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Diptera: Culicidae: Aedini) Based on Morphological Data from All Life Stages

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Diptera: Culicidae: Aedini) Based on Morphological Data from All Life Stages Al 14I Zoological Journal of the Linnean Society, 2006, 148, 1-101. With 2 figures Phylogeny and classification of Finlaya and allied taxa (Diptera: Culicidae: Aedini) based on morphological data from all life stages JOHN F. REINERTI, RALPH E. HARBACH2 and IAN J. IUTCHING2 Center for Medical, Agricultural and Veterinary Entomology, United States Department of Agriculture, Agricultural Research Service, 1600/1700 SW 23rd Drive, Gainesville, FL 32608-1067, USA 2Department of Entomology, The Natural History Museum, Cromwell Road, London SW7 5BD, UK Received July 2005; accepted for publication March 2006 The phylogenetic relationships and generic assignments ofFinlaya and related taxa of uncertain taxonomic position in the classification of Aedini proposed by Reinert et al. (2004) are explored using 232 characters from eggs, fourth- instar larvae, pupae, adults and immature habitat coded for 116 exemplar species. The ingroup comprises 74 species, including 41 currently classified as Finlaya, 25 formerly placed in this group and eight related species. The outgroup includes four non-aedine species and 38 aedine species representing all generic-level taxa of the polyphyletic Ochierotatus and major clades within the polyphyletic Aedes in figure 4 of Reinert et al. (2004). Data are analysed in a total-evidence approach using implied weighting. The analysis produced four most parsimonious cladograms and ambiguity among the four is relatively minor concerning just four parts of the cladograms, two of which are linked. The strict consensus tree corroborates the monophyly of the 13 genera proposed by Reinert et al. (2004) that are included in the analysis. Overall, the results show remarkable congruence with those of Reinert et al. (2004) despite differences in the taxa included and morphological characters examined in the two studies. Seventeen genera, of which 11 are new, are proposed for monophyletic clades based on the strength of character support and application of the principle of equivalent rank. Bruceharrisonius stat. nov., Macleaya stat. nov., Molpemyia stat. nov. and Pseudoskusea stat. nov. are raised from subgeneric rank within Ochlerotatus; Hulecoeteomyia stat. nov. and Phagomyia stat. nov. are resurrected from synonymy with Finlaya; and Collessius gen. nov. (Ochlerotatus mac- farlanei Edwards), Dahliana gen. nov. (Culex geniculatus Olivier), Dobrotworskyius gen. nov. (Aedes tubbu- tiensis Dobrotworsky), Georgecraigius gen. nov. (Culex atropalpus Coquillett), Gilesius gen. nov. (Culex pulchriventer Giles), Himalaius gen. nov. (Finlaya gilli Barraud), Jarnellius gen. nov. (Culex varipalpus Coquil- lett), Jihlienius gen. nov. (Aedes chungi Lien), Patmarksia gen. nov. (Leucomyia australiensis variety papuensis Taylor), Rampamyia gen. nov. (Culex notoscriptus Skuse) and Vansoinerenis gen. nov. (Aedes pulchrithorax Edwards) are introduced for the type species indicated in parentheses, and their allies. Additionally, Hors fallius subgen. nov. (Culex fluviatilis Lutz) and Lewnielsenius subgen. nov. (Aedes muelleri Dyar) are introduced as subgenera of Georgecraigius and Jarnellius, respectively. As is usual with generic-level groups of Aedini, these genera are polythetic taxa that are diagnosed by unique combinations of characters. The analysis also shows that Oc. (Protomacleaya) is a polyphyletic assemblage of species, which is retained as a non-monophyletic taxon until the included species can be classified into monophyletic groups. Journal compilation © 2006 The Linnean Society of London, Zoological Journal of the Linnean Society, 2006, 148, 1-101. No claim to original US government works. ADDITIONAL KEYWORDS: cladistics - mosquitoes - systematics - taxonomy. INTRODUCTION ognized species, 32 subspecies, nine infrasubspecific forms/varieties and nine nomina dubia. Reinert, Har- Aedini (sensu Belkin, 1962) is the largest tribe in fam- bach & Kitching (2004) conducted a phylogenetic anal- ily Culicidae (Diptera). It includes 1235 currently rec- ysis of higher-level relationships within the tribe using 172 morphological characters from all life stages Corresponding author. E-mail: and 119 exemplar species representing the 12 genera [email protected] and 56 subgenera that were recognized at the time. All Journal compilation © 2006 The Linnean Society of London, Zoological Journal of the Linnean Society, 2006, 148, 1-101 No claim to original US government works. 2 J. F. REINERT ET AL. of the genera except Aedes and Ochierotatus were and subgroups are by no means sacrosanct and we recovered as monophyletic. (The authors of all aedine already know, from life history studies, that reassess- taxa are cited in Appendix 4.) In addition to 12 for- ment of the present placing of some species will be merly recognized genera, Reinert et cii. (2004) pro- needed. This applies particularly to rockpool species. posed generic status for 32 subgenera of Aedes and Belkin (1962) followed the classification of Knight & Ochierot at us, raised a dade consisting of Oc. (Finiaya) Marks (1952) but stated that Finiaya as understood at and its relatives (Kochi Group) to generic rank as present is a large heterogeneous assemblage of genus Faniaya, elevated Downsiomyia from synonymy species . and I believe that a natural classification with Finiaya as the generic name for a dade compris- of Finiaya cannot be developed until the immature ing Oc. (Fin.) niveus and its relatives (Niveus Group), stages are carefully studied. Tanaka, Mizusawa & and described a new genus, Tanakaius, to accommo- Saugstad (1979) recognized that previous attempts at date Oc. (Fin.) togoi and Oc. (Fin.) savoryi (Togoi classification of Finiaya did not reflect a natural clas- Group). Genus Aedes was restricted to the former sub- sification and proposed nine groups for species of Fin- genus Aedes and genus Ochierotatus was limited to a iaya occurring in the Japanese and Korean area based dade consisting of the type species, six related species primarily on characters of the male genitalia and lar- and members of subgenus Rusticoidus. These actions vae. Schick (1970) revised the Terrens Group and left species previously included in subgenera Aedimor- Arnell & Nielsen (1972) revised the Varipalpus Group phus and Cci ncraedes of Aedes and subgenera Bruce- based on characters of the adult and immature stages. harrisonius, Chaetocruiomyia, Finiaya, Macieaya, Zavortink (1972) removed all New World species from Moipemyia, Ochierotatu.s, Protomacieaya and Pseu- Finiaya and placed them in subgenera Aztecaedes, doskusea of Ochierotatus without generic assign- Gymnometopa, Protomacleaya and Ochierotatus of ments. The present study is the first step toward Aedes and subgenus Conopostegus of Haemagogus resolving the relationships and generic placements of based on features of the adults, pupae and larvae. these taxa. It treats species and species-groups of Reinert (1990, 1993, 1999b, 2003) transferred some uncertain taxonomic position (incertae sedis) that species to other subgenera, i.e. Kenknightia, Moipe- were previously included in subgenus Finiaya of myia, Zavortinkius and Bruceharrisonius. Reinert Ochierotatus, and species previously removed from (2000a) elevated Ochierotatus to generic rank and Finiaya by Zavortink (1972) and placed in subgenera transferred subgenus Finiaya to that genus. Reinert Protomacleaya and Ochierotatus of Ochierot at us. (2002e) separated subgenus Finiaya and related sub- Edwards (1932) provided the first comprehensive genera of Ochierotatus into Assemblages and Sub- classification of Finiaya Theobald (as a subgenus of assemblages based on features of the female genitalia. Aedes). He recognized eight groups within Finiaya He mentioned that Subgenus Finiaya, as currently (designated groups A—H) based on a few morphological configured, contains several diverse groups that are characters of adults. Barraud (1934) followed markedly different in the female genitalia; therefore, Edwards arrangement of genera and subgenera. the above definition of the subgenus is somewhat Knight (1948) recognized that Edwards Group F con- imprecise and indicates the need for a revision of the tained more or less unrelated forms and therefore entire taxon. However, this revision should include all divided it into eight divisions (denoted by Roman life stages as suggested by Belkin (1962). Finally, numerals). Knight & Marks (1952) revised and Reinert et al. (2004), as noted above, formally limited expanded the classification of Edwards and added genus Finiaya to the Kochi Group. numerous subgroup categories. They also based their Few molecular phylogenetic studies of aedine taxa classification almost exclusively on adult characters have been published. Wesson, Porter & Collins (1992) and indicated that a number of the groups were unnat- examined the relationships of six aedine species (four ural assemblages. Whereas Knight & Hull (1951), Aedes, one Haemagogus, one Psorophora) based on when examining the Philippine fauna, utilized the rDNA ITS2 sequence data. Besansky & Fahey (1997) group categories of Knight & Marks (1952) (they noted included three Aedes and one Haemagogus among 13 this paper was in press), subsequent authors, i.e. mosquito species in a study of relationships based on Bohart (1957), Marks & Hodgkin (1958), Macdonald & sequence data for the nuclear protein-coding white Traub (1960), Lee et cii. (1982) and Lu & Ji (1997), pro- gene. Kumar, Black & Rai (1998) investigated the vided some modifications to their classification, again relationships of 15 culicine
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