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 biosistemático de Corethrellidae a nivel Mundial, con su único género Corethrella, proporciona una completa recopilación de todo lo conocido para el grupo, tanto desde el punto de vista taxonómico como bionómico. Las descripciones de cada especie se realizan primariamente sobre la base de adultos, resumen toda la información bionómica y proveen un mapa donde se muestra su distribución. Se brindan claves para especie de cada región. Se reconoce un total de 97 especies, 52 de las cuales son nuevas. Se describen siete especies fósiles, siendo dos de ellas nuevas para la Ciencia. Para una fácil referencia, todas las especies son catalogadas en una tabla, incluyendo 13 nuevos sinónimos. Debido a que sus tipos se hallan dañados 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 análisis cladístico interpreta la mayoría de las especies actuales y fósiles (algunas no pueden ser interpretadas actualmente) y provee la base para interpretaciones zoogeográficas y bionómicas. Las especies de Corethrella se hallan entre 50°N y 50°S, aunque la mayoría se encuentran entre 30°N y 30°S y por debajo de 1500 metros de elevación. Debido a que las hembras adultas son atraídas por el llamado de ranas macho y se alimentan de su sangre, las especies están restringidas a las áreas donde se hallan ranas. Los patrones filogenéticos sugieren conecciones Gondwánicas para los linajes más antiguos del género. Al menos un linaje se ha dispersado desde el Nuevo Mundo hacia el sudeste de Asia, y algunas especies se hallan en islas volcánicas del Caribe, indicando otras instancias de dispersión. Con seguridad aún quedan muchas más especies por ser descubiertas. Los patrones filogenéticos indican que los inmaduros de las especies de Corethrella se han movido repetidamente entre habitats ubicados a nivel del suelo y fitotelmata, siendo probablemente
This catalog of the biting midges (Diptera: Ceratopogonidae) of the New World north of Mexico includes 603 species and 39 genera. The full citation, province or state of the type locality and distribution by province and/or state of each species is given. The most recent revision for each genus, subgenus and species group is cited.
Distribution and habitat preferences of species of Chaoboridae in the Holarctic Region are described. Coexistence of larvae of different Chaoborus species in various water bodies reflects differences in geographical distribution, habitat preference, and phyletic relationships. Extant sister species with similar habitat preferences are allopatric whereas those with differences may be sympatric. This suggests that the development of ecological divergence must historically precede coexistence of the immatures of species. Species which coexist often differ markedly morphologically and are not immediately related phylogenetically.Corethrella and Chaoborus (Sayomyia) species that occur in the Holarctic Region and Chaoborus pallidus may have southern, tropical phyletic affinities. Distributions of some North American chaoborids suggest the presence of ecological barriers in northwestern U.S.A. that prevent southern extensions of the ranges of boreal species.
The male genital tract of Diptera is surveyed, based on whole dissections, with emphasis on nematocerous infraorders and especially the Culicomorpha. The genital tracts of all available nematocerous families are described and illustrated, including, for the first time, those of the families Thaumaleidae, Dixidae, Hesperinidae, and Pachyneuridae, and, as an outgroup, the mecopteran family Nannochoristidae (Mecoptera). On the basis of outgroup comparison, the Diptera + Mecoptera + Siphonaptera share the synapomorphies of accessory gland continuous with the vas deferens and U-shaped vasa deferentia. Further synapomorphies of the male genital tract provide evidence of the monophyly of the Mecoptera + Siphonaptera, Boreidae + Siphonaptera, and the Blephariceridae + Psychodidae + Trichoceridae + Anisopodidae + Brachycera. The Bibionomorpha, Culicomorpha, and Ptychopteromorpha share the synapomorphy of paired, two-to three-chambered accessory gland complexes, with secondary losses in Corethrellidae + Culicidae. It is concluded that there is no convincing evidence for the monophyly of the Chironomoidea. A sperm pump with an ejaculatory apodeme and the absence of a spermatophore are considered to be part of the ground plan of Diptera, but, because of uncertainty as to the identity of the basal lineage within Diptera and the homology of the sperm pump outside of Diptera, the polarity of these features cannot be presently interpreted.
The pupae of species in each of the 45 genera of Ceratopogonidae known in this stage are diagnosed and described. A standard set of terms is provided, with a glossary, for all pertinent structures of the pupal stage within a context of other Culicomorpha. The variety of terms provided by previous authors are synonymized. Some pupal structures are directly related to developing adult structures and these are discussed. A key to the genera (and to the subgenera of Forcipomyia Meigen) allows for their identification. Pupation and pupal behaviour is summarized. A table of all previous descriptions of each stage of the immatures (egg, larva, pupa) is provided, showing that 13% of all validly named extant Ceratopogonidae are known as pupae. This study examined 45% of these species. All species known as fossil pupae are discussed. A phylogenetic analysis based primarily on pupal characters confirms the relationships between the subfamilies as well as the relationships between the genera in Leptoconopinae, Forcipomyiinae and Dasyheleinae. The question of the monophyly of the Culicoidini remains unresolved. Results confirm the paraphyly of the Ceratopogonini and, for the first time, the Sphaeromiini sensu lato, which is divided into Hebetulini (new tribe), Johannsenomyiini Crampton (new status) and Sphaeromiini sensu novum. Sphaeromiini sensu novum includes Sphaeromias Curtis, Leehelea Debenham, Homohelea Kieffer and Xenohelea Kieffer and forms the sister group of the Palpomyiini. Other genera in Sphaeromiini sensu lato not known as pupae are discussed. The genus Mallochohelea Wirth is shown to be polyphyletic and one group of species is therefore recognized as members of the new genus Anebomyia (type species = Mallochohelea atripes Wirth). A number of species previously placed in Stilobezzia Kieffer are shown to belong to Schizonyxhelea Clastrier. Study of the type species of the monotypic genus Nemoromyia Liu and Yu showed it to be a member of the Palpomyia distincta species group and the generic name is therefore a new junior synonym of Palpomyia. Examination of pupae showed that the following species were misplaced and are now in other genera as follows: Schizonyxhelea brevicostalis (Das Gupta & Wirth), 1968: 28 (Stilobezzia) new combination, Schizonyxhelea bulla (Thomsen), 1935: 289 (Stilobezzia) new combination, Schizonyxhelea caribe (Lane & Forattini), 1958: 208 (Stilobezzia) new combination, Schizonyxhelea diminuta (Lane & Forattini), 1958: 209 (Stilobezzia) new combination, Schizonyxhelea obscura (Lane & Forattini), 1958: 216 (Stilobezzia) new combination, Schizonyxhelea panamensis (Lane & Forattini), 1958: 218 (Stilobezzia) new combination, Schizonyxhelea thomsenae (Wirth), 1953: 83 (Stilobezzia) new combination, Schizonyxhelea scutata (Lane & Forattini), 1961: 92 (Stilobezzia) new combination, Pellucidomyia geari (de Meillon & Wirth), 1981: 547 (Macropeza) new combination, Dibezzia prominens (Johannsen) 1932: 435 (Johannsenomyia) new combination, Nilobezzia theileri (de Meillon & Wirth, 1981:552) (Sphaeromias) ...
Study of all flies (Diptera) collected for one year from a four-hectare (150 x 266 meter) patch of cloud forest at 1,600 meters above sea level at Zurquí de Moravia, San José Province, Costa Rica (hereafter referred to as Zurquí), revealed an astounding 4,332 species. This amounts to more than half the number of named species of flies for all of Central America. Specimens were collected with two Malaise traps running continuously and with a wide array of supplementary collecting methods for three days of each month. All morphospecies from all 73 families recorded were fully curated by technicians before submission to an international team of 59 taxonomic experts for identification.Overall, a Malaise trap on the forest edge captured 1,988 species or 51% of all collected dipteran taxa (other than of Phoridae, subsampled only from this and one other Malaise trap). A Malaise trap in the forest sampled 906 species. Of other sampling methods, the combination of four other Malaise traps and an intercept trap, aerial/hand collecting, 10 emergence traps, and four CDC light traps added the greatest number of species to our inventory. This complement of sampling methods was an effective combination for retrieving substantial numbers of species of Diptera. Comparison of select sampling methods (considering 3,487 species of non-phorid Diptera) provided further details regarding how many species were sampled by various methods.Comparison of species numbers from each of two permanent Malaise traps from Zurquí with those of single Malaise traps at each of Tapantí and Las Alturas, 40 and 180 km distant from Zurquí respectively, suggested significant species turnover. Comparison of the greater number of species collected in all traps from Zurquí did not markedly change the degree of similarity between the three sites, although the actual number of species shared did increase.Comparisons of the total number of named and unnamed species of Diptera from four hectares at Zurquí is equivalent to 51% of all flies named from Central America, greater than all the named fly fauna of Colombia, equivalent to 14% of named Neotropical species and equal to about 2.7% of all named Diptera worldwide. Clearly the number of species of Diptera in tropical regions has been severely underestimated and the actual number may surpass the number of species of Coleoptera.Various published extrapolations from limited data to estimate total numbers of species of larger taxonomic categories (e.g., Hexapoda, Arthropoda, Eukaryota, etc.) are highly questionable, and certainly will remain uncertain until we have more exhaustive surveys of all and diverse taxa (like Diptera) from multiple tropical sites.Morphological characterization of species in inventories provides identifications placed in the context of taxonomy, phylogeny, form, and ecology. DNA barcoding species is a valuable tool to estimate species numbers but used alone fails to provide a broader context for the species identified.
Estimations of tropical insect diversity generally suffer from lack of known groups or faunas against which extrapolations can be made, and have seriously underestimated the diversity of some taxa. Here we report the intensive inventory of a four-hectare tropical cloud forest in Costa Rica for one year, which yielded 4332 species of Diptera, providing the first verifiable basis for diversity of a major group of insects at a single site in the tropics. In total 73 families were present, all of which were studied to the species level, providing potentially complete coverage of all families of the order likely to be present at the site. Even so, extrapolations based on our data indicate that with further sampling, the actual total for the site could be closer to 8000 species. Efforts to completely sample a site, although resource-intensive and time-consuming, are needed to better ground estimations of world biodiversity based on limited sampling.
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