Location: Mosquito and Fly Research Unit
Title: Phylogeny and classification of tribe Aedini (Diptera: Culicidae). Authors
|Harbach, Ralph - BMNH, LONDON, UK|
|Kitching, Iam - BMNH, LONDON, UK|
Submitted to: Zoological Journal of the Linnean Society
Publication Type: Peer Reviewed Journal
Publication Acceptance Date: January 1, 2009
Publication Date: December 1, 2009
Citation: Reinert, J.F., Harbach, R.E., Kitching, I.J. 2009. Phylogeny and classification of tribe Aedini (Diptera: Culicidae). Zoological Journal of the Linnean Society. 157:700-794. Interpretive Summary: Generic-level taxa included in the large, heterogenous, mosquito tribe Aedini were analyzed using cladistic procedures to determine their phylogenetic relationships. Numerous morphological characters of all life stages of the included species were evaluated. The study, the fourth in the evaluation of Aedini, provides for the first time a natural classification of the largest and most complex tribe of the family Culicidae. Unfortunately, because of insufficient specimens or published descriptions a few species could not be placed with certainty in a recognizable generic-level taxon. Since a number of species evaluated in the present study are important in the transmission of pathogens to humans and domesticated animals the above information will be of value in epidemiological and control programs. This work was completed at the Center for Medical, Agricultural and Veterinary Entomology of the United States Department of Agriculture. The article will be published in the Zoological Journal of the Linnean Society.
Technical Abstract: The phylogeny and classification of tribe Aedini are delineated based on a cladistic analysis of 336 characters from eggs, fourth-instar larvae, pupae, adult females and males, and immature stage habitat coded for 270 exemplar species, including an outgroup of four species from different non-aedine genera. Analyses of the data set under implied weights, implemented by TNT version 1.1, with values of the concavity constant K ranging from 7 to 13 each produced a single most parsimonious cladogram (MPC). The MPCs obtained with K values of 7–9 were identical, and that for K = 10 differed only in small changes in the relationships within one subclade. Because values of K < 7 and > 10 produced large changes in the relationships among the taxa, the stability of relationships exemplified by the MPC obtained from the K = 9 analysis is used to interpret the phylogeny and classification of Aedini. Clade support was assessed using parsimony jackknife and symmetric resampling. Overall, the results reinforce the patterns of relationships obtained previously (Reinert et al., 2004, 2006, 2008) despite differences in the taxa and characters included in the analyses. With two exceptions, all of the groups represented by two or more species were once again recovered as monophyletic taxa. Thus, the monophyly of the following genera and subgenera is corroborated: Aedes, Albuginosus, Armigeres (and its two subgenera), Ayurakitia, Bothaella, Bruceharrisonius, Christophersiomyia, Collessius (and its two subgenera), Dahliana, Danielsia, Dobrotworskyius, Downsiomyia, Edwardsaedes, Finlaya, Georgecraigius (and its two subgenera), Eretmapodites, Geoskusea, Haemagogus (and its two subgenera), Heizmannia (and subgenus Heizmannia), Hopkinsius (and its two subgenera), Howardina, Hulecoeteomyia, Jarnellius, Kenknightia, Lorrainea, Macleaya, Mucidus (and its two subgenera), Neomelaniconion, Ochlerotatus (subgenera Chrysoconops, Culicelsa, Gilesia, Pholeomyia, Protoculex, Rusticoidus and Pseudoskusea), Opifex, Paraedes, Patmarksia, Phagomyia, Pseudarmigeres, Rhinoskusea, Psorophora (and its three subgenera), Rampamyia, Scutomyia, Stegomyia, Tanakaius, Udaya, Vansomerenis, Verrallina (and subgenera Harbachius and Neomacleaya), Zavortinkius and Zeugnomyia. In addition, the monophyly of Tewarius, newly added to the data set, is confirmed. Heizmannia (Mattinglyia) and Verrallina (Verrallina) were found to be paraphyletic with respect to Heizmannia (Heizmannia) and Verrallina (Neomacleaya), respectively. Fifteen additional genera, three of which are new, and 12 additional subgenera, 11 of which are new, are proposed for monophyletic clades, and a few lineages represented by a single species, based on branch support, the number and nature of the characters that support the branches, and the principle of equivalent rank. Aedimorphus stat. nov., Cancraedes stat. nov., Cornetius stat. nov., Geoskusea stat. nov., Levua stat. nov., Lewnielsenius stat. nov., Rhinoskusea stat. nov. and Sallumia stat. nov., which were previously recognized as subgenera (Reinert et al., 2008) of various genera, are elevated to generic status. Catageiomyia stat. nov. and Polyleptiomyia stat. nov. are resurrected from synonymy with Aedimorphus, and Catatassomyia stat. nov. and Dendroskusea stat. nov. are resurrected from synonymy with Diceromyia. Bifidistylus gen. nov. (type species: Aedes lamborni Edwards) and Elpeytonius gen. nov. (type species: Ochlerotatus apicoannulatus Edwards) are described as new for species previously included in Aedes (Aedimorphus), and Petermattinglyius gen. nov. (type species: Aedes iyengari Edwards) and Pe. (Aglaonotus) subg. nov. (type species: Aedes whartoni Mattingly) are described as new for species previously included in Aedes (Diceromyia). Four additional subgenera are recognized for species of Ochlerotatus, including Oc. (Culicada) stat. nov. (type species: Culex canadensis Theobald), Oc. (Juppius) subg. nov. (type species: Grabhamia caballa Theobald), Oc. (Lepidokeneon) subg. nov. (type species: Aedes spilotus Marks) and Oc. (Woodius) subg. nov. (type species: Aedes intrudens Dyar), and seven are proposed for species of Stegomyia: St. (Actinothrix) subg. nov. (type species: Stegomyia edwardsi Barraud), St. (Bohartius) subg. nov. (type species: Aedes pandani Stone), St. (Heteraspidion) subg. nov. (type species: Stegomyia annandalei Theobald), St. (Huangmyia) subg. nov. (type species: Stegomyia mediopunctata Theobald), St. (Mukwaya) subg. nov. (type species: Stegomyia simpsoni Theobald), St. (Xyele) subg. nov. (type species: Stegomyia desmotes Giles) and St. (Zoromorphus) subg. nov. (type species: Aedes futunae Belkin). Due to the unavailability of specimens for study, many species of Stegomyia are without subgeneric placement. As is usual with generic-level groups of Aedini, the newly recognized genera and subgenera are polythetic taxa that are diagnosed by unique combinations of characters. The analysis corroborates the previous observation that ‘Oc. (Protomacleaya)’ is a polyphyletic assemblage of species.