Therefore, the length of its antennal club is greater than that of any other genus of the subfamily. Several other features differentiate the species of Patia , particularly the number of sulci and pseudosulci and its contour or perimeter. A pattern was observed, especially in P. rhetes , in which the central sulci are disintegrated and surrounded by several pseudosulci in the basal antennomeres and along the club; in the medial and distal antennomeres, the pseudosulci merge to form sulci aggregates. Pseudosulci are uncommon on the distal antennomeres, even in P. rhetes . The central sulci are irregular and have a discontinuous edge when they are disaggregated and accompanied by pseudosulci; those that are elliptical and have a continuous contour, seldom have pseudosulci nearby. Another noteworthy feature is the different shapes of the distal antennomere that occur in the subspecies of P. cordillera and P. orise . In the former, this antennomere is quite elongated and ends at a point, whereas the distal one is less elongated and its apex blunt. Patia rhetes shows the lowest number of scaleless antennomeres (n = 11) while P. cordillera sspp. the largest (n = 14 or 15). The variety of the sensilla present in the Patia species does not provide specific differences, except in P. cordillera sororna where we discovered the presence of a claviform sensillum that does not present in any other species or subspecies. In our small sample size, we did not observe sexual dimorphism.

DNA barcoding regions have been used for identifying organisms and delimiting species. Our research focused on butterflies belonging to the Enantia jethys species complex (Lepidoptera: Pieridae) in Mexico, specifically to resolve the taxonomic problem of the number of species in the group. We used the standard segment of approximately 650 base pairs of the mitochondrial cytochrome oxidase subunit I (COI) gene. Our study is the first to use DNA sequences to examine phylogenetic relationships of this complex species. Three phylogenetic inference methods used were parsimony, maximum likelihood, and Bayesian inference. Twospecies delimitation methods also were used: generalized mixed Yule-coalescent and Poisson tree process (bPTP). We used all the analyses to obtain 155 COI sequences and a persistent clade with four monophyletic groups: three corresponding to Enantia albania (Bates), Enantia jethys (Boisduval), and Enantia mazai Llorente, and a fourth corresponding to a new potential cryptic species, which must be described.

Resumen. Se estudia e ilustra la estructura corionica de cuatro especies de dos generos de la subfamilia Coliadinae: Colias (C. dimera, C. erate, y C. eurytheme) y Zerene (Z. c. cesonia). Se reexaminan y comparan descripciones previas de C. dimera, C. eurytheme y Z. c. cesonia, con nuevos materiales provenientes de Mexico y Colombia. Ademas, por primera vez se describe el corion de C. erate y la microrreticula en tres de las cuatro especies; que se pensaba ausente en ambos generos, por su carencia en C. dimera. Con el uso del microscopio electronico de barrido (MEB), se efectuaron tomas fotograficas de cuatro regiones corionicas (micropilar, perimicropilar o apical, ecuador, y base); estas se describen, ilustran, y comparan tabularmente. Las tomas en el MEB permiten analizar y resolver que C. dimera presenta caracteres de generos menos derivados dentro de los Coliadinae, mientras que en C. eurytheme se expresan caracteres encontrados en generos mas derivados. Sin embargo, caracteres reticulares del area b...

Resumen. Se describe y compara la arquitectura corionica de tres especies pertenecientes a generos poco derivados de las Coliadinae: Nathalis plauta (Doubleday, 1847), N. i. iole (Boisduval, 1836), y Kricogonia lyside (Godart, 1819). La seleccion constituye un refinamiento de estudios anteriores del corion en este grupo, al reconocer la presencia de microrreticula en las tres especies. El microscopio electronico de barrido (MEB) se incorpora con la tecnica de tincion, enriqueciendo la descripcion y analisis de la arquitectura corionica en las regiones apical, ecuatorial y basal. Se tipifican dos clases de macrorreticula hexagonal y dos de macrorreticula rectangular; se identifica un estado intermedio entre ambas, mediante el concepto de ‘eje zigzagueante’, con implicaciones taxonomicas y filogeneticas. Los resultados generales del estudio permiten analizar la sistematica de Coliadinae con implicaciones clasificatorias y biogeograficas.

Resumen. Se complementa la descripcion e ilustracion relativamente detallada del corion en la subfamilia Biblidinae, al agregarse 11 especies distintas a las presentadas recientemente, en los generos Hamadryas, Marpesia, Eunica, Catonephele, Myscelia, Asterope, Epiphile, Callicore, y Cyclogramma. Los ejemplares provienen de Mexico y Colombia. El estudio se realizo a partir de una muestra de 35 hembras, de las cuales 15 tuvieron huevos maduros. Se caracterizaron los principales rasgos estructurales del corion: color, forma, tamano y relieve: diferenciacion perimicropilar y formacion de ejes y costillas. Se hace una distincion de rasgos especializados en las zonas apical y ecuatorial, asi como el grado de engrosamiento de los ejes. Se siguen los tipos de huevos reconocidos recientemente, asi mismo la nomenclatura de las descripciones. Los resultados confirman los tipos de huevo y arquitectura propias de cada taxon supragenerico. Ademas, apoyan la propuesta de emplear al corion como caracter taxonomico, al e...

Resumen. Se estudia e ilustra la estructura corionica de cuatro especies de dos generos de la subfamilia Coliadinae: Anteos y Rhabdodryas. Se reexamina la descripcion de Anteos maerula y A. clorinde con nuevos materiales provenientes de Mexico y Colombia. Por primera vez se inspecciona el relieve corionico de A. menippe y su microrreticula, ademas de la de Rhabdodryas trite, que se pensaba ausente. Con el uso del microscopio electronico de barrido (MEB) se efectuaron tomas fotograficas de cuatro regiones corionicas (micropilar, perimicropilar o apical, ecuador y base) en las tres especies de Anteos; estas se describen, ilustran, y comparan tabularmente. Las tomas en el MEB permiten analizar y resolver que la microrreticula es tambien una seccion externa del exocorion (relieve): una matriz poligonal multiple en cada unidad macrorreticular. Hasta ahora su presencia en los generos estudiados, asi como en Prestonia, Aphrissa, y Phoebis, permiten postularla como una sinapomorfia en el grupo llamado ‘Catopsilia...

Resumen. Se presenta una lista de 1331 taxones a nivel especifico, pertenecientes a 462 generos, 24 subfamilias, y seis familias de Papilionoidea (sensu lato) del estado de Oaxaca; que constituyen 68% de tales taxones registrados para Mexico. Con esto se actualiza la lista de mariposas diurnas del estado con 165 registros nuevos a nivel especifico, de los cuales 17 tambien son registros nuevos para este pais. De las especies y subespecies oaxaquenas, hasta ahora se han registrado 259 endemicas de Mexico, que representan 19.45% del total en el estado y 57.5% de los endemicos que ocurren en el pais. Estas cifras y porcentajes indican que Oaxaca es el estado con mayor riqueza y endemismo de Mexico. La alta riqueza de Papilionoidea de Oaxaca y su endemismo, se debe principalmente a tres factores: 1. la asociacion estrecha que existe entre las mariposas diurnas y el bosque tropical perennifolio; 2. el alto porcentaje de fauna endemica asociada a comunidades xericas del noroeste del estado, los valles centrales...

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.

A new species of Cyllopsis R. Felder, 1869, is described and illustrated from the highlands of Chiapas, Mexico. Cyllopsis tomemmeli A. Warren & Nakahara, sp. nov., is currently known from 13 specimens (9 males and 4 females) collected on March 26–28, 1959, southeast of San Cristóbal de Las Casas. Despite extensive studies on the butterfly fauna of this region, this species has not since been encountered. We discuss possible relationships between this new species and other species of Cyllopsis.

We present the results of the first examination and comparative analysis of the ultrastructure of the antennae in seven species of Leptidea Billberg, 1820 (Pieridae). The results add further support to the hypothesis that the subfamily Dismorphiinae is composed of the tribes Leptideini and Dismorphiini. We summarize the results in a table and discuss the ultrastructure similarities and differences between the two tribes. Following a previously proposed framework, we trace the progression of character states and discuss the significance of the results in the context of varying phylogenetic hypotheses for Leptidea, with special emphasis on biogeography. We include a brief description of the terms we employ to described the main characters and types of sensilla of the antennal club of Leptidea. Remarkable antennal sexual dimorphism is noted in L. gigantea.

"El orden Phasmatodea ha sido poco estudiado en México y no existe un trabajo formal que enliste las especies presentes en el país, o una clave de determinación taxonómica que permita la identificación a nivel genérico. En este trabajo se realizó una revisión bibliográfica exhaustiva con el fin de obtener un listado de especies de Phasmatodea registradas en México. Se registró un total de 17 géneros y 65 especies de Phasmatodea para el territorio mexicano. Se presenta una clave genérica ilustrada de identificación con su respectivo glosario terminológico especializado."

We propose a revised classification for the genus Elzunia Bryk, 1937, with four species and 20 subspecies. We describe two new subspecies, Elzunia humboldt carlosi Le Crom and Llorente, ssp. nov., and Elzunia humboldt willmotti Le Crom, Llorente and Andrade, ssp. nov. We recognize five additional unnamed subspecies but do not describe them because they are represented by too little material. A detailed examination of 854 specimens from 31 collections allowed us to define 24 stable phenotypes and delimit their geographic distribution. We present a diagnosis and description for each taxon and provide images of the wing coloration pattern and male genitalia, and a distribution map. We also present a key for all the species and subspecies. We conducted a multivariate statistical analysis of measurements of the discal cell (DC) veins, and with this we explore the species groupings by these characters. Lastly, we discuss mimetic interactions of this genus with species of other Ithomiini and Heliconiinae.

Resumen La familia Riodinidae es muy diversa en la región Neotropical; en México aún es escaso el conocimiento acerca de la biología de la mayoría de sus especies. Para el presente trabajo, se realizaron recolectas exhaustivas y sistemáticas de Riodinidae en varias localidades de la región Loxicha, Oaxaca, México, particularmente en dos con bosque tropical subcaducifolio. Además se examinaron los registros previos para esa región, en un gradiente altitudinal de los 70 a 2 500 m; la mayoría recolectados por personal del Museo de Zoología (Facultad de Ciencias, UNAM). A partir de la revisión de la base de datos MARIPOSA y del trabajo de campo, se alcanzó la cifra de 4 029 ejemplares de esta región; de los cuales, 2 306 se obtuvieron del bosque tropical subcaducifolio (bts) durante el presente estudio. El total de especies para la región, arrojó un total de 52 especies, distribuidas en 22 géneros y dos subfamilias; la subfamilia Riodininae la más diversa (50 spp.). Mediante estimadores paramétricos se calculó la riqueza de esta familia para las dos localidades de bts estudiadas y en el total de la región; alcanzando valores teóricos mayores al 80 %. Al comparar los resultados con otras listas de especies del Pacífico mexicano, se concluye que en la región Loxicha se encuentra la mayor riqueza de la familia Riodinidae. En el bosque tropical subcaducifolio se efectuaron transectos por dos años, que permitieron distinguir los microhábitats de más de 40 especies, lo cual se ilustra y describe en dibujos y fotografías in situ. En este tipo vegetacional se registra la mayoría de especies de la región Loxicha. Abstract Riodinidae is very diverse in the Neotropical region; in Mexico the knowledge about the biology of most of its species is still scarce. For the present work, exhaustive and systematic collections of Riodinidae were carried out in 14 localities of the Loxicha region, Oaxaca, Mexico, particularly in two with tropical semi-deciduous forest. In addition, previous records for this region were examined, in an altitude range of 70 to 2 500 m; the majority collected by the staff of the Museo de Zoología (Facultad de Ciencias, UNAM). We obtained 4 029 specimens from the fieldwork and MARIPOSA database, 2 306 of them are present in the tropical semi-deciduous forest (TSF). The total number of species for the region is 52, distributed in 22 genera and two subfamilies; the most diverse subfamily was Riodininae (50 species). Using parametric estimators the richness of the family was calculated for the two locations of TSF studied and the entire region, reaching theoretical values greater than 80 %. When comparing the results with other lists of species of the Mexican Pacific, it is concluded that the Loxicha region harbours the greatest richness of Riodinidae. In the tropical semi-deciduous forest, transects were carried out for two years, allowing microhabitats to be distinguished from more than 40 species, which are described and illustrated with drawings and in situ photographs. In this vegetation type most species of the Loxicha region are recorded. Rev. Biol. Trop. 66(2): 503-558. Epub 2018 June 01.