Comprehensive phylogeny of Pieridae butterflies reveals strong correlation between diversification and temperature.

Temperature is thought to be a key factor influencing global species richness patterns. We investigate the link between temperature and diversification in the butterfly family Pieridae by combining next generation DNA sequences and published molecular data with fine-grained distribution data. We sampled nearly 600 pierid butterfly species to infer the most comprehensive molecular phylogeny of the family and curated a distribution dataset of more than 800,000 occurrences. We found strong evidence that species in environments with more stable daily temperatures or cooler maximum temperatures in the warm seasons have higher speciation rates. Furthermore, speciation and extinction rates decreased in tandem with global temperatures through geological time, resulting in a constant net diversification.

Anurocampa (Lepidoptera: Notodontidae): two new species, systematics and immature stages.

Notodontidae (Lepidoptera, Noctuoidea) comprise over 4,000 described species distributed worldwide, among which nearly half are restricted to the Neotropics. Morphology of adults and immatures of Notodontidae have been broadly investigated and many larval, pupal, and adult characters were found to be synapomorphies of subfamilies and tribes. Despite this, the current classification of Notodontidae remains unsettled as most recent classification systems are contradictory due to reliance on incomplete global sampling and, many taxa, especially in the Neotropics, are still informally classified as incertae sedis. Anurocampa Herrich-Shäffer was recently treated as an incertae sedis genus, and immature and adult characters may provide further evidence for its systematic position among the Notodontidae. With this goal in mind, the present study describes the immature stages of Anurocampa mingens Herrich-Shäffer from Brazil and describes two new species in the genus from Costa Rica based on morphology and mitochondrial DNA: Anurocampa markhastingsi Chacón and St Laurent sp. nov. and Anurocampa abelardochaconi Chacón and St Laurent sp. nov. and discusses the systematic position of Anurocampa.

Prominent moths (Lepidoptera: Notodontidae) of Colombia.

Based on literature review, curatorial work in national and international collections, information available on online databases, and field work conducted from 2017 to 2022, we provide the first checklist for the family Notodontidae in Colombia. A total of 515 species (51 endemics), 122 genera, 7 subfamilies as well as 108 new records for Colombia are presented. These data position Colombia as the country with the third highest diversity of prominent moths in the world, based on current knowledge. Furthermore, 239 known hostplants are listed for 91 species. The distribution map of the family shows the highest concentration of national records in the Andean and Pacific regions. Future surveys in less explored zones, long-term monitoring, expansion and maintenance of biological collections, and working collaboratively with communities will surely increase the known diversity as well as the conservation of prominent moths in Colombia.

A global phylogeny of butterflies reveals their evolutionary history, ancestral hosts and biogeographic origins.

Butterflies are a diverse and charismatic insect group that are thought to have evolved with plants and dispersed throughout the world in response to key geological events. However, these hypotheses have not been extensively tested because a comprehensive phylogenetic framework and datasets for butterfly larval hosts and global distributions are lacking. We sequenced 391 genes from nearly 2,300 butterfly species, sampled from 90 countries and 28 specimen collections, to reconstruct a new phylogenomic tree of butterflies representing 92% of all genera. Our phylogeny has strong support for nearly all nodes and demonstrates that at least 36 butterfly tribes require reclassification. Divergence time analyses imply an origin ~100 million years ago for butterflies and indicate that all but one family were present before the K/Pg extinction event. We aggregated larval host datasets and global distribution records and found that butterflies are likely to have first fed on Fabaceae and originated in what is now the Americas. Soon after the Cretaceous Thermal Maximum, butterflies crossed Beringia and diversified in the Palaeotropics. Our results also reveal that most butterfly species are specialists that feed on only one larval host plant family. However, generalist butterflies that consume two or more plant families usually feed on closely related plants.

A diversification relay race from Caribbean-Mesoamerica to the Andes: historical biogeography of Xylophanes hawkmoths.

The regions of the Andes and Caribbean-Mesoamerica are both hypothesized to be the cradle for many Neotropical lineages, but few studies have fully investigated the dynamics and interactions between Neotropical bioregions. The New World hawkmoth genus Xylophanes is the most taxonomically diverse genus in the Sphingidae, with the highest endemism and richness in the Andes and Caribbean-Mesoamerica. We integrated phylogenomic and DNA barcode data and generated the first time-calibrated tree for this genus, covering 93.8% of the species diversity. We used event-based likelihood ancestral area estimation and biogeographic stochastic mapping to examine the speciation and dispersal dynamics of Xylophanes across bioregions. We also used trait-dependent diversification models to compare speciation and extinction rates of lineages associated with different bioregions. Our results indicate that Xylophanes originated in Caribbean-Mesoamerica in the Late Miocene, and immediately diverged into five major clades. The current species diversity and distribution of Xylophanes can be explained by two consecutive phases. In the first phase, the highest Xylophanes speciation and emigration rates occurred in the Caribbean-Mesoamerica, and the highest immigration rates occurred in the Andes, whereas in the second phase the highest immigration rates were found in Amazonia, and the Andes had the highest speciation and emigration rates.

Phylogeny of gracillariid leaf-mining moths: evolution of larval behaviour inferred from phylogenomic and Sanger data.

Gracillariidae is the most taxonomically diverse cosmopolitan leaf-mining moth family, consisting of nearly 2000 named species in 105 described genera, classified into eight extant subfamilies. The majority of gracillariid species are internal plant feeders as larvae, creating mines and galls in plant tissue. Despite their diversity and ecological adaptations, their phylogenetic relationships, especially among subfamilies, remain uncertain. Genomic data (83 taxa, 589 loci) were integrated with Sanger data (130 taxa, 22 loci), to reconstruct a phylogeny of Gracillariidae. Based on analyses of both datasets combined and analyzed separately, monophyly of Gracillariidae and all its subfamilies, monophyly of the clade "LAMPO" (subfamilies: Lithocolletinae, Acrocercopinae, Marmarinae, Phyllocnistinae, and Oecophyllembiinae) and relationships of its subclade "AMO" (subfamilies: Acrocercopinae, Marmarinae, and Oecophyllembiinae) were strongly supported. A sister-group relationship of Ornixolinae to the remainder of the family, and a monophyletic leaf roller lineage (Callicercops Vári + Parornichinae) + Gracillariinae, as sister to the "LAMPO" clade were supported by the most likely tree. Dating analyses indicate a mid-Cretaceous (105.3 Ma) origin of the family, followed by a rapid diversification into the nine subfamilies predating the Cretaceous-Palaeogene extinction. We hypothesize that advanced larval behaviours, such as making keeled or tentiform blotch mines, rolling leaves and galling, allowed gracillariids to better avoid larval parasitoids allowing them to further diversify. Finally, we stabilize the classification by formally re-establishing the subfamily ranks of Marmarinae stat.rev., Oecophyllembiinae stat.rev. and Parornichinae stat.rev., and erect a new subfamily, Callicercopinae Li, Ohshima and Kawahara to accommodate the enigmatic genus Callicercops.

The Bombycoidea and Mimallonoidea (Lepidoptera) of a research station in the grasslands of Southern Brazil.

We report the Bombycoidea and Mimallonoidea (Lepidoptera) observed at the the Universidade Federal do Rio Grande do Suls (UFRGS) Agronomic Research Station in Eldorado do Sul, Rio Grande do Sul, Brazil from September 7th to November 6th, 2018. This research station is situated in the Campos Sulinos (southern Brazilian grasslands) environment. As part of our inventory, we report the following Bombycoidea: 29 species of Sphingidae, 26 species of Saturniidae, at least seven species of Apatelodidae, and at least four species of Bombycidae. Within Mimallonidae we recorded eight species. New state records include the following Sphingidae: Xylophanes alineae, X. crenulata, and Perigonia stulta; and Mimallonidae: Mimallo grisea and the genus Menevia. Our study represents the first inventory of moths from this location and one of the few from the Campos Sulinos more broadly.

Food Plant Shifts Drive the Diversification of Sack-Bearer Moths.

AbstractLepidoptera are a highly diverse group of herbivorous insects; however, some superfamilies have relatively few species. Two alternative hypotheses for drivers of Lepidoptera diversity are shifts in food plant use or shifts from concealed to external feeding as larvae. Many studies address the former hypothesis but with bias toward externally feeding taxa. One of the most striking examples of species disparity between sister lineages in Lepidoptera is between the concealed-feeding sack-bearer moths (Mimallonoidea), which contain about 300 species, and externally feeding Macroheterocera, which have over 74,000 species. We provide the first dated tree of Mimallonidae to understand the diversification dynamics of these moths in order to fill a knowledge gap pertaining to drivers of diversity within an important concealed-feeding clade. We find that Mimallonidae is an ancient Lepidoptera lineage that originated in the Cretaceous ∼105 million years ago and has had a close association with the plant order Myrtales for the past 40 million years. Diversification dynamics are tightly linked with food plant usage in this group. Reliance on Myrtales may have influenced diversification of Mimallonidae because clades that shifted away from the ancestral condition of feeding on Myrtales have the highest speciation rates in the family.

Is Sexual Conflict a Driver of Speciation? A Case Study With a Tribe of Brush-footed Butterflies.

Understanding the evolutionary mechanisms governing the uneven distribution of species richness across the tree of life is a great challenge in biology. Scientists have long argued that sexual conflict is a key driver of speciation. This hypothesis, however, has been highly debated in light of empirical evidence. Recent advances in the study of macroevolution make it possible to test this hypothesis with more data and increased accuracy. In the present study, we use phylogenomics combined with four different diversification rate analytical approaches to test whether sexual conflict is a driver of speciation in brush-footed butterflies of the tribe Acraeini. The presence of a sphragis, an external mating plug found in most species among Acraeini, was used as a proxy for sexual conflict. Diversification analyses statistically rejected the hypothesis that sexual conflict is associated with shifts in diversification rates in Acraeini. This result contrasts with earlier studies and suggests that the underlying mechanisms driving diversification are more complex than previously considered. In the case of butterflies, natural history traits acting in concert with abiotic factors possibly play a stronger role in triggering speciation than does sexual conflict. [Acraeini butterflies; arms race; exon capture phylogenomics; Lepidoptera macroevolution; sexual selection; sphragis.].

The Wild Silkmoths (Lepidoptera: Bombycoidea: Saturniidae) of Colombia: a database of occurrence points and taxonomic checklist.

Based on the review of literature and biological collections, information retrieved from public online databases, and from fieldwork conducted between 2015 and 2020, we provide more than 3500 occurrence records and an updated checklist of Colombian Saturniidae, annotated with distribution data for all species. In the first checklist of Colombian saturniids published two decades ago, a total of 184 species were cited; in the current update the number has risen to 653 species/subspecies classified in 55 genera, representing all six recognized Neotropical subfamilies. The Andean and Pacific regions are the richest, accounting for about three quarters of all species listed herein. We call attention to the fact that for most speciose saturniid genera in the Neotropics, there remains a significant need for further taxonomic and systematic research with objective and integrative approaches. Likewise, it is necessary to continue the study of Saturniidae moths in Colombia, strategically increasing the sampling efforts in specific under-sampled natural regions to further document the diversity of this family in the country.

Revision of Roelofa Schaus, 1928 (Lepidoptera, Mimallonidae, Roelofinae) with a description of a new species.

The mimallonid genus Roelofa is revised. It is the only genus belonging to the recently erected subfamily, Roelofinae.Both sexes and genitalia are figured for all previously described species in the genus, and all are redescribed. The species Roelofa maera stat. rev. is no longer considered a synonym of R. narga based on morphological differences. We describe a new species from Guatemala: R. monzoni St Laurent, Herbin Kawahara, sp. n. which is most similar to the widespread Central American R. hegewischi.

A new species of Cicinnus Blanchard (Lepidoptera, Mimallonidae, Cicinninae) from the mangrove ecoregions of Brazil.

A new species of Cicinnus from northern coastal Brazil is described and illustrated: C. litoralis sp. n.. The species is known from five males collected in the mangrove ecoregions of the Brazilian states of Pará and Maranhão (the Pará mangrove and Maranhão mangrove ecoregions respectively). This area of Brazil is poorly sampled for Mimallonidae, and the new species represents the first mimallonid considered endemic to these ecoregions. We also provide a discussion regarding the type species of Cicinnus, C. orthane Blanchard, focusing on populations from the Cerrado and the Atlantic Forest of Brazil.

Cicinnus chambersi: a new species of sack-bearer moth (Lepidoptera, Mimallonidae, Cicinninae) from southeastern Arizona, USA.

A new species of cicinnine Mimallonidae, Cicinnus chambersi sp. nov., is described from the Sky Islands Region of southern Arizona, USA. The new species is closely related to C. mexicana (Druce), type locality Veracruz, Mexico, based on morphology and genetics. The other Cicinnus species known from the United States, the common C. melsheimeri (type locality Pennsylvania, USA) is morphologically and genetically distinct from both C. chambersi and C. mexicana. The new species is compared to C. mexicana and C. melsheimeri, as well as other Mexican Cicinnus. The life history of C. chambersi is unknown, but its description should facilitate future studies on this rarely reported North American mimallonid, a species which may have only recently become established in the United States. Cicinnus chambersi is the fifth known Mimallonidae species from the United States, and the first described from the country in nearly half a century.

New species of Procinnus Herbin and Micrallo St Laurent and C. Mielke (Lepidoptera, Mimallonidae, Druenticinae) from the Brazilian Cerrado.

Two new species of Mimallonidae are described from the Brazilian Cerrado. One species belongs to Procinnus Herbin, 2016 and the second to Micrallo St Laurent and C. Mielke, 2016, both considered to belong to Druenticinae St Laurent and Kawahara, 2018. Procinnus incanus sp. n. is described from Bahia, Goiás, Distrito Federal, Minas Gerais, and São Paulo and is characterized by its wing habitus and male genitalia. Micrallo macro sp. n., the second species to be placed in Micrallo, is described from Minas Gerais. It differs from M. minutus St Laurent and C. Mielke, 2016 by larger size and distinct genitalia of both the male and female. All taxa involved are figured along with their genitalia. Holotypes designated here are deposited in the Coll. Padre Jesus S. Moure, in the Departamento de Zoologia, Universidade Federal do Paraná, Curitiba, Paraná, Brazil.

Reclassification of the Sack-bearer Moths (Lepidoptera, Mimallonoidea, Mimallonidae).

A backbone molecular phylogeny of Mimallonidae, based on 47 species and 515 loci, was recently published. That study resolved some of the major relationships in the family, but taxon sampling was limited and a classification of the family was not formally presented for all species. Here morphological phylogenetic analyses in parsimony and maximum likelihood (ML) frameworks were conducted that included 192 species and 55 morphological characters. A phylogenetic analysis was also conducted on the morphological dataset with a topological constraint based on the 515 locus tree from the previous study. Results show that nearly all species can be confidently placed in a genus using morphological phylogenetics. The presence of a frenulum, a character that was historically used to distinguish major groups of Mimallonidae, varies within and among genera. Based on our phylogenetic results, the classification of Mimallonidae is revised, which now includes 291 species in 41 genera. Descriptions of three new genera are included: Fatellalla gen. n., Citralla gen. n., and Lepismalla gen. n. The following taxonomic changes were made in the present article: 43 new/revived combinations (in Aceclostria Vuillot, Arcinnus Herbin, Bedosia Schaus, Bedosiallo St Laurent & Kawahara, Cicinnus Blanchard, Citralla, Druentica Strand, Fatellalla, Lacosoma Grote, Lepismalla, Mimallo Hübner, Procinnus Herbin, Psychocampa Grote, Roelmana Schaus, and Thaelia Herbin), two new species-level synonyms (melini Bryk is synonymized with viemanda Schaus, jaruga Jones is synonymized with hamata Walker), one revived synonymy (roscida Dognin is resynonymized with externa Moore), seven new statuses (in Druentica, Macessoga Schaus, and Trogoptera Herrich-Schäffer), six revived statuses (in Aceclostria, Cicinnus, Druentica, Psychocampa, and Zaphanta Dyar), and one new designation of nomen nudum. In order to alleviate nomenclatural problems, twelve lectotypes are designated (for Tolypidaamaryllis (Schaus), Trogopteraalthora Schaus, Adalgisacroesa Schaus, Alheitapulloides (Dognin), LacosomabriasiaSchaus, Lacosomadiederica Schaus, Lacosomaraydela Schaus, Psychocampalacuna (Schaus), Cicinnuscorallina Dognin, Cicinnuslatris Schaus, Cicinnussolvens Schaus, Cicinnustuisana Schaus) as well as a neotype for Mimallodespecta Walker (= Cicinnusdespecta). This paper also provides apomorphies for each genus and a morphological key to genera. Annotations are given to aid researchers in understanding all changes made herein, and images of male and female and their genitalia are present for nearly all type species.

The identity of Cicinnus orthane Blanchard, 1852 (Lepidoptera, Mimallonidae, Cicinninae), type species of Cicinnus Blanchard, 1852.

In order to clarify the identity of the type species of one of the most diverse Mimallonidae genera, Cicinnus Blanchard, 1852, we designate a lectotype for Cicinnus orthane Blanchard, 1852 with photographs of the lectotype (dorsal and ventral) and its genitalia figured for the first time. Cicinnus orthane, described from Chile, is a close morphological match for a southeastern Brazilian species, which we also illustrate for comparative purposes. Because the lectotype of C. orthane is female, we include both sexes of this related Brazilian species in order to facilitate the figuring of male genitalia as well as comparisons of the female genitalia. We are therefore able to establish sound morphological characteristics for Cicinnus sensu stricto, namely in the males the bifid configuration of the gnathos, a complex juxtal arrangement fused to the phallus, and largely membranous valvae; and in the females a wide and narrow lamella antevaginalis, lack of setae covered bulbous masses on either side of the lamella antevaginalis, well-sclerotized and posteriorly protruding tergite VIII, and reduced, stout apophyses anteriores which are about one quarter the length of the apophyses posteriores.

Phylogenetics of moth-like butterflies (Papilionoidea: Hedylidae) based on a new 13-locus target capture probe set.

The Neotropical moth-like butterflies (Hedylidae) are perhaps the most unusual butterfly family. In addition to being species-poor, this family is predominantly nocturnal and has anti-bat ultrasound hearing organs. Evolutionary relationships among the 36 described species are largely unexplored. A new, target capture, anchored hybrid enrichment probe set ('BUTTERFLY2.0') was developed to infer relationships of hedylids and some of their butterfly relatives. The probe set includes 13 genes that have historically been used in butterfly phylogenetics. Our dataset comprised of up to 10,898 aligned base pairs from 22 hedylid species and 19 outgroups. Eleven of the thirteen loci were successfully captured from all samples, and the remaining loci were captured from ≥94% of samples. The inferred phylogeny was consistent with recent molecular studies by placing Hedylidae sister to Hesperiidae, and the tree had robust support for 80% of nodes. Our results are also consistent with morphological studies, with Macrosoma tipulata as the sister species to all remaining hedylids, followed by M. semiermis sister to the remaining species in the genus. We tested the hypothesis that nocturnality evolved once from diurnality in Hedylidae, and demonstrate that the ancestral condition was likely diurnal, with a shift to nocturnality early in the diversification of this family. The BUTTERFLY2.0 probe set includes standard butterfly phylogenetics markers, captures sequences from decades-old museum specimens, and is a cost-effective technique to infer phylogenetic relationships of the butterfly tree of life.

A global checklist of the Bombycoidea (Insecta: Lepidoptera).

BACKGROUND: Bombycoidea is an ecologically diverse and speciose superfamily of Lepidoptera. The superfamily includes many model organisms, but the taxonomy and classification of the superfamily has remained largely in disarray. Here we present a global checklist of Bombycoidea. Following Zwick (2008) and Zwick et al. (2011), ten families are recognized: Anthelidae, Apatelodidae, Bombycidae, Brahmaeidae, Carthaeidae, Endromidae, Eupterotidae, Phiditiidae, Saturniidae and Sphingidae. The former families Lemoniidae and Mirinidae are included within Brahmaeidae and Endromidae respectively. The former bombycid subfamilies Oberthueriinae and Prismostictinae are also treated as synonyms of Endromidae, and the former bombycine subfamilies Apatelodinae and Phitditiinae are treated as families. NEW INFORMATION: This checklist represents the first effort to synthesize the current taxonomic treatment of the entire superfamily. It includes 12,159 names and references to their authors, and it accounts for the recent burst in species and subspecies descriptions within family Saturniidae (ca. 1,500 within the past 10 years) and to a lesser extent in Sphingidae (ca. 250 species over the same period). The changes to the higher classification of Saturniidae proposed by Nässig et al. (2015) are rejected as premature and unnecessary. The new tribes, subtribes and genera described by Cooper (2002) are here treated as junior synonyms. We also present a new higher classification of Sphingidae, based on Kawahara et al. (2009), Barber and Kawahara (2013) and a more recent phylogenomic study by Breinholt et al. (2017), as well as a reviewed genus and species level classification, as documented by Kitching (2018).

Revision of the genus Reinmara Schaus, 1928 (Lepidoptera, Mimallonoidea, Mimallonidae) with the descriptions of four new species from South America.

The mimallonid genus Reinmara Schaus, 1928 is revised. The three previously described species, R. enthona (Schaus, 1905), R. minasa Schaus, 1928, and R. wolfei Herbin & C. Mielke, 2014 are redescribed and the females of each are described and figured for the first time. Additionally, we describe four new species, two Andean: R. andensissp. n. and R. occidentalissp. n., and two Brazilian: R. atlanticasp. n. and R. igneasp. n.. The new species R. ignea and R. atlantica are likely of conservation concern due to their rarity in collections and their apparent endemism to an endangered biome, the Brazilian Atlantic Forest.

The identity of Arhodia egenaria Walker, 1866 (Lepidoptera, Mimallonoidea, Mimallonidae) and a new synonym of Cicinnus melsheimeri (Harris, 1841).

The holotypes of Arhodia egenaria Walker, 1866 and Cicinnus primolus Schaus, 1928, syn. n., were examined. Both names are junior synonyms of C. melsheimeri (Harris, 1841). Cicinnus melsheimeri (as Perophora egenaria), sensu Hampson, 1904, is a misidentification of C. bahamensis St Laurent & McCabe, 2016.