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New Classification for the Composite Genus Aedes

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New Classification for the Composite Genus Aedes Journal of the American Mosquito Contol Association, 16(3):175-188, 2000 Copyright O 2000 by the American Mosquito Control Association, Inc. NEW CLASSIFICATION FOR THE COMPOSITE GENUS AEDES (DIPTERA: CULICIDAE: AEDINI), ELEVATION OF SUBGENUS OCHLEROTATUS TO GENERIC RANK, RECLASSIFICATION OF THE OTHER SUBGENERA. AND NOTES ON CERTAIN SUBGENERA AND SPECIES JOHN F REINERT' Center for Medical, Agricultural, and Veterinary Entomology (CMAVE), United States Department of Agriculture, Agricultural Research Sewice, 1600/1700 SW 23rd Drive, Gainesville, FL 32608 ABSTRACT The composite gents Aedes is divided into 2 genera, Aedes and, Ochlerotatus, on the basis of consistent primary characters of the female and male genitalia. Ochlerotatus is separated into 2 sections. Ad- ditional supplemental features of the female and male genitalia, 4th-stage larvae, and pupae are provided for the separation of the genera and sections as weII as a discussion of exceptions and comparisotrr. ihis classifi- cation is based on a morphological examination of specimens of over 65% of the currenily recognized species and all subgenera previously included in Aedes and representative material of all subgeneri and lenera of tribe Aedini. Published literature was examined and evaluated. All currently recognized subgenera are issigned to the appropriate genus. The proposed new generic classification provides better defined genera and a m6re natural arrangement of included taxa. Armigerini is formally recognized as a synonyrn, in part, of Aedini. KEY WORDS Aedes, Ochlerotatus, Aedin| Armigerini, ruxonomy INTRODUCTION define and was often characteized by a combina- tion The original description of Aedes by Meigen of characters or the lack of characters as typ- ified "This (1818), written in German and also in Latin, was by the statement of Hopkins (1952): genus very brief and included a single species, Aedes ci- very large has larvae which are distinguished nereus Meigen. A translation to English, provided more by the absence of peculiar characters than by by the late George C. Steyskal, Gainesville, FL, of their presence." This has resulted in a generic level the description follows: antennae extending for- taxon that is poorly defined, generally without de- ward, filiform, l4-segmented, in male long plu- finitive features, and undoubtedly heterogeneous. mose, in female bristly; proboscis extending for- Recent actions (Reinert l999c,ZOOOb. 2000c) have ward, as long as dorsum ofthorax; palpi very short; partially remedied this by removing several groups wings scaly, lying down. Christophers (1960) from the genus (i.e., Ayurakitia Thurman and, Ver- discussed the history and rationale for not using the rallina Theobald were restored to generic rank, and diaeresis in the generic name Aedes: also, this is Sinoaedes Gong and Lu was placed in synonymy covered by Article 27 of the International Code of with subgenas Mattinglyia Lien of Heizmannia Zoolo gical Nomenclature. Ludlow). Investigators who have examined Aedes have The lst comprehensive treatment of Aedes was consistently recognized that it was heterogeneous. published by Edwards (1932). He included 24 sub- ... Typical comments concerning the genus are . genera in the genus, most of which had previously its internal classification is in great need of revi- been treated as genera. Edwards' definition of Ae- "A sion" (Belkin 1962), polymorphic genus; most des contained many characters that were variable characters extremely variable" (Tbnaka et al. 1979). and few that were constant. "A Subsequent authors and very large heterogeneous genus in which have followed his arrangement of the genus and subgenera, or species groups within major subgen- subgenera, but with some modification. era, are far more readily characteized than tlre ge- Edwards (1932, l94l) indicated that Aedes could, (Lee nus itself" et al. 1982). Mattingly (t97liin be divided into numerous subgenera primarily on his keys to the life stages of mosquito genera of the basis of the structure of the male genitalia and the world, noted numerous exceptions in genus Ae- the form of the male maxillary palpus. He divided des. Harbach and Kitching (1998) in their phylog- the subgenera into 2 main groups. The lst group eny and classification of Culicidae discussed the was defined primarily by the male genitalia having problematic nature of genus. the As a result of its claspettes present, arising from the basosternal area heterogeneous nature, Aedes has been difficult to of the gonocoxite and normally in the form of a fingedike process bearing a hooked filament api- rAlso collaborator, Walter Reed Biosvstematics Unit cally, and a phallosome that was undivided, scoop- (WRBU), National Museum of Natural ilistorv. Smith- shaped, and without teeth. The 2nd group possessed sonian Institution, Washington, DC 20560. male genitalia with structures corresponding to the 175 t76 JounNr- oF THE AMERTCINMosqurro CoNrnol AssocIATroN VoL. 16, No. 3 claspettes but in the form of a hairy basal lobe or sory parts published in Mosquito Systematics plaque, never as a fingerlike process with a hooked Newsletter and Mosquito Systematics between appendage, and a phallosome that was divided into 1970 and 1978 by these authors and the late Jean lateral plates that were variously toothed. Edwards L. Laffoon. "Some (1941) stated, however, that small and anomalous subgenera do not fit easily into either of MATERIALS AND METHODS these groups .. ." Belkin (1962) proposed a division of ttte genus During the last 30 plus years, I have had the on characters of the male genitalia (i.e., proctiger opportunity to examine the morphology of speci- with cercal setae present or absent and the aedeagus mens of all the 43 currently recognized subgenera simple or complex) and the 4th-stage larvae with of Aedes and the 1O other genera of Aedini (i.e., seta l2-I present or absent. His study was based Armigeres Theobald,2 Ayurakitia, Eretmapodites principally on material from the South Pacific area Theobald, Haemagogus Williston, Heizmannia, and included 12 subgenera of Aedes and 2 other Optfex Hutton, Psorophora Robineau-Desvoidy, genera of Aedini. He recognized that the genus Udaya Thurman, Verrallina, and Zeugnomyia must be studied on a worldwide basis before it Leicester). I have studied all life stages where avail- could be truly evaluated, understood, and possibly able but have devoted special attention to a study reorganized. of the female genitalia of the subgenera and genera Even though Aedes has not been evaluated on a of the tribe. I am convinced that the structures of worldwide basis since Edwards (1932), consider- the female genitalia offer some of the best charac- able work has been done on the included subgen- ters for partitioning generic, subgeneric, and group era. Much of this work benefited from the works level taxa. In several pending papers, I will treat of Belkin (1951, L952, 1953, 1962) in which the these structures in detail. The male genitalia also stability of the chaetotaxy of the 4th-stage larva have been extensively studied and offer outstanding and pupa was established and its taxonomic im- characters for grouping similar taxa of Aedini at all portance was demonstrated. In 1962, Belkin set a levels. Characters described for the male genitalia new standard for providing detailed descriptions are in the prerotation sense. Now that the 4th-stage and illustrations of the life stages of subgenera in- larvae of more species have been accurately de- cluded it Aedes. Following this format, numerous scribed and illustrated in detail it has become evi- revisionary studies of subgenera have been pub- dent that this stage presents a wealth of characters lished, such as Marks (1962,1964) (Chaetocruio- not only for distinguishing species but also for or- myia Theobald); Belkin (1968) (Nothoskusea gafizing and defining higher level taxa. The pupal Dumbleton); Knight (1968) (Finlaya Theobald, in and egg stages have not received thorough evalu- part); Huang (1968) (Huaedes Huang), (1972, ations, but as more species in various groups, sub- 1977, 1979, 1990, 1997) (Stegomyia Theobald); genera, and genera are described and illustrated, Berlin (1969) (Howardina Theobald); Zavortink character trends are emerging. Unfortunately, (1970) (Abraedes Zavortink), (1972) (Abraedes, worldwide, a considerable number of species in Ae- Aztecaedes Zavortink, Gymnometopa Coquillett, dini still have not been adequately described and Kompia Aitken, and Protomacleaya Theobald); illustrated in all stages. Schick (1970) (Protomacleaya, in part); Reinert Illustrations showing examples of the primary (1970, 1973b) (Diceromyia Theobald), (1973a) and supplemental characters used below to distin- (Aedimorphus Theobald), (1973c) (Bothaella guish the 2 genera and the 2 sections of Ochlero- Reinert), (1976a) (Edwardsaedes Belkin and In- tatus can be seen in Figs. 1-4. Parts A{ of Fig. du s i u s E,dw ards), ( 1976b) (Rhino sku se a Edwards), 1 are original. Other figures are electronically mod- (1979) (Isoaedes Reinert), (1981) (Paraedes Ed- ified and reconfigured from those in the following wards), (1982a) (Belkinius Reinert), (1985) (Scu- publications: Figs. lE, 2A,28, and 4C, 4D from tomyia Theobald), (1986) (Albuginosus Reinert), Zavortktk (1972); Figs. 2E-2I and 3A-3C, 3E, 3F (1990) (Kenkni ghtia Reinert), (1993) (Molpemy i a from Reinert (1973a); Figs. 3H, 3I from Reinert Theobald), (1999a) (Rusticoidus Shevchenko and Prudkina), ( 1999b) (Zavortinkius Reinert), (2000a) 'Li and Lu (1997:28O-281) removed Armigeres
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