The Frog-Biting Midges of the World (Corethrellidae: Diptera)

This worldwide biosystematic study of Corethrellidae, with its single genus Corethrella Coquillett, provides a complete compilation of all that is known for the group, both taxonomically and bionomically. Descriptions of each species are based primarily on the adults, summarize all bionomic information, and provide a map showing its distribution. Keys to the species of each region are provided. A total of 97 extant species is recognized, with 52 of these being new. Seven fossil species are described with two of these being new to science. All species, including 13 new synonyms, are cataloged in a table for easy reference. Seven extant species are of uncertain status because of damaged or missing types. Lectotypes and, depending on the species, some paralectotypes, are designated for the following species: C. inepta (Annandale), C. pallitarsis Edwards, C. picticollis Edwards, C. ananacola Dyar, C. calathicola Edwards, and C. brakeleyi (Coquillett). A cladistic analysis interprets most extant and fossil species (some are not interpretable at the present time) and provides the basis for zoogeographic and bionomic interpretation. Worldwide, Corethrella species are found between 50°N and 50°S but most are found between 30°N and 30°S and below 1500 meters in elevation. Because female adults are attracted to the call of male frogs and feed on their blood, species are restricted to areas where there are frogs. Phylogenetic patterns suggest Gondwanan connections for earlier lineages within the genus. At least one lineage has dispersed from the New World to southeast Asia and some species are located on volcanic islands in the Caribbean, indicating further instances of dispersal. It is certain that many more species are yet to be discovered. Phylogenetic patterns indicate that the immatures of Corethrella species have repeatedly moved between ground-dwelling habitats and phytotelmata, with the plesiotypic habitat likely being ground-dwelling. Some lineages have diversified within phytotelmata. Fossil, cladistic and morphological evidence indicates that Corethrella females have been feeding on calling frogs since at least the Early Cretaceous. Females likely hear their frog hosts using the Johnston’s Organ. There is some evidence of host specificity as well as selection of particular biting sites for some species of Corethrella. The females of at least some species of Corethrella transmit Trypanosoma Gruby between calling frogs and this association is also likely an ancient one.Este estudio biosistematico de Corethrellidae a nivel Mundial, con su unico genero Corethrella, proporciona una completa recopilacion de todo lo conocido para el grupo, tanto desde el punto de vista taxonomico como bionomico. Las descripciones de cada especie se realizan primariamente sobre la base de adultos, resumen toda la informacion bionomica y proveen un mapa donde se muestra su distribucion. Se brindan claves para especie de cada region. Se reconoce un total de 97 especies, 52 de las cuales son nuevas. Se describen siete especies fosiles, siendo dos de ellas nuevas para la Ciencia. Para una facil referencia, todas las especies son catalogadas en una tabla, incluyendo 13 nuevos sinonimos. Debido a que sus tipos se hallan danados o perdidos, siete especies actuales ostentan un status incierto. De acuerdo a la especie, son designados lectotipos o paralectotipos de las siguientes especies: C. inepta (Annandale), C. pallitarsis Edwards, C. picticollis Edwards, C. ananacola Dyar, C. calathicola Edwards, y C. brakeleyi (Coquillett). El analisis cladistico interpreta la mayoria de las especies actuales y fosiles (algunas no pueden ser interpretadas actualmente) y provee la base para interpretaciones zoogeograficas y bionomicas. Las especies de Corethrella se hallan entre 50°N y 50°S, aunque la mayoria se encuentran entre 30°N y 30°S y por debajo de 1500 metros de elevacion. Debido a que las hembras adultas son atraidas por el llamado de ranas macho y se alimentan de su sangre, las especies estan restringidas a las areas donde se hallan ranas. Los patrones filogeneticos sugieren conecciones Gondwanicas para los linajes mas antiguos del genero. Al menos un linaje se ha dispersado desde el Nuevo Mundo hacia el sudeste de Asia, y algunas especies se hallan en islas volcanicas del Caribe, indicando otras instancias de dispersion. Con seguridad aun quedan muchas mas especies por ser descubiertas. Los patrones filogeneticos indican que los inmaduros de las especies de Corethrella se han movido repetidamente entre habitats ubicados a nivel del suelo y fitotelmata, siendo probablemente