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- Title
Phylogeny and classification of Aedini (Diptera: Culicidae), based on morphological characters of all life stages.
- Authors
Reinert, John F.; Harbach, Ralph E.; Kitching, Ian J.
- Abstract
Higher-level relationships within Aedini, the largest tribe of Culicidae, are explored using morphological characters of eggs, fourth-instar larvae, pupae, and adult females and males. In total, 172 characters were examined for 119 exemplar species representing the existing 12 genera and 56 subgenera recognized within the tribe. The data for immature and adult stages were analysed separately and in combination using equal (EW) and implied weighting (IW). Since the classification of Aedini is based mainly on adult morphology, we first tested whether adult data alone would support the existing classification. Overall, the results of these analyses did not reflect the generic classification of the tribe. The tribe as a whole was portrayed as a polyphyletic assemblage ofAedesandOchlerotatuswithin which eight (EW) or seven (IW) other genera were embedded. Strict consensus trees (SCTs) derived from analyses of the immature stages data were almost completely unresolved. Combining the adult and immature stages data resulted in fewer most parsimonious cladograms (MPCs) and a more resolved SCT than was found when either of the two data subsets was analysed separately. However, the recovered relationships were still unsatisfactory. Except for the additional recovery ofArmigeresas a monophyletic genus, the groups recovered in the EW analysis of the combined data were those found in the EW analysis of adult data. The IW analysis of the total data yielded eight MPCs consisting of three sets of two mutually exclusive topologies that occurred in all possible combinations. We carefully studied the different hypotheses of character transformation responsible for each of the alternative patterns of relationship but were unable to select one of the eight MPCs as a preferred cladogram. Overall, the relationships within the SCT of the eight MPCs were a significant improvement over those found by equal weighting. Aedini and all existing genera exceptOchlerotatusandAedeswere recovered as monophyletic.Ochlerotatusformed a polyphyletic assemblage basal toAedes. This group includedHaemagogusandPsorophora, and alsoOpifexin a sister-group relationship withOc. (Not.)chathamicus.Aedeswas polyphyletic relative to seven other genera,Armigeres,Ayurakitia,Eretmapodites,Heizmannia,Udaya,VerrallinaandZeugnomyia. With the exception ofAe. (Aedimorphus),Oc. (Finlaya),Oc. (Ochlerotatus) andOc. (Protomacleaya), all subgenera with two or more species included in the analysis were recovered as monophyletic. Rather than leave the generic classification of Aedini in its current chaotic state, we decided a reasonable and conservative compromise classification would be to recognize as genera those groups that are‘weighting independent’, i.e. those that are common to the results of both the EW and IW analyses of the total data. The SCT of these combined analyses resulted in a topology of 29 clades, each comprising between two and nine taxa, and 30 taxa (includingMansonia) in an unresolved basal polytomy. In addition to ten genera (Armigeres,Ayurakitia,Eretmapodites,Haemagogus,Heizmannia,Opifex, Psorophora, Udaya,VerrallinaandZeugnomyia), generic status is proposed for the following:(i) 32 existing subgenera ofAedesandOchlerotatus, including nine monobasic subgenera within the basal polytomy, i.e.Ae. (Belkinius),Ae. (Fredwardsius),Ae. (Indusius),Ae. (Isoaedes),Ae. (Leptosomatomyia),Oc. (Abraedes),Oc. (Aztecaedes),Oc. (Gymnometopa) andOc. (Kompia); (ii) three small subgenera within the basal polytomy that are undoubtedly monophyletic, i.e.Ae. (Huaedes),Ae. (Skusea) andOc. (Levua), and (iii) another 20 subgenera that fall within the resolved part of the SCT, i.e.Ae. (Aedes),Ae. (Alanstonea),Ae. (Albuginosus),Ae. (Bothaella),Ae. (Christophersiomyia),Ae. (Diceromyia),Ae. (Edwardsaedes),Ae. (Lorrainea),Ae. (Neomelaniconion),Ae. (Paraedes),Ae. (Pseudarmigeres),Ae. (Scutomyia),Ae. (Stegomyia),Oc. (Geoskusea),Oc. (Halaedes),Oc. (Howardina),Oc. (Kenknightia),Oc. (Mucidus),Oc. (Rhinoskusea) andOc. (Zavortinkius). A clade consisting ofOc. (Fin.)kochi,Oc. (Fin.)poiciliusand relatives is raised to generic rank asFinlaya, andDownsiomyiaVargas is reinstated from synonymy withFinlayaas the generic name for the clade comprisingOc. (Fin.)leonis,Oc. (Fin.)niveusand their relatives. Three other species ofFinlaya− Oc. (Fin.)chrysolineatus,Oc. (Fin.)geniculatusandOc. (Fin.)macfarlanei− fall within the basal polytomy and are treated asOc. (Finlaya)incertae sedis.Ochlerotatus(Ochlerotatus) is divided into three lineages, two of which,Oc. (Och.)atropalpusandOc. (Och.)muelleri, are part of the basal polytomy. The remaining seven taxa ofOc. (Ochlerotatus) analysed, including the type species, form a reasonably well-supported group that is regarded asOchlerotatus s.s. Ochlerotatus(Rusticoidus) is retained as a subgenus withinOchlerotatus s.s. Ochlerotatus(Nothoskusea) is recognized as a subgenus ofOpifexbased on two unique features that support their sister-group relationship. A new genus,Tanakaiusgen. nov., is proposed forOc. (Fin.)togoiand the related speciesOc. (Fin.)savoryi. The taxonomic status and generic placement of all currently valid species of Aedini are listed in an appendix. © 2004 The Linnean Society of London,Zoological Journal of the Linnean Society, 2004,142, 289−368.
- Subjects
PHYLOGENY; MOSQUITOES; BIOLOGICAL evolution; DEVELOPMENTAL biology; CLADISTIC analysis; ANIMAL morphology
- Publication
Zoological Journal of the Linnean Society, 2004, Vol 142, Issue 3, p289
- ISSN
0024-4082
- Publication type
Article
- DOI
10.1111/j.1096-3642.2004.00144.x