Systematics & Taxonomy Research
- Blood-feeding and Fruit-feeding Moths of Calpini Tribe (Noctuidae), Their Phylogeny and Classification
- DNA barcoding of Palearctic butterflies
- Higher Classification of Hesperiidae
- Systematics of Genus Calisto
- Systematics and Classification of Ithomiinae
- Taxonomy of Monarch butterflies
- Phylogenetics and other projects - Kawahara Lab
Blood-feeding and Fruit-feeding Moths of Calpini Tribe (Noctuidae), Their Phylogeny and Classification
The noctuid moth subfamily Calpinae is comprised of about 360 species worldwide. Calpinae is defined by the structure of the proboscis which consists of fish hook-like barbs used for tearing the skin of fruits and mammals. This subfamily consists of four tribes: Anomini, Calpini, Phyllodini, and Scoliopterygini. Calpini consists of about 200 species in the following genera: Calyptra, Cecharismena, Eudocima, Goniapteryx, Gonodonta, Hypsoropha, Oraesia, Pharga, Phyprosopus, Plusiodonta, Psammathodoxa and Radara.
The tribe Calpini is cosmopolitan in its distribution. The genus Calyptra is considered to be Old World in its distribution with a high concentration of diversity in South and Southeast Asia, yet one species, C. canadensis, occurs in the northeastern United States and Canada. Genera Cecharismena, Goniapteryx, Hypsoropha, Pharga, Phyprosopus, and Psammathodoxa are mainly found in the New World, while Eudocima is found in the Old World tropics. Genus Gonodonta can be found in subtropical and tropical regions, with seven species occurring in Arizona, Florida and Texas. Species of Oraesia, Plusiodonta, and Radara are common in the Old and New World tropics.
At least five genera within the Calpini are considered to be primary piercers of both hard and soft-skinned fruits; nine species in the genus Calyptra have been recorded piercing the skin of mammals and feeding on their blood. Bänziger divides these feeding behaviors into three categories: skin piercers and blood feeders, primary fruit piercers and secondary fruit piercers. Primary fruit piercers are able to penetrate fruit, while secondary piercers are only capable of piercing fruit damaged previously by primary piercers or other animals.
It has been hypothesized that blood-feeding behavior evolved from fruit piercing. This hypothesis has never been tested, and cannot be tested until the relationships of Calyptra and related genera are known. McGuire Center doctoral researcher Jennifer Zaspel helped reconstruct the phylogenetic relationships among the genera in Calpini. She also learned valuable information on the origin(s) of blood feeding in the genus Calyptra and if there is, in fact, a directional progression of feeding types in these moths.
DNA barcoding of Palearctic butterflies
Higher Classification of Hesperiidae
Butterflies in the family Hesperiidae, commonly known as skippers, comprise between 4,000 and 5,000 species worldwide, constituting about 20 percent of the Earth's butterfly fauna. Despite this diversity, the family has historically been under-studied, compared to the other families of butterflies, and there has been no general consensus on the definitions of or relationships between higher taxa in the family.
Recent studies on the higher classification of the family, conducted by Andrew Warren and colleagues, have sought to define the major genealogical lineages (subfamilies and tribes) of the world's skippers, through combined analyses of adult morphology and DNA sequence character data.
The results support the recognition of seven subfamilies of Hesperiidae, with the following relationships: (Coeliadinae, (Euschemoninae, (Eudaminae, (Pyrginae, (Heteropterinae, (Trapezitinae, Hesperiinae)))))).
The subfamily Pyrginae can be divided into seven tribes, as follows: ((Celaenorrhinini, (Pyrrhopygini, Tagiadini)), (Carcharodini, (Erynnini, (Achlyodidini, Pyrgini)))). The tribal classification of the Hesperiidae is not as well supported as that of the Pyrginae, and will require the addition of more characters and taxa to fully resolve. Aeromachini reside at the base of the Hesperiinae, followed by a paraphyletic grade of mostly Old World taxa, but also including Megathymini (Giant Skippers) and New World genera such as Perichares and Orses. Terminal clades of Hesperiinae are better supported, and include: (Baorini, (Taractrocerini, (Thymelicini, (Calpodini, (new tribe, (Hesperiinae, new tribe)))))).
We are continually seeking to expand our data matrix through the addition of more taxa, and eventually hope to include molecular and morphological character data for all genera of Hesperiidae.
Systematics of Genus Calisto
Butterfly species in the genus Calisto are highly successful, but are now seriously endangered due to habitat loss. Of the 103 butterfly genera on Hispaniola, the genus Calisto comprises 20 percent of the butterfly fauna. This genus is an extraordinary example of butterfly radiation (their spread and evolving diversity). There are some 40 Calisto species and subspecies that have adapted to every possible environment on the island, from lowland desert where temperatures reach 120 degrees, to 10,000-foot-high mountains where frost covers the ground at night.
Calisto have extremely local ranges, and many species are rare and endangered. In the exceptionally dry Hispaniolan lowlands, four Calisto species survive on seemingly unpalatable Bunch Grass (Uniola virgata). In the highlands, several isolated species are associated with various bamboos. Some Calisto are known from just several specimens, and nothing is yet known about their biology. Many Calisto species are endangered and will probably go extinct in coming decades. There is little protection of their habitats, and the few national parks suffer from illegal logging, grazing and agriculture. Andrei Sourakov conducted most of the recent Calisto research. His work described the biology of many species and reconstructs their evolutionary history.
Systematics and Classification of Ithomiinae
The nymphalid butterfly subfamily Ithomiinae contains some 360 species occurring only in the neotropical region, from the southwestern U.S.A. to Uruguay. Closely related to the subfamily Danainae (milkweed butterflies), it represents one of the largest neotropical butterfly radiations. Ithomiine biology has been extensively studied for many years, and currently these butterflies are a model group for research into biogeography, evolution and genetics.
Despite much work on ithomiine systematics over the last 50 years, until recently there existed no phylogeny, or hypothesis of evolutionary relationships, between ithomiine species and genera. McGuire Center curator Keith Willmott is working with colleagues in the U.S. and Brazil to study the morphology and genetics of these butterflies to derive a phylogeny for all genera and species groups. This phylogeny will be used to check the existing higher-level classification (tribes, genera) and propose changes where necessary. He is also working with Gerardo Lamas, from the Museo de Historia Natural, Universidad Nacional Mayor de San Marcos in Lima, to revise the systematics and classification of all ithomiine genera that have not been studied in the past 50 years. This work involves about 200 species, of which about 10 percent are undescribed. Results should provide a stable framework for testing evolutionary hypotheses in this subfamily and for further studies of their ecology and biology in the field.
Learn more: Illustrated Checklist of Ithomiinae
Taxonomy of Monarch butterflies
Most insect revisions and classifications are based mainly on the characters of adult insects. However, when the taxonomic status of a taxon is in question, molecular data and/or hybridization crosses can be employed. In order to clarify the taxonomic status of Danaus erippus, which has often been classified as a subspecies of Monarch butterfly, Danaus plexippus, hybridization studies were performed by Mirian Medina Hay-Roe. The results revealed the presence of pre- and postzygotic isolation between these two taxa. Pupal inviability and Haldane rule effects (male hybrids only, the homogametic sex in butterflies) were also observed. These results reinforce the hypothesis that plexippus and erippus are separate, reproductively isolated species. Chemical analysis of their cuticular hydrocarbons shows quantitative differences which may imply reproductive isolation between these two species.