The butterflies (Lepidoptera, Papilionoidea) of the Parque Estadual Intervales and surroundings, São Paulo, Brazil / As borboletas (Lepidoptera, Papilionoidea) do Parque Estadual Intervales e arredores, São Paulo, Brasil

Abstract The Global South has witnessed increasing sampling of its immense biological diversity during the past century. However, the diversity of many regions remains unknown, even at pristine and highly threatened places, such as in the Atlantic Forest; and for bioindicator, umbrella, and flagship groups. The present study reports the first butterfly list of the Parque Estadual Intervales, São Paulo, Brazil and surroundings, a key protected area in the last massive continuous of the Atlantic Forest. We compiled data from museums and four years of field work, under three sampling methods. We also aimed at providing resources to support conservation efforts by analyzing 27 years of climatic data (detailed in the Supplementary Material, in English and in Portuguese), discussing our results also for non-academics, and producing scientific outreach and educational material. A companion article dealt with the experiences of science outreach and capacity development, and illustrated the butterfly catalogue of the species sampled in the park. We found 312 species that sum to 2,139 records. The museum had 229 species (432 records), and we sampled 142 species (1,682 individuals), in a total effort of 36,679 sampling hours (36,432 trap and 247 net and observation hours). The richest families were Nymphalidae (148 species) and Hesperiidae (100 species). Most species were sampled exclusively by active methods (79.8%), but other sources (passive sampling, citizen science, etc.) also found unique records. We found the highest diversity metrics from January to May, and we demonstrated that winter months had less richness and abundance. We illustrated the 20 species common to all regions, and listed those that were found more than seven months in the year, as well as the most abundant species in trap sampling, with forest dwellers as well as species common to open and fragmented areas. The dominant species in our trap datasets was the iridescent white morpho, Morpho epistrophus (Fabricius, 1796), and we suggest it to become the park butterfly mascot.

Resumo O Sul Global testemunhou crescente amostragem de sua imensa diversidade biológica durante o século passado. Entretanto, a diversidade de muitas regiões permanece desconhecida, mesmo em locais pristinos e altamente ameaçados, como na Mata Atlântica; e para grupos bioindicadores, guarda-chuva e emblemáticos. O presente estudo reporta a primeira lista de borboletas do Parque Estadual Intervales, São Paulo, Brasil e arredores, uma unidade de conservação chave no último maciço contínuo de Mata Atlântica. Compilamos dados de museus e quatro anos de campo, sob três métodos de amostragem. Também visamos oferecer recursos para apoiar os esforços de conservação, analisando 27 anos de dados climáticos (detalhados no Material Suplementar, em inglês e em português), discutindo nossos resultados numa linguagem também para não acadêmicos, e produzindo material de divulgação científica e educativos. Um artigo irmão tratou das experiências de divulgação científica e capacitação, e ilustrou o catálogo de borboletas das espécies amostradas no parque. Encontramos 312 espécies em 2.139 registros. O museu tinha 229 espécies (432 registros), e amostramos 142 espécies (1.682 indivíduos), em um esforço total de 36.679 horas de amostragem (36.432 armadilhas e 247 horas de rede e observação). As famílias mais ricas foram Nymphalidae (148 espécies) e Hesperiidae (100 espécies). A maioria das espécies foi amostrada exclusivamente por métodos ativos (79,8%), mas outras fontes (passiva, ciência cidadã, etc.) também encontraram registros únicos. Encontramos as maiores métricas de diversidade de janeiro a maio, e demonstramos que os meses de inverno tiveram menos riqueza e abundância. Ilustramos as 20 espécies comuns a todas as regiões, e listamos aquelas que foram encontradas em mais de sete meses no ano, bem como as espécies mais abundantes em armadilhas, com espécies florestais e também comuns em áreas abertas e fragmentadas. A espécie dominante em nossas armadilhas foi a morfo branca iridescente, Morpho epistrophus (Fabricius, 1796), e sugerimos que se torne a borboleta mascote do parque.

An overview of arginussa species group of Memphis Hbner, [1819], with the description of a new species from Panama (Lepidoptera: Nymphalidae: Charaxinae).

Memphis Hbner, [1819] is a nymphalid butterfly genus exclusive to the Neotropics. It includes species with iridescent bluish or greenish coloration on the upper side of the wings while the underside is cryptic, resembling dead leaves. This paper aims to describe a remarkable new species, M. smalli Riley Dias sp. nov., from the remote Atlantic slopes of Panama based on molecular and morphological analyses. We also review the taxonomy of species herein included in the arginussa species group based on distances analyses of DNA sequence data. The arginussa species group, as defined here, includes M. arginussa (Geyer, 1832), M. eubaena (Boisduval, 1870) stat. rest., M. onophis (Felder Felder, 1861) stat. rest., M. lemons (Druce, 1877), M. neidhoeferi (Rotger, Escalante Coronado, 1965), M. perenna (Godman Salvin, [1884]), M. lankesteri (Hall, 1935) stat. rest., M. paulus Costa Orellana, 2014, M. pithyusa (Felder, 1869), M. herbacea (Butler Druce, 1872) and M. smalli sp. nov. Anaea pithyusa morena Hall, 1935 syn. nov. is recognized as a synonym of Nymphalis pithyusa Felder, 1869. The new species and its closest ally, M. herbacea, are illustrated, including characters of the head, labial palpus, wings, legs, male and female genitalia and their distribution map.

The Granulate Cutworm (Lepidoptera: Noctuidae): Biological Parameters Under Controlled Conditions, Host Plants, and Distribution in the Americas.

Feltia subterranea (Fabricius), commonly known as the granulate cutworm, is a common species of owlet moths (Noctuidae) of major agricultural importance, widely distributed in Nearctic and Neotropical regions. This study was conducted to determine the species biological parameters, gather information about its larval host plants, and assess the agricultural significance of this species in the Americas. The viability of the egg, larval, pupal stages, and prepupal period was 98, 98, and 100%, respectively, under laboratory conditions. The average duration of the egg, larval, pupal stages, and prepupal period was 3, 17, 4, and 13 d, respectively. All laboratory-reared larvae developed through five instars. The growth ratio was 1.93 for females and 1.85 for males. The duration of the larval stage was significantly longer in females than in males from the fourth instar. The duration of the pupal stage was significantly shorter in females than in males. When larval and pupal stage durations were combined, there were no significant differences in total development time as a function of sex. In total, 159 botanical taxa belonging to 41 families were recorded as host species for F. subterranea. The families with the greatest number of host species were Fabaceae (22), Poaceae (19), Asteraceae (16), Brassicaceae (13), Solanaceae (12), Amaranthaceae (7), Cucurbitaceae (7), and Malvaceae (5). It is noteworthy that the large number of native weeds used by F. subterranea as host plants could represent a significant source of infestation of crops in the agricultural landscape.

Flight over the Proto-Caribbean seaway: Phylogeny and macroevolution of Neotropical Anaeini leafwing butterflies.

Our understanding of the origin and evolution of the astonishing Neotropical biodiversity remains somewhat limited. In particular, decoupling the respective impacts of biotic and abiotic factors on the macroevolution of clades is paramount to understand biodiversity assemblage in this region. We present the first comprehensive molecular phylogeny for the Neotropical Anaeini leafwing butterflies (Nymphalidae, Charaxinae) and, applying likelihood-based methods, we test the impact of major abiotic (Andean orogeny, Central American highland orogeny, Proto-Caribbean seaway closure, Quaternary glaciations) and biotic (host plant association) factors on their macroevolution. We infer a robust phylogenetic hypothesis for the tribe despite moderate support in some derived clades. Our phylogenetic inference recovers the genus Polygrapha Staudinger, [1887] as polyphyletic, rendering the genera FountaineaRydon, 1971 and Memphis Hübner, [1819] paraphyletic. Consequently, we transfer Polygrapha tyrianthina (Salvin & Godman, 1868) comb. nov. to Fountainea and Polygrapha xenocrates (Westwood, 1850) comb. nov. to Memphis. We infer an origin of the group in the late Eocene ca. 40 million years ago in Central American lowlands which at the time were separated from South America by the Proto-Caribbean seaway. The biogeographical history of the group is very dynamic, with several oversea colonization events from Central America into the Chocó and Andean regions during intense stages of Andean orogeny. These events coincide with the emergence of an archipelagic setting between Central America and northern South America in the mid-Miocene that likely facilitated dispersal across the now-vanished Proto-Caribbean seaway. The Amazonian region also played a central role in the diversification of the Anaeini, acting both as a museum and a cradle of diversity. We recover a diversification rate shift in the Miocene within the species-rich genus Memphis. State speciation and extinction models recover a significant relationship between this rate shift and host plant association, indicating a positive role on speciation rates of a switch between Malpighiales and new plant orders. We find less support for a role of abiotic factors including the progressive Andean orogeny, Proto-Caribbean seaway closure and Quaternary glaciations. Miocene host plant shifts possibly acted in concert with abiotic and/or biotic factors to shape the diversification of Anaeini butterflies.

Resurgence of a forgotten Southern Brazil endemic species: taxonomic position, redescription, and spatio-temporal distribution of Porosagrotis carolia Schaus, 1929 (Lepidoptera: Noctuidae: Noctuinae).

The taxonomy of the hitherto unplaced taxon Porosagrotis carolia Schaus, 1929 is evaluated and the name is taken out of taxonomic limbo and combined with Feltia Walker, 1856. Feltia carolia comb. nov., described from a single female specimen, is redescribed, including the first description of the male. Head, thorax and its appendages, and male and female genitalia are illustrated through photographs and line drawings; photographs of the holotype and a distribution map for the species is provided. Additionally, the spatio-temporal occurrence of the species in Rio Grande do Sul was assessed by monthly standardized samplings with light traps from January of 1998 to December of 1999 in ten different physiographical zones of the state. Our results suggest that F. carolia comb. nov. is univoltine and goes through prepupal and pupal estival diapauses. The assessment of the distribution of the species, based on the standardized samplings and specimens deposited in entomological collections, suggests that F. carolia comb. nov. is restricted to the southern Brazilian Campos, a mostly neglected environment discontinuously distributed in mountainous plateaus of the southern Mata Atlântica biome.

Two new species of Euptychia Hübner, 1818 from the upper Amazon basin (Lepidoptera, Nymphalidae, Satyrinae).

Two new species of Euptychia Hübner, 1818 are described from the upper Amazon basin: Euptychia attenboroughi Neild, Nakahara, Fratello & Le Crom, sp. n. (type locality: Amazonas, Venezuela), and Euptychia sophiae Zacca, Nakahara, Dolibaina & Dias, sp. n. (type locality: Acre, Brazil). Their unusual facies prompted molecular and phylogenetic analyses of one of the species resulting in support for their classification in monophyletic Euptychia. Diagnostic characters for the two species are presented based on wing morphology, wing pattern, presence of androconial patches on the hindwing, and genitalia. Our results indicate that the projection of the tegumen above the uncus, previously considered a synapomorphy for Euptychia, is not shared by all species in the genus. The adults and their genitalia are documented, and distribution data and a map are provided.

ResumoDuas novas espécies de Euptychia Hübner, 1818 são descritas da bacia do alto Rio Amazonas: Euptychia attenboroughi Neild, Nakahara, Fratello & Le Crom, sp. n. (localidade tipo: Amazonas, Venezuela), Euptychia sophiae Zacca, Nakahara, Dolibaina & Dias, sp. n. (localidade tipo: Acre, Brasil). Suas aparências não usuais levaram a análises molecular e filogenética de uma das espécies que sustentam sua classificação como Euptychia. São apresentados caracteres diagnósticos para as duas espécies baseados na morfologia e no padrão de coloração de suas asas, presença de manchas androconiais nas asas posteriores e suas genitálias. Os resultados indicam que a projeção do tegume sobre o unco, previamente considerada uma sinapomorfia para Euptychia, não é compartilhada por todas as espécies do gênero. São ilustrados os adultos e suas genitálias, bem como dados e mapa de distribuição.

ResumenSe describen dos nuevas especies de Euptychia Hübner, 1818 para la cuenca alta del Río Amazonas: Euptychia attenboroughi Neild, Nakahara, Fratello & Le Crom, sp. n. (localidad típica: Amazonas, Venezuela) y Euptychia sophiae Zacca, Nakahara, Dolibaina & Dias, sp. n. (localidad típica: Acre, Brasil). Sus apariencias inusuales motivaron el análisis molecular y filogenético de una de las especies, lo cual sustenta su clasificación como Euptychia. Se presentan caracteres diagnósticos para las dos especies basados en la morfología y patrón de coloración alar, presencia de parches androconiales en las alas posteriores y sus genitales. Los resultados indican que la proyección del tegumen sobre el uncus, previamente considerada una sinapomorfía para Euptychia, no es compartida por todas las especies del género. Se ilustran los adultos y sus genitales, y se proporcionan datos y mapa de distribución.

Notes on the geographic distribution and subspecific taxonomy of Sais rosalia (Cramer) (Lepidoptera, Nymphalidae, Ithomiini), including the first records in Paraguay

Notes on the geographic distribution and subspecific taxonomy of Sais rosalia (Cramer) (Lepidoptera, Nymphalidae, Ithomiini), including the first records in Paraguay. This paper provides comments on the subspecific taxonomy and geographic distribution of Sais rosalia (Cramer, 1779) (Lepidoptera, Nymphalidae, Ithomiini), as well as an up-to-date distributional map, complemented with unpublished distributional data based on specimens deposited in the Coleção Entomológica Pe. Jesus S. Moure, Curitiba, Brazil and the Museo de Historia Natural, Lima, Peru. The following synonyms are proposed: Sais rosalia camariensis Haensch, 1905 syn. rev. as junior subjective synonym of Papilio rosalia Cramer, 1779 and Sais rosalia brasiliensis Talbot, 1928 syn. rev. as junior subjective synonym of Sais rosalia rosalinde Weymer, 1890. Additionally, the first country records of Sais rosalia in Paraguay, including the southernmost record of the species, are documented.

Biology and external morphology of immature stages of the butterfly, Diaethria candrena candrena.

The biology and the external morphology of immature stages of Diaethria candrena candrena (Godart) (Lepidoptera: Nymphalidae: Biblidinae) are described. Immature D. c. candrena found on Allophylus spp. (Sapindaceae) were collected in Curitiba, Paraná, Brazil and reared in the laboratory. Morphological descriptions and illustrations are given, based on observations using electronic, stereoscopic, and optic microscopes, the latter two attached to camera lucida. Results are compared and discussed with immature stages of other species of Biblidinae described to date.

External morphology and ultra-structure of eggs and first instar of Prepona laertes laertes (Hübner, [1811]), with notes on host plant use and taxonomy.

The external morphology and the tegument ultra-structure of Prepona laertes laertes (Hübner, [1811]) (Lepidoptera: Nymphalidae: Charaxinae) eggs and first instar larvae feeding on Inga spp. (Fabaceae) in a forest fragment in Joinville, Santa Catarina, Brazil, are described. Descriptions of the morphology with illustrations are presented, based upon observations through scanning electron microscopy and stereoscopic and optic microscopes attached to a camera lucida. Descriptions and illustrations of the head capsule, chaetotaxy, tegument, and setae are presented. The taxonomy, morphological characters, and host plant use of Prepona laertes immature stages are discussed.

Alternative techniques to study characters of the genitalia in Lepidoptera

The present note aims to describe two alternative methods for observing genitalia in Lepidoptera. The first one provides means to examine both male and female genitalia without spoiling the scales of the abdomen, preserving it attached to the thorax and aesthetically similar to an unexamined specimen. The second one provides ways of observing certain characters on the male genitalia in a non-destructive way, and does not depend on time-consuming removing and dissection of the abdomen. It is expected that the presented techniques will help on morphological studies and on identifying similar species which consistently differ in genitalic armatures.

[Biological aspects and external morphology of immature stages of Memphis moruus stheno (Prittwittz) (Lepidoptera: Nymphalidae)]. / Aspectos biológicos e morfologia externa dos imaturos de Memphismoruus stheno (Prittwitz) (Lepidoptera: Nymphalidae).

Immature stages of Memphis moruus stheno (Prittwittz) were subject of a behavior, biological and morphological study. The morphological study was carried out through observation by stereoscopic microscopy with camera lucida and micrometric lens attached for illustrations and measurements, respectively; and scanning electron microscopy for ultrastructural analysis. Behavioral data were obtained through direct observation in the field and laboratory. Eggs were laid on the underside of leaves of two Lauraceae species. The first to third instars build frass chains, the fourth and the fifth instars build a conical shelter using a single leaf of the host plant. Before pupation, larvae bend its body ventrally and the pupae are incapable of movement. Descriptions, illustrations and photographs of egg, all five instars and pupa are given. The cephalic morphology and chaetotaxy of first instar were described and illustrated. Results are compared with other Charaxinae immature stages.

Aspectos biológicos e morfologia externa dos imaturos de Memphismoruus stheno (Prittwitz) (Lepidoptera: Nymphalidae) / Biological aspects and external morphology of immature stages of Memphis moruus stheno (Prittwittz) (Lepidoptera: Nymphalidae)

Immature stages of Memphis moruus stheno (Prittwittz) were subject of a behavior, biological and morphological study. The morphological study was carried out through observation by stereoscopic microscopy with camera lucida and micrometric lens attached for illustrations and measurements, respectively; and scanning electron microscopy for ultrastructural analysis. Behavioral data were obtained through direct observation in the field and laboratory. Eggs were laid on the underside of leaves of two Lauraceae species. The first to third instars build frass chains, the fourth and the fifth instars build a conical shelter using a single leaf of the host plant. Before pupation, larvae bend its body ventrally and the pupae are incapable of movement. Descriptions, illustrations and photographs of egg, all five instars and pupa are given. The cephalic morphology and chaetotaxy of first instar were described and illustrated. Results are compared with other Charaxinae immature stages.

Alternative techniques to study characters of the genitalia in Lepidoptera.

The present note aims to describe two alternative methods for observing genitalia in Lepidoptera. The first one provides means to examine both male and female genitalia without spoiling the scales of the abdomen, preserving it attached to the thorax and aesthetically similar to an unexamined specimen. The second one provides ways of observing certain characters on the male genitalia in a non-destructive way, and does not depend on time-consuming removing and dissection of the abdomen. It is expected that the presented techniques will help on morphological studies and on identifying similar species which consistently differ in genitalic armatures.

Morfologia do exoesqueleto de adultos de Memphis moruus stheno (Pritwittz) (Lepidoptera, Nymphalidae, Charaxinae) / Exoskeleton morphology of the adult of Memphis moruus stheno (Pritwittz) (Lepidoptera, Nymphalidae, Charaxinae)

O presente trabalho visa fornecer subsídios para estudos morfológicos comparativos de Charaxinae neotropicais. Memphis moruus stheno (Prittwitz, 1865) é a segunda espécie neotropical de Anaeini e a primeira do gênero Memphis a ter sua morfologia detalhada. A morfologia externa é apresentada com descrições, ilustrações e imagens de microscopia eletrônica de varredura de estruturas da cabeça e apêndices cefálicos, região cervical, tórax e apêndices torácicos, abdome e genitálias masculina e feminina. A morfologia de M. moruus stheno é comparada com uma espécie do gênero proximamente relacionado Zaretis Hübner, [1819].

This study is intended to allow further comparative morphological studies on the Neotropical Charaxinae. Memphis moruus stheno (Prittwitz, 1865) is the second species of Anaeini and the first in the genus Memphis to have its morphology published in detail. The external morphology is presented with descriptions, line drawings and scan electron microscopy of the head and its appendages, cervix, thorax and its appendages, abdomen and both male and female genital armatures. Memphis moruus stheno is compared with a species of its closely related genus Zaretis Hübner, [1819].