New Classification for the Composite Genus Aedes

Home , Aedes, Aedes cinereus, Aedini, Eretmapodites, Fredwardsius, Ochlerotatus

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 classification 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 from (Fredwardsiu.s Reinert); Tyson (1970) (Mucidus Aedini and placed it in a new monogeneric tribe, Armi- Theobald); Arnell and Nielsen (1972) (Ochlero- gerini. However, I believe Annigeres should be retained tatus Lynclr Anibalzaga, in part); Arnell (1976) in Aedini because the new tribe is not equivalent in rank (Ochlerotatus, in part); Abercrombie (1977) relative to other tribes of Culicidae and the characters giv- (page (Christophersiomyia Barraud); and Tanaka et al. en by Li and Lu 281) to define Armigerini are similar to members of other genera of Aedini (e.g., male gen- (1979) (Aedes Meigen, in part; Finlaya, in part; italia with the gonostylus having multiple gonostylar part; part). Addi- Ochlerotatus, in Stegomyia, in claws in a row also is found in numerous species of sub- tional stability in the nomenclature of all life stag- gents Aedimorphus of Aedesi and the larval siphon with- es of Culicidae has resulted from the publication out a pecten also is found in several species of Eretma- Taxonornists' Glossary of Mosquito Anatomy by podites). Therefore, Armigerini is hereby formally Harbach and Knight (1980, 1982) and its precur- recognized as a synonym, in part, of Aedini. SeprerasBn2000 Npw CI-.qssrFrcerroNron Aross t77

(1976a); Figs. 1D, 2C,2D, and 44, 48 from Arnell ing a minute to small spicule) that are situated more (1976); Fig. 3G from Reinert (1981); Fig. 3J from or less medially (Fig. lC); male genitalia have the Reinert (1982a); and Fig. 3D from Reinert (1993). claspette developed as a basal mesal lobe bearing Figures are not drawn to the same scale. setae (Figs. 28, 2G, 2H) or otherwise developed, but not developed as a single, elongate, columnar TAXONOMIC ACTIONS stem with a terminal bladelike filament. and have the gonostylus variously developed (Figs. 2E and On the basis of my examination of the morphol- 3E-3J) (e.g., with distal portion expanded, or bi- ogy of over 65Vo of the curently recognized spe- furcated, or trifurcated, or with rows or patches of cies (Knight and Stone 19771' Y:night 1978; Ward short stout spicules, or with more than I gonostylar 1984, 1992) and all subgenera previously included claw, or without a gonostylar claw); and 4th-stage it Aedes, representative material of all subgenera larvae have seta l2-l absent (Fig. 3B), the and genera of Aedini, and the published literature, ventral brush (seta 4-X) is grid (some I hereby propose the division of the composite ge- borne on a groups also may have a few precratal (Fig. nus Aedes into 2 genera, Aedes Meigen and, Och- setae) 3C), and (Fig. (single lerotatus Lynch Arribalzaga, and the latter genus seta 4-M is normally branched 3A') in into 2 sections. This action necessitates the resto- some species). ration of Ochlerotatus to generic rank. The divi- Genus Ochlerotatus: Male genitalia have the sions are based on consistent primary features of claspette developed as a moderately long to long, the female and male genitalia, and supplemental narrow, columnar stem with a terminal flattened fil- features of the female and male genitalia, 4th-stage ament (Figs. 2A-2D) (except in subgenera Geosku- larvae, and one pupal character. No apparent con- sea, Howardina, Kenlcnightia, and Rhinoskusea), sistent differences in adult habitus were noted for and the gonostylus is moderately long to long, rel- the 2 genera. atively n,urow throughout its length but with the I do not lightly undertake the division of genus distal portion niurower and somewhat curved mes- Aedes becatse many of the species transferred to ally, and with a single gonostylar claw that is long, gents Ochlerotatus are vectors of disease patho- nrurow, uniformly thick tlroughout its length, and gens, have medical and veterinary importance, and attached at the apex of the gonostylus (Figs. 24, have long been known as Aedes. However, I believe 2C)'. the creation of more natural and better defined gen- Section.L' Fourth-stage larvae have seta l2-lpre- era are valid reasons for proposing this change and sent (Fig. 4A) (except in 6 species, see discussion outweigh the initial inconvenience of the generic section below), the ventral brush (seta 4-X) is at- change of the affected species. Fortunately, the tached to a grid (Fig. 4B) (some groups also may transfer of numerous species from Aedes (mascu- have a few precratal setae) or in the 2 small-sized line) to Ochlerotatus (masculine) does not result in subgenera Molpemyia (Fig. 3D) and, Mucidus it is changing the gender of the species names (Inter- attached to a very large and posteriorly projecting national Code of hological Nomenclature, Article boss, seta 4-M is branched (Fig. aA) (rarely single), 3lb). The proposed Z-letter abbreviation is Oc. for and seta 3-V (and also normally seta 3-Itr,fV) is at Ochlerotatus, and Ae. remains the same for Aedes. least 2 times the length (often much longer) of seta Three-letter abbreviations used for subgenera fol- 5-V (and also normally seta 5-III,fV) @ig. 4,A) (ex- low Reinert (197 5, 1982b, 1992). cept variable in subgenus Protomacleaya, some Finlaya, and a few other species); and pupal setae Primary characters defining the genera 1-6-VI normally have 3 or more of the setae (Fig. Genus Aedes: Female genitalia have the insula branched lD). tonguelike and without setae (Fig. 1C); male geni- Section 1L' Fourth-stage larvae do not have seta talia have the proctiger without setae (Fig. 2E) and 12-I (Fig. 4C), and the ventral brush (seta 4-X) is the phallosome has the aedeagus consisting of 2 attached to a small boss (Fig. 4D), seta 4-M is sin- lateral plates that usually bear teeth laterally and/or gle (Fig. 4C) (rarely branched), and seta 3-V (and apically (Figs. 28, 2E 2I,2J). also normally seta 3-m,IV) is shorter than, or nor- Genus Ochlerotatus: Female genitalia have the mally less than 1.5 times the length of, seta 5-V insula liplike and bearing well developed setae lat- (and also normally seta 5-III,IV) (Fig. aC), but very erally (Figs. 1A, 1B); male genitalia have the proc- rarely is seta 3-V (and seta 3-m,ry) near 2 times tiger bearing cercal setae that are tiny and all the the length of seta 5-V (and seta 5-I[,ry); and pupal same size (Figs. 2A, 2C), and the phallosome has setae 1-6-VI are single (Fig. 1E) (few species with the aedeagus simple and scooplike, troughlike, or 1-3 of the setae with 2, 3 branches). tubelike (Figs. 2A, 2C).

Key based on female genitalia Supplemental features supporting the partition of the genera Insula tonguelike, without setae . . . Aedes Genus Aedes: Female genitalia have the insula ''lli usually with a few tiny tuberculi (often each bear- I'll'Tl* lll ll-:i:i i'T,,"iii*i JounNlr- or rnE AIragnrcANMosQUITo CoNTRoL AssocIATIoN VoL. 16, No. 3

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Key based on male genitalia however, many appeared to have utility in distinguishing (e.g., Proctiger without cercal setae,raedeagus consisting other taxa subgenera and species groups). of 2 lateral plates usually bearing teeth laterally Other consistent distinguishing features for genera and./orapically...... Aedes separating these may be discovered when a detailed Proctiger pith tiny cercal setae; aedeagussimple examination is made of structures not fully and scooplike, troughlike, or tubelike evaluated during this study, such as mouthparts of ....Ochlerotatus 4th-stage larvae, chaetotaxy and structure of lst- tExcept for a few species(see discussionsec- stage larvae and pupae, and the structure of eggs. tion). Few molecular studies dealing with species in Ae- dini have been published (e.9., Wesson et al. 1992, Kumar et al. 1998). Unfortunately, these studies Key based on 4th-stage larvae treated only a few species and genera; however, l. Setal2-I absent ... . . 2 they generally appear to provide additional support Seta 12-I presentl.... Ochlerotatus,Sectionl for the above proposed separation ofgenera. Future 2. Setaeof ventral brush attachedto srid' molecular studies should evaluate and compare ...... Aedesmuch larger numbers of species, subgenera, and Setaeof ventral brush attachedto small boss . . genera of Aedini and should link their results with " morphological features. iE;;;ili;.; *:*l Exceptions to the characters for the genera and ;;;i";i"3:Y'::"XXl sections given above, of which I am aware, are discussed below. Also, other comments pertaining to Subgenera included in each genus the classiflcation of genera and subgenera of Aedini Genus Aedes: Aedes, Aedimorphus, Alanstonea are provided. Mattingly, Albuginosus, Belkinius, Bothaella, Can- In male genitalia of genus Aedes, the phallosome craedes Edwards, Christophersiomyia, Diceromyia, has the aedeagus divided into 2 lateral plates that Edwardsaedes, Fredwardsius, Huaedes, Indusius, normally bear several stout lateral and./or apical teeth Isoaedes, Izptosomatomyra Theobald, Lorrainea and the plates are usually not, or are only weakly, Belkin, Neomelaniconion Newstead, Paraedes, fused apically. Species of subgenus Christophersi- Pseudarmigeres Stone and Knight, Scutomyia, Sku- omyia appear to have 2lateral plates of the aedeagus sea Theobald, and Stegomyia. that have the distal portions strongly curved mesally Genus Ochlerotatus, Section I: Chaetocruio- and fused at the apex and are without teeth (except myia, Finlaya, Geoskusea Edwards, Halaedes Ae. gombokensis Mattingly, which has 2, 3 short Belkin, Kenknightia, Levua Stone andBohm, Ma- stout teeth laterally). Species of subgenus Bothaella c I eay a Ttteobald., M o lp emy i a, M uc i du s, N o tho sku - have the distal portion of the lateral plates of the sea, Ochlerotatus, Protomacleaya, Pseudoskusea aedeagus curved mesally and fused at the apex, and Theobald, Rhinoskusea, Rusticoidus, and Tavor- each plate bears 3-8 short or long teeth (teeth are tinkius. weakly developed in Ae. helenae Reinert). Species Genus Ochlerotatus, Section II: Abraedes, Azte- in subgenus Lorranea appear to have the aedeagus caedes, Gymnometopa, Howardina, and, Kompia. divided into 2 lateral plates in which the apices are lightly fused and slightly extended posteriorly. The aedeagus of some subgenera (e.9., Alanstonea, Di- DISCUSSION ceromyia, and Isoaedes) have numerous long, stout, During this study, only morphological characters curved teeth laterally and apically, but the apices of that consistently separated genera Aedes and, Och- the lateral plates appear to be strongly fused. The lerotatus were selected as primary characters. Char- aedeagus is unknown and the claspette is only partly acters listed as supplemental features are very use- known for the monotypic subgenus Indusius (see ful in partitioning the genera and sections but have Reinert 1976a). some exceptions. The primary characters combined The gonostylus of the male genitalia in genus with the supplemental features reliably distinguish Aedes normally is variously developed (e.g., distal the 2 genera and 2 sections of Ochlerot@trrr. Nu- portion is expanded, or is bifurcate, or is trifurcate, merous other characters, in all life stages, also were or has rows or patches of short stout spicules, or examined and evaluated but were found to be too has 2 or more gonostylar claws, or has the gonos- inconsistent to be of value at the generic level; tylar claw absent), but rarely is it developed as in

Fig. l. (A_{) Female genitalia showing structures; (D, E) pupal segment VI showing setae. (A, D) Ochlerotatus (Ochlerotatus) scapularis (Rondani); (B, E) Ochlerotatus (Aztecaedes) ramirezi (Vargas and Downs); (C) Aedes (Ae- dimorphus) domesticus (Theobald). Abbreviations used: Ce, cercus; H, hinge; I, insula; LVL, Iower vaginal lip; PGL, postgenital lobe; SE, spermathecal eminence; Tu, tuberculus; UVL, upper vaginal lip; VI, abdominal segment VI. Larger numbers in circles on D and E signify distinctive setae. r80 JounNar op r-gr AvrentcAN MosQUno CoNrRoL AssocnrloN Vol. 16, No. 3

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IN *7., \-1t cl i'E_ Pr Ae SerreN{ssn 2000 NEw CLASSTFICATIoN FoR AEDES 181 genus Ochlerotatus. However, a few species have kusea) postspiraculosus (Dobrotworsky), the gon- the gonostylus moderately long to long, relatively ostylus bears 2 or more stouter setae subapically narrow throughout its length, and with a single ter- that are somewhat like the terminal gonostylar claw minal gonostylar claw (e.g., Ae. (Aedimorphus) na- but thinner. The gonostylar claw is sinuous in Oc. tronius Edwards, Ae. (Diceromyia) kanarensis E,d- (Rusticoidus) rusticus (Rossi). wards, Ae. (Diceromyia) micropterus (Giles), Ae. In Aedes,2 species, Ae. (Diceromyia) kanarensis (Diceromyia) reginae Edwards, Ae. (Isoaedes) ca- and Ae. (Stegonryia) calceatus Edwards, have a few vatic us Reinert, Ae. (Neome lanic onion) berg erardi very long, stout setae on the proctiger of the male Pajot and Geoffroy, Ae. (Neomelaniconion) mono- genitalia, but because of their unusual development, trichus Edwards, and species of subgenera Cftn's- I consider these to be a departure from the normal. tophersiomyia and Huaedes) except that the gon- Also, a few species of Edwards' (1932) Group C ostylar claw is usually differently developed. In of subgenus Aedimorphus (e.9., Ae. argenteopunc- subgenus Stegomyia the gonostylus superficially re- tatus (Tlteobald), Ae. minutus (Theobald), and Ae. sembles those of genus Ochlerotatus, but the distal punctothoracis (Theobald)) have a few tiny cercal portion is somewhat expanded and the gonostylar setae on the proctiger; their presence appears to be claw is attached subapically (except the gonostylar a reversal. claw appears to be attached apically irt Ae. noval- Within Aedini, the claspettes and the basal mesal bopictus Barraud [see Huang 1972]). Subgenera lobes are homologous structures that can be iden- Aedes, Indusius, Neomelaniconion, and Skusea tified as such by their connection basomesally to have the gonostylus attached subapically to the the spiculate, more or less narrow, somewhat gonocoxite and the gonocoxite has the sternal area troughlike, aedeagal guide (see Reinert [1999b] as developed into an apical projection. The gonostylar adapted from Wood [1991]; :interbasal lobe, in claw is absent in Ae. (Stegomyia) desmotes (Giles) part, of Freeborn ll924l;:median lobe of Schick and subgenera Aedes, B elkinius, Edwardsaede s, In- :interbasal fold of Tbnaka et al. dusius, and Paraedes. The latter 2 subgenera have [1970]: Il979l) that is situated ventral to the phallosome. Freeborn short spicules forming filelike ridges on the distal (1924) should be examined for a comparative ex- portion of the gonostylus. The gonostylus of Ae. amination of this structure within Culicidae. This (Fredwardsius) vittatus (Bigot) has the distal por- structure has apparently been nearly lost in species tion greatly expanded and bears numerous, rela- of subgenus Geoskusea because it appears to be tively long spicules in rows on one side, but this reduced to the spiculate aedeagal guide and a short species also bears a narrow, strongly curved, gon- thin strip extending laterally onto the ostylar claw. The gonostylus is bifurcate in all or basosternal portion gonocoxite some species of subgenera Aedes, Aedimorphus, of the mesal surface of the and D ic e ro my ia, Edw ar ds ae de s, P arae de s, and Skus ea. bears 1 or a few short, thin, inconspicuous setae. Ttvo or more gonostylar claws are present in many Species of subgenus Howardina (see Berlin 1969) species of subgenus Aedimorphus. Species of sub- have the claspette developed as a spiculate, some- genus Pseudarmigeres usually have, in addition to what elongate lobe, with the basal portion usually the gonostylar claw, several stout spicules on the somewhat expanded, and with a single, apical, mesal margin that resemble gonostylar claws. Spe- stout, setalike filament (2 apical setae in Oc. au- cies of subgenus Skusea possess a comblike row of reolineatus (Berlin) and Oc. ioliota (Dyar and, closely set, stout, blunt-tipped spicules. Knab), but I seta is shorter and slightly thinner). The gonostylus of the male genitalia of Oc. bry- Also, according to Berlin (1969), the claspette is gooi (Brunhes) (subgenus Zavortinkius) has the completely absent in the male genitalia of tl;Le 2 middle portion broadly expanded. In subgenus Ze- species in the Whitmorei Group of subgenus Flo- vaa, tllLe gonostylus bears a pair of short, stout, wardina; however, this should be reevaluated to see curved, and pointed gonostylar claws subapically. if this structure has been reduced to only an aedea- The gonostylar claw of Oc. (Geoskusea) leabaenen- gal guide. The development of the short claspette sis (Brug) arrd Oc. (Geoskusea) baisasi (Knight and bearing a stout setalike filament in subgenus Flo- Hull) is relatively short and has a somewhat flared wardina is somewhat similar to that of species in apex; the apex in the latter species also has a few subgenus Kenknightia of Ochlerotalus (see Reinert small notches. In Oc. (Finlaya) elsiae (Barraud), 1990). Species of subgenus Levua and Oc. (Och- Oc. (Finlaya) shortti (Barraud), and Oc. (Pseudos- lerotatus) antipodeus Edwards have a stout setalike

Fig. 2. Male genitalia showing structures. (A, C, E) Genitalia minus right gonocoxite and gonostylus, (E) minus tergum IX; (8, D, G, H) claspette (G and H enlarged); (R I, J) aedeagus and left paramere (enlarged). (A,B) Ochler- otatus (Aztecaedes) ramirezi; (C, D) Ochlerotatus (Ochlerotatus) scapularis; (E-G) Aedes (Aedimorphus) mediolineatus (Theobald); (H, l) Aedes (Aedimorphus) stenoetrus (Theobald); (J) Aedes (lsoaedes) cavaticus. Abbreviations used: Ae, aedeagus; Cf; claspette filament; Cl, claspette; CSe, cercal seta; CSt, claspette stem; Gc, gonocoxite; GC, gonostylar claw; Gs, gonostylus; Par, paramere; Pr, proctiger. l JounNlr- oF THE AMERTCANMoseulTo CoNTRoL AssocnrroN VoL. 16, No. 3

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fllament attached apically to a moderately long, co- lobe of the claspette. Additionally, considerable lumnar, claspette stem. work is needed on the internal classification of spe- In Ochlerotatu,s, male genitalia of subgenus Rfti- cies included in subgenus Ochlerotatus because, as noskusea are unusual in that the claspette does not currently known worldwide, it is very heterogenous have a filament and is bifurcate in 2 species, and and poorly defined. the gonostylus is expanded distally or near mid- Very short maxillary palpi of males are found in length and bears a subapical, longitudinally striated, Aedes in subgenera Aedes, Belkinius, Bothaella, short, gonostylar claw. The claspette of Oc. (No- Cancraedes, Christophe rsiomyia, Huaedes, Lepto- thoskusea) chathamicus (Dumbleton) has the fila- somatomyia, Paraedes, and most species of Geo- ment absent and is developed as a bluntly rounded, skusea, but they are also found in Ochlerotatus in lobelike process that is densely covered with short, subgenera Nothoskusea and Rhinoskusea. Else- thin setae. In subgenus Halaedes, the claspette ap- where in Aedini, short maxillary palpi of males are pears to be developed as an oblong plaque bearing found in genera Heizmannia, Verrallina, Zeugno- a number of curved lanceolate setae. is connected myia, and a few species of Haemagogus. with its mate by a spiculate aedeagal guide, and the Subgenus Gymnometopa shares several features dorsal lateral portion is fused with the basomesal with genus Haemagogus. Zavortink (1972) alluded portion of the tergal surface of the gonocoxite. In to a remote relationship among subgenera subgenus Pseudoskusea, the claspette is much re- Ab raede s, Azte c aede s, Gymnometopa, Howardirm, duced (similar to subgenus Geoskusea), is restricted and Kompia. This relationship is supported here as to a spiculate aedeagal guide and a short thin strip demonstrated by the characterization of Section II that extends laterally onto the basosternal portion of Ochlerotatus. of the mesal surface of the gonocoxite, and bears Fourth-stage larvae of subgenera Abraedes, Az- 2 or a few short, thin, inconspicuous setae. The tecaedes, Kompia, and Oc. muelleri (Dyar) share claspette of subgenus Rusticoidus and some species the unusual feature of the posterior pair of seta in of subgenus Ochlerotatus has the stem moderately the ventral brush (seta 4-X) being short, which ap- to greatly swollen subapically or at about midlength pears to be unique for Aedini. (e.9., Oc. aenigmaticus (Cerqueira and Costa) and The absence of seta l2-I in the 4th-stage larvae Oc. atlanticus (Dyar and Knab)), other species pos- is unusual for genus Ochlerotatus. However, l2-I sess a short, thumblike process on the stem suba- is absent in 4th-stage larvae in subgenus Howar- pically or at about midlength (e.g-, Oc. aurifer (Co- dina, tlne monotypic subgenera Abraedes, Azte- quillett), Oc. clelandi (Taylor), Oc. diantaeus caedes, Gymnometopa, and Kompia (all in Section (Howard, Dyar, and Knab), Oc. imperfeclas (Dob- II), 6 species of subgenus Ochlerotatus (i.e., Oc. rotworsky), Oc. intrudens (Dyar), Oc. nigrithorax impiger daisetsuzanus (Tanaka, Mizusawa, and (Marquart), Oc. pullatus (Coquillett), Oc. sagax Saugstad) [see Tanaka et al. 19791, Oc. laguna (Ar- (Skuse), Oc. sapiens (Marks), Oc. thibaulti (Dyar nell and Nielsen). Oc. monticola (Belkin and and Knab), and Oc. vittiger (Skuse), whereas other McDonald), Oc. muelleri, and Oc. varipalpus (Co- species appear to have an additional basolateral, quillett)), and Oc. (Chaetocruiomyia) tulliae (Tay- caudally directed, long process bearing setae in ad- lor) (see Marks 1962). dition to the columnar stem and filament (e.9., Oc. Setae of the ventral brush (seta 4-X) are attached aenigmaticus, Oc. atlanticus, Oc. pertina.r (Grab- to a grid in Section I of Ochlerotarur, which is dif- ham), Oc. serratus (Theobald), and Oc. tormentor ferent from Section II but similar to Aedes. Tlae (Dyar and Knab)). The claspette filament is rela- ventral brush is attached to a weakly developed grid tively short and transversely annulated in species of in 4th-stage larvae of subgenera Christophersi- subgenus Rusticoidus. Detailed comparative studies omyia and Maclcaya and some species of subgen- on the development of the claspette and its rela- era Finlaya and Stegomyia, whereas in 6 other spe- tionship with the basomesal structures (lobes, setal cies, Ae. (Alanstonea) brevitibia (Edwards), Ae. groups, specialized setae, etc.) of the gonocoxite (Leptosomatomyia) aurimargo Edwards, Ae. are needed. For example, I have observed a narrow (Stegomyia) aobae Belkin, Ae. (Stegomyia) futunae sclerotized attachment between these structures in Belkin, Ae. (Stegomyia) robinsoni Belkin, and Ae. a number of species, and in others it is difficult to (Stegomyia) tulagiensis Edwards, the ventral brush determine if the structure is a basomesal lobe of the is attached to a small boss. In the 4th-stage larva tergal surface of the gonocoxite or a dorsolateral of Oc. (Abraedes) papago (Zavortink), the boss at (-

Fig. 3. Fourth-stage larva showing structures. (A) Thorax; (B) abdominal segment I; (C, D) abdominal segments Vm, X and appendages, and segment VII in C; (E-J) gonostylus of male genitalia. (A-C, E) Aedes (Aedimorphus) vexans; (D) Ochlerotatus (Molpemyia) pecuniosus (Edwards); (E) Aedes (Aedimorphus) vexans; (F) Aedes (Aedimor- phus) pampangensis (Ludlow); (G) Aedes (Paraedes) bonneae Mattingly; (H) Aedes (Indusius) pulverulentus; (l) Aedes (Edwardsaedes) imprimens (Walker); (J) Aedes (Belkinius) aurotaeniatus. Abbreviations used: B, boss; G, grid; I, VII, VIII, and X, abdominal segments I, VII, VIII, and X with appendages; M, mesothorax; R prothorax; S, siphon; t metathorax. Larger numbers in circles signify distinctive setae. 184 JounNnLoF THEAMERrcer Moseurro CoNrnol AssocrenoN

irY t" l 1l

.,;' SppmNrsen2000 Nrw ClassmrcnnoN ron Arors 185 the base of the ventral brush of segment X is weak- monotypic subgenus Kompia in which their ab- ly developed. The ventral brush is attached to a senceI consider to be a reversal),prespiracular se- small but heavily pigmented boss in genera Eret- tae are absent,upper calypter of the wing has setae mapodites and Haemagogus. on the margin, mesopostnotumis without scalesor Larval seta 3-V is only slightly longer than seta setae (except in Oc. (Rusticoidus) lepidonotus (Ed,- 5-V and seta S-trI,fV is slightly longer than seta 3- wards) [see Reinert l999al and 4 speciesof sub- [I,IV in Oc. (Macleaya) tremulus (Theobald). genus Zavortinkius lsee Reinert 1999b1), dorsal Members of the Sexlineatus Section and 2 spe- margin of the mesomeron is above the base of the cies (Ioliota Group) of the Walkeri Section of sub- hindcoxa (except rn Ae. desmofes),and the ungues genus Howardina haye seta 13-P developed in the of the male foretarsus are unequal in size (except 4th-stage larvae. This seta is also present in sub- ungues are equal in size in Ae. (Indusius) pulver- gents Bothaella and the 4 species of subgenus ulentus Edwards); female genitalia are without low- Stegornyia, Ae. hoguei Belkin, Ae. robinsoni, Ae. er vaginal sclerites; pupae have seta 1-I well de- tulagiensis, and Ae. upolensis Marks. veloped and with a number of branches, and the Fourth-stage larvae of subgenera Aedes and Rus- paddle has only I seta (i.e., seta l-Pa); and 4th- ticoidus, and a few species of subgenus Ochlero- stagelarvae have a well developedpecten on a nor- tatus (i.e., Oc. crinifer (Theobald) [see Arnell mally moderately long siphon and setae9,10-T are 1976), Oc. hexodontus (Dyar), Oc. hexodontus hok- well developed. kaidensis (Tanaka, Mizusawa, and Saugstad), and Aedes has a distribution in the Old World except Oc. punctor (Kirby) [see Tanaka et a]. 19791) are for Ae. (Aedes)cinereus, Ae. (Aedimorphus)vexans unusual for Aedini in the development of accessory (Meigen), Ae. (Stegomyia)aegypti, andAe. (Stego- setae on the siphon. myia) albopicras (Skuse)that are found in the New The 4th-stage larva of Ae. (Alanstonea) breviti- World. The 2 species of subgenusStegomyia are bia lras seta 0-X present, which is unusual for Ae- believed to have been introduced to this region. In dini. North America, Ae. cinereus exhibits noticeable Seta 13-CT is present on the pupal metanotum morphological variability (Wood et al. 1979, Bick- of a few species of Aedes, that is, Ae. (Belkinius) ley 1980) and is reported from much of the United aurotaeniatus Edwards, a few African species of Statesand Canada(see Darsie and Ward 1981).The subgenus Aedimorphus, and the following species primary range occupied by genus Aedes is the Af- of subgenus Stegomyia: Ae. futunae, Ae. rotwnae rotropical and Oriental regions, but a few species Belkin, and,Ae. upolensis (see Reinert 1980). Setae occur in the Palaearctic and Nearctic regions (see 13,14-CT are present in Ae. krombeizi Huang (see above).Section I of Ochlerotatzs is essentiallycos- Tewari and Hiriyan 1990). mopolitan in distribution. Section fl of Ochlerota- Normally, 3 or more of setae l-6-VI are tur occurs in the New World and has a distribution branched in pupae of genus Ochlerotatus Section I, that is primarily Neotropical and centered in the but most of these setae are single in Section II and American Mediterranean Region (sensu Belkin in some species of subgenera Finlaya, Macleaya, 1962) and as reported by Berlin (1969) and Zavor and Molpemyia, the Varipalpus Group, and Oc. ttnk (1972). However, Section II extends north to muelleri. the southern part of the United States (Oc. The pupal paddle has a fringe of hairlike spicules (Abraedes)papago and Oc. (Kompia) purpureipes in subgenera Belkinius, Lorrainea, most Stegomyia (Aitken) in southern Arizona; and Oc. (Howardina) (except, e.g., Ae. aegypti (Linnaeus) and the Den- bahamensis(Berlin) that was probably intoduced drophilus Group), afew Diceromyia (e.9., Ae. fran- into southern Florida [Pafume et al. 1988]) and ciscoi Mattingly, Ae. iyengari Edwards, Ae. peris- south to northern Argentina along the Andes. kelatus (Giles), and Ae. tiptoni @ebine) of Aedes, and subgenus Abraedes of Ochlerotarzs. A similar ACKNOWLEDGMENTS hairlike fringe of spicules is found in the pupal paddle of genera Artnigeres, Eretmapodites, Heizman- Grateful appreciation is expressed to Donald R. nia, Udaya, and Zeugnomyia. Barnard and Herbert Oberlander (CMAVE) for pro- In addition to the previously mentioned charac- viding facilities; to Ralph E. Harbach (The Natural ters that distinguish Aedes and Ochlerotatus, some History Museum, London, United Kingdom), characters, in various combinations, are useful in Thomas J. Zavortink (Departmentof Biology, Uni- separating them from other genera of Aedini, such versity of San Francisco, San Francisco, CA), Ri- as, adults have postspiracular setae (except the chard C. Wilkerson and M. A. M. Sallum (WRBU),

Fig. 4. Fourth-stage larva showing structures. (A, C) Thorax and abdominal segments I-VI; (B, D) abdominal segments VIII and X with appendages. (A, B) Ochlerotatus (Ochlerotatus) scapularis; (C,D) Ochlerotatus (Aztecaedes) ramirezi. Abbreviations used: B, boss; G, grid; I-VI, VIII, and X, abdominal segments I-VI, Vru, and X; M, mesothorax; R prothorax; 1l metathorax; S, siphon. Larger numbers in circles signify distinctive setae. 186 Jouruval oF THE AMERTcaN Mosgurro CoNrnol AssocrerroN VoL.16, No.3

Bruce A. Harrison (North Carolina Department of cation of the Culicidae (Diptera). Syst Entomol23:327- Environmental and Natural Resources, Winston-Sa- 370. lem, NC), and Richard E Darsie, Jr. (Florida Med- Harbach RE, Knight KL. 1980. Taxonomists' glossary of mosquito anatomy Marlton, ical Entomology Laboratory, Vero Beach, FL) for NJ: Plexus Publishing, Inc. Harbach RE, Knight KL. 1982. Corrections and additions reviewing the manuscript; to Young T, Sohn (illus- to Taxonomists' glossary of mosquito anntomy. Mosq trator assigned to organization preceding WRBU) Syst (1981) 13:20r-217. for preparing the original illustrations used in Figs. Hopkins GHF, 1952. Mosquitoes of the Ethiopian Region lA-lC; to Keira Dooley (CMAVE) for the elec- I.-Larval bionomics of mosquitoes and taxonomy of tronic modification and reconfiguration of the fig- culicine larvae 2nd edition. London, United Kingdom: ures; and to the editors of the Contributions of the British Museum (Natural History). 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nids of Canada Part 6. Hull, Quebec, Canada: Canadian Zavortink TJ. 19'72. Mosquito studies (Diptera, Culicidae) Government Publishing Centre. 390 pp. XXVIII. The new world species formerly placed in Ae- Zavortink TJ. 1970. Mosquito studies (Diptera, Culici- des (Finlaya). Contrib Am Entomol Inst (Ann Arbor) dae) XXII. A new subgenus and species of Aedes from 8(3):1-206. Arizona. Contrib Am Entomol Inst (Ann Arbor\ 7(l\: l-11.