Distribution, diversity, endemism, and ecology of Nymphalid butterflies (Lepidoptera: Nymphalidae) in the Loxicha Region, Oaxaca, Mexico / Distribución, diversidad, endemismo y ecología de las mariposas Nymphalidae (Lepidoptera: Nymphalidae) en la Región Loxicha, Oaxaca, México

Abstract Introduction: 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 Nymphalidae 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.

Resumen 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.

Geographical distribution of Emesis/ Fabricius (Lepidoptera: Riodinidae) in Mexico: Updated checklist and temporal patterns.

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.

Emesis planeca n. comb. (Lepidoptera: Riodinidae): a new combination revealed by molecular evidence with a description of its morphological variation.

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.

Challenges for organismic taxonomical revisions in the age of phylogenomics: A response to Zhang et al. (2019).

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).

Distribution and diversity of Papilionidae and Pieridae (Lepidoptera: Papilionoidea) in Loxicha Region, Oaxaca, Mexico

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.

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.

Geographical distribution of Lasaia Bates, 1868 (Lepidoptera: Riodinidae) across the biogeographical provinces of Mexico.

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.

Two new genera of metalmark butterflies of North and Central America (Lepidoptera, Riodinidae).

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.

Diversity and Resource Use Patterns of Anthophile Insects in Cuatro Ciénegas, Coahuila, Mexico.

Bees and flies are the main pollinators responsible for the maintenance of plant diversity and crop production. Studies of pollinator communities have focused on bees; however, community structures may vary among groups of pollinators. We describe and compare the diversity and resource use patterns of six Apoidea and two Diptera anthophile families in Cuatro Ciénegas in the northeast of Mexico. We sampled insects in two distinct geographic units (valley and mountains) and two seasons (rainy and dry). Spatial and temporal patterns in species composition and abundance of these families were identified and the diversity in each family compared. The spatial and temporal patterns in species composition, diversity, and resource use were different for each family. Overall, diversity was higher during the rainy season. Only fly families and the Andrenidae presented season-specific differences in species composition, in contrast with the other Apoidea. Two bee families visited more plant species per insect species during the rainy season, whereas flies visited the same number of plant species in both seasons. Apidae, Megachilidae, and Bombyliidae visited more plant species in the mountains, while the other families exhibited no differences between the mountains and the valley. The variation in the community structures of each group may reflect distinct life histories, resources needs during larval stages, and foraging behaviors. These differences are relevant in management and conservation programs that could benefit one group of pollinators while harming others with different characteristics. The traits of some families indicate their potential value as pollinators.

New records of bee flies (Diptera, Bombyliidae) from Cuatro Ciénegas, Coahuila, Mexico.

Forty one new records of species of Bombyliidae are reported for Coahuila in northeastern Mexico. Nine of these species are reported for the first time for the country. The specimens were collected in the Cuatro Ciénegas Basin and Sierra La Madera mountains during 2007-2013. The modified distributions of species are discussed. The gaps in the distribution of many species suggest an undersampling of this group of insects in the north of Mexico.

Patrones de distribución de las especies de la tribu Meliponini (Hymenoptera: Apoidea: Apidae) en México / Distribution patterns of species of the Meliponini (Hymenoptera: Apoidea: Apidae) tribe in Mexico

Basados en la comparación de 43 trazos individuales, se analizaron los patrones de distribución de la tribu Meliponini en México usando el método panbiogeográfico. Se encontraron cuatro trazos generalizados y 13 nodos panbiogeográficos. Tres trazos generalizados forman un componente Neotropical que corre por la Costa del Pacífico o por la Costa del Golfo, uniéndose en el norte de Chiapas donde se bifurca al sur hacia América del Centro, y hacia el noreste por la Península de Yucatán. Ocupa ocho provincias biogeográficas: Costa del Pacífico Mexicano, Eje Volcánico Transmexicano, Depresión del Balsas, Sierra Madre del Sur, Sierra Madre Oriental, Golfo de México, Chiapas y Península de Yucatán. Los trazos generalizados y los nodos coinciden con otros previamente reconocidos para otros taxones.