OCCURRENCE of MESOPOSTNOTAL SETAE and SCALES in the FAMILY Culicidael

SEPTEMBER1990 131

OCCURRENCE OF MESOPOSTNOTAL SETAE AND SCALES IN THE FAMILY CULICIDAEl

BRUCE A. HARRISON~ AND WILLIAM E. BKKLEY~

ABSTRACT. A surveyof specimensand the literature wasconducted to determine the presence,distribution and possiblephylogenetic significance of mesopostnotalsetae and/or scaleson adultsthroughout the family Culicidae. The distributionin the family is tabulated with detailednotes for the speciesin eachtaxon having the character,and 4 species with the characterare illustrated. The presenceand frequencyof the characterin varioustaxa of the tribes Aedini, Culicini and Sabethini are discussedhighlighting differences in given taxa between the Eastern and Western Hemispheres. Phylogeneticconsiderations regarding the correlation of the character with the subtropics/tropics, phytotelmicimmature habitats,and the origin of the current distributionof the characterare discussed.

INTRODUCTION He alsorecognized 10 generawhose species had or usually had setaeon the Mpn. To many mosquito taxonomistsand identifiers in the The mosquitotaxon usuallyassociated with the pres- Holarctic Region the mesopostnotumof mosquitoesis ence of Mpn setae is the tribe Sabethini, but not all usuallybare and not usedin taxonomicdecisions. On the sabethinespecies possess them. Some New World work- other hand,workers in the subtropicsand tropicsmust be ershave cometo rely on Mpn setaeor scalesas a constant alert for the presenceor absenceof scalesand/or setaeon characterin sabethines,e.g., Galindo et al. (1951) made thisstructure, as they canbe highlysignificant for separat- the followingstatement in the descriptionsof 2 new Wyeo- ing variousmosquito taxa. This structurehas been called myia species:“Postnotum bare exceptfor the usualtuft of the metathorax,metanotum or the postnotumby numer- hairspresent in all sabethines.”Belkin (1962) statedthat ous authorsover the years. Harbach and Knight (1980) the Mpn in sabethinesis often keeled and with setae or designatedthis structure “mesopostnotum”(Mpn), fol- scales,but frequentlymay be bare and smooth. Similarly, lowing Patton and Evans (1929). We are usingthe term speciesin the subfamilyCulicinae, particularly the tribes “mesopostnotum” instead of “postnotum” as used by Aedini and Culicini are generallyconsidered to lack Mpn Owen (1977) and McAlpine (1981) to conform with the setae,l?owever, there are exceptionsas noted below. terminologyused for the postnotumof the metathoraxof Actually, an appreciablenumber of speciesare known culicids,i.e., metapostnotum(Owen 1977, Harbach and to deviatefrom what mightbe considereda norm, i.e., that Knight 1980). Theobald (19Ola) found that the presence setae or scales on the Mpn is a sabethine character. of setaeon the mesopostnotum(as metanotum) was valu- Furthermore,the referencesnoting many of thesedeviates able in separating genera, and Theobald (19Olb: 95) are obscureor not generally known. Consequently,we mentioned that setae (as chetae) on this structurewere decided to undertake a survey of the specieswith and very importantfor characterizingthe genusWyeomyia and without Mpn setae and/or scalesto: (1) assembleand that setae and scaleswere important for recognizingthe reviewthe literature on this character;(2) determinehow genus Trichoprosopon. In the same paper, Theobald widespreadthis characteris in the family Culicidae;and classifiedthe genera Sabethes and Eretmapodites in the (3) seeif the charactermay be usefulin understandingthe sectionhaving the mesopostnotum(as metanotum)“ nude”. phylogenyof the Culicidae. This may have been an oversighton his part, or the Mpn structureswere rubbed off of the specimenshe studied. MATERIALS AND METHODS Edwards (1932) also recognizedthe value of Mpn setae, andin keysor descriptionshe referred to 15 generawhose Much of thissurvey involved a searchthrough published specieslacked Mpn setae, or nearly alwayslacked them. speciesdescriptions, particularly in those genera where the Mpn characteris variable. Although we have not examined every speciesin taxa such as Wyeomyia, Aedes or 1Theviewsof the authorsdonotpurport to reflect thepositionof the Culex(Melanoconion), we tried to confirmthe presenceof Departmentof the Army or the Departmentof Defense (para 4-3), AR Mpn setaeand/or scaleson all of the speciesin the smaller 360-S. taxawhere the characterhas been listedas present. In this ‘Department of Entomology,Walter Reed Army Institute of Re- search,Washington, DC, 20307-5100.Address for reprints: Walter regard,we were fortunate to haveexamined many species Reed BiosystematicsUnit, MuseumSupport Center, Smithsonian Insti- of different taxa over the lastfew years,and to haveaccess tution, Washington,DC, 20560. to the mosquitocollection located at the NationalMuseum 3651640th Avenue, University Park, MD, 20782. 132 MOSQUITO SY$XJNATICS VOL. 22, No. 2 of Natural History, Museum Support Center, Smithsonian Mpn setae. Reinert (1974: 95) reported that the male Institution, Washington, DC. The classification and the holotype ofAe. (VerraZZina)viriZis (Leicester) has one long names of the taxa follow Knight and Stone (1977), Knight and one short setae on the Mpn. In addition we found one (1978), Ward (1984) and Gaffigan and Ward (1985). The male and one female of 13 specimens of viriZisfrom Sabah, numbers listed in the various tribes, genera and subgenera Malaysia, with a single short seta on the Mpn. The second are close estimates (usually underestimates) and not to be species, Marks sp. No. 171 of Lee et al. (1987: 253), is an construed as final numbers. We believe there are hun- undescribed member of the subgenus VerraZZinafrom the dreds of culicid species still to be discovered, particularly Australasian Region. We were unable to find other spe- among the sabethines and other groups utilizing natural cies ofAedes with this character. plant receptacles. In Table 1 the numbers appearing in (2) The genus Armigeres has only one species that has parentheses refer to numbered notes in the results section, Mpn setae and a small group of pale scales, i.e., fzavus and after each taxon one or more authorities are listed. In (Leicester), in the subgenusLeicestetia. Macdonald (1960) addition, one or both of us have examined specimens in all discussed this peculiarity and stated that the Mpn is bare of the genera and subgenera listed in this study, and for on all the other species in Leicestetia. We have examined most of the species mentioned. When we use the term 20 of the 46 species in the genus, including 13 of the 31 “bare” it means the absence of setae and scales. This strict species in the subgenusArmigeres, and found this charac- definition was not used by some earlier authors, e.g., Lane ter only onflavus. Thurman (1959) and Stone et al. (1959) (1953) used nude to mean the absence of setae, even when placed flavus in a separate subgenus, Leicesteriomyia, scales were present. using the Mpn character as a major portion of their justification. However, Macdonald (1960) synonymized RESULTS AND NOTES this subgenus under Leicestetia on the basis of his very detailed and comprehensive morphological and ecological Table 1 shows the distribution of the Mpn character in studies of this subgenus. Armigeresflavus has both setae the family Culicidae as determined by our survey. Of the and scales on the mesopostnotum. During this study we 3 subfamilies, Anophelinae, Culicinae and Toxorhynchiti- examined 12 specimens offlavus and noted a range of l-6 nae, only the culicine tribes, Aedini, Culicini and Sabethini setae (mode 3) and a range of O-3 scales (mode 0). include species with Mpn setae and/or scales. Tables 2-3 (3) Eretmapodites is a genus confined to the Afrotropi- provide the frequency of this character in the genera cal Region. According to Edwards (1941) this is the only included in these 3 tribes. To date, no reports have been taxon in Africa with species having Mpn setae. He noted found of Mpn setae and/or scales on members of the other the presence of a small tuft of bristles in chrysogaster 7 tribes of Culicinae, i.e., Aedeomyiini, Culisetini, Ficalbi- Graham, dracenae Edwards, forcipulatus Edwards, gra- ini, Hodgesiini, Mansoniini, Orthopodomyiini, and Ura- hami Edwards, inomatus Newstead, intermedius Edwards, notaeniini. Furthermore, we have not seen Mpn setae penicillatus Edwards, semisimplicipes Edwards, silvestris and/or scales on species in these tribes. Table 4 lists the Ingram and de Meillon, and subsimplicipes Edwards. On primary oviposition habitats of the taxa containing species the basis of Edwards’ statement we infer that the following with Mpn setae/scales. species also possessMpn setae: argynrus Edwards, Zeu- Nine genera are currently recognized in the tribe Aedini, copus Graham, melanopous Graham, oedipodeios Gra- and Mpn setae and/or scales have been found on only ham, and plioleucus Edwards. Additionally, Edwards certain species of the following 5 genera, Aedes (1941) noted that quinquevittatus Theobald, and tonsus (cosmopolitan), Armigeres (Oriental), Eretmapodites Edwards lack Mpn setae. (Afrotropical), Haemagogus (Neotropical) and Heizman- Our observations of a small number of specimens from nia (Oriental). All 3 genera in the tribe Culicini, i.e., Culex Cameroon, Sierra Leone, and Ivory Coast confirm the (cosmopolitan), Deinocerites (Neotropical and subtropi- presence of Mpn setae on chrysogaster,dracenae, grahami, cal) and Galindomyia (Neotropical), have representatives Zeucopus,oedipodeios, semisimplicipes, and silvestris. We with this character. Nine of the 12 genera in the Sabethini have seen specimens of quinquevittatuswith the Mpn bare. have species with this character: Johnbelkinia (Neotropi- Since 1941, parvipluma Edwards, productus Edwards Cal), Limatus (Neotropical), Phoniomyia (Neotropical), and wansoni Edwards have been elevated from subspecies Runchomyia (Neotropical), Sabethes (Neotropical), Shan- to species. The subspecies,brevis Edwards and conchobius noniana (Neotropical), Trichoprosopon (Neotropical), Edwards are still subspecies and subspecies stanZeyiEd- Tripteroides (Oriental) and Wyeomyia (Nearctic and wards has been synonymized under oedipodeios. We have Neotropical). The expression of this character in the not seen specimens of these taxa and do not know if they above genera is as follows (numbers coincide with num- have Mpn setae. Regarding forcipulatus, Edwards (1941: bers in Table 1). 234) noted that the Mpn setae were reduced to l-2 or (1) Aedes is the largest genus in the family, with over 940 absent. species. However, only 2 species are known to possess An additional 24 species of Eretmapodites have been SEYI-EMBER1990 133

Table 1. Distribution of mesopostnotalsetae/scales in the family Culicidae.

Mpn Setae and/or Scales Taxon Present Presentor Absent Basic absent references

Anophelinae 0 Belkin (1962) Culicinae +/o See below Aedeomyiini 0 Belkin (1962) Aedini +/o See below Aedes +/om* Reinert (1974) and Lee et al. (1987) Armigeres +/o(2) Macdonald (1960) Eretmapodites +/o(3) Edwards (1941) Haemagogus + /o(4) Zavortink (1972) and Arnell(1973) Heizmannia + IO(5) Mattingly (1970) and presentstudy opij&x Belkin (1%2) Psorophora Carpenter and LaCasse(1955) Udaya Mattingly (1958) Zeugnomyia Baisasand Feliciano (1953) Culicini +/o See below Cl&X +/o (6) Rozeboom and Komp (1948) and presentstudy Deinocetites +/o (7) Adames (1971) Galindomyia +/o (f-9 Stone and Barreto (1969) and present study Culisetini Belkin (1962) Ficalbiini Belkin (1962) Hodgesiini Belkin (1962) Mansoniini Belkin (1%2) Orthopodomyiini Belkin (1962) Sabethini + /o(9) See below John belkinia + (10) Zavortink (1979a) Limatus + (11) Edwards(1932) Malaya 0 Belkin (1962) Maorigoeldia 0 Belkin (1962) Phoniomyia + 02) Theobald (1903), Lane (1953) and Correa and Ramalho (1956) Runchomyia + (13) Zavortink (1979a) Sabethes + (14) Edwards(1932) Shannoniana + (15) Zavortink (1979a) Topomyia 0 Edwards(1932) and Thurman (1959) Trichoprosopon + (16) Lane (1953) andZavortink (1979a, 1981) Ttipteroides + /o(17) Mattingly (1981) Wyeomyia + (18) Lane (1953) and Belkin et al. (1970) Uranotaeniini 0 Belkin (1%2) Toxorhynchitinae 0 Belkin (1%2) *Refer to numberednotes in text. 3.34 MOSQUITOSYSIEMATICS VOL. 22, No. 2

Table 2. Frequencyof mesopostnotalsetae/scales in the tribesAedini and Culicini with includedgenera.

Category Number species Number species Frequencyof Zoogeographical with Mpn in category Mpn character regions*where character in category characterfound

Aedini 55 1,146 0.048 AA, AT, NT, 0 Aedes 2 940 0.002 -0 Armigeres 1 46 0.022 0 Eretmapodites 18** 44 0.409 AT Haemagogus 9 32 0.281 NT Heizmannia 25 30 0.833 0 opt@ 0 1 SP Psorophora 0 47 NA/NT Udaya 0 2 0 Zeugnomyia 0 4 0

Culcini 12 759 0.016 NT cl&x 4 740 0.005 NT Deinocerites 7 18 0.389 NT Galindomyia 1 1 1.0 NT

*Australasian(AA), Afrotropical (AT), Nearctic (NA), Neotropical (NT), Oriental (0), South Pacific (SP). **Probably an underestimateas 24 speciesof Eretmapodites were unavailablefor examination.

Table 3. Frequencyof mesopostnotalsetae/scales in the tribe Sabethiniand includedgenera.

Number species Number species* Frequencyof Zoogeographical Category with Mpn in category Mpn character regions** where Character in category characterfound

Sabethini 226* 377* 0.599 NA, NT, AA, 0 Johnbelkinia 3 3 1.0 NT Limatus 8 8 1.0 NT Malaya 0 12 AT, 0 Maorigoeldia 0 1 SP Phoniomyia 23 23 1.0 NT Runchomyia 11 11 1.0 NT Sabethes 30 30 1.0 NT Shannoniana 3 3 1.0 NT Topomyia 0 38 0 Trichoprosopon I3 13 1.0 NT Tripteroides 19*** 119 0.160 AA,0 Wyeomyia 116 116 1.0 NA, NT

*Many speciesof Sabethiniare undescribed,and many more remain to be discovered. **Australasian(AA), Afrotropical (AT), Nearctic (NA), Neotropical (NT), Oriental (0), South Pacific (SP). ***Probably an underestimateas 14 speciesof subgenusRachisoura were unavailablefor examination.

describedsince 1941. White (1980) listed44 speciesin this ited in comparisonwith the British andFrench collections genus.Most of thesemore recentlydescribed species were andwe havenot been able to examinespecimens of the 24 differentiatedon the basisof male genitaliccharacters, and more recentlydescribed species. Rickenbach and Eouzan other characterssuch as Mpn setaewere rarely mentioned. (1970) describedferaraiandgermaini; Ferrara andEouzan The SmithsonianAfrotropical mosquitocollection is lim- (1974) describedrickenbachi and adami; and Rickenbach SEFTEMBERWO 135 and Lombrici (1975) redescribedleucopus andproductus communis (L&ester), complex (Theobald),covelli Barraud, and describedbrenguesi and jani. These 8 specieswere demeilioni MattinglyJiuzerea@&ester), greenii (Theobald), differentiatedby genitalia,and Mpn setaewere not men- himalayensis Edwards,indica (Theobald), kana Tanaka, tioned. Rickenbachet al. (1976) listed32 speciesof Eret- Mizusawa and Saugstad,kanhsienensis Tung, lii Wu, mapodites in Cameroon, but they did not mention Mpn macdonaldi Mattingly, mattinglyi Thurman, persimilis setaeand/or scales. Mattingly,propinqua Mattingly, proxima Mattingly, reidi (4) Of the 32 speciescurrently recognizedin the Neo- Mattingly,scanloni Mattingly,scintillans Ludlow, taiwanen- tropical genus,Haemagogus, the 8 species(4 unnamed) sis Lien, andviridis Barraud. We examined17 of the above recognizedby Zavortink (1972) in the subgenusConopos- 25 speciesand confirmedthe presenceof Mpn setae,and tegus apparentlydo not exhibit Mpn scalesor setae,and in the caseof complex and taiwanensis, scales also. Seven only 9 of the 24 speciesin the subgenusHaemagogus have of the remaining 8 speciesof subgenusHeizmannia are Mpn setae or scales. Those 9 speciesbelong to the describedas havingMpn setae. We have not seenspeci- SplendensSection of Arnell(1973: 69), whichis character- mensor the descriptionof kanhsienensis Tung. ized, in part, by havingl-2 smallposteromesal setae on the (6) C&X is the second largest genus in Culicidae, mesopostnotum.They are: aeritinctus Galindo andTrap- containingover 740 speciesin 20 subgenera. In major ido, argyromerisDyar and Ludlow, boshelli Osorno-Mesa, revisionsthis genusis generally describedas having the celeste Dyar and Nunez Tovar, chalcospilans Dyar, iridi- mesopostnotumbare (Lane 1953,Belkin 1962,Tanaka et color Dyar, ZuciferHoward, Dyar and Knab, regalis Dyar al. 1979). However, this is an oversight,as at least one andKnab, andsplendens Williston. Membersof both sub- specieswith Mpn setae has been known in the subgenus generawere examinedduring this studyand no deviations Melanoconion sincethe report of Rozeboom and Komp from the abovestatements were noted. (1948). In addition,we have discoveredMpn setae on 3 (5) The genusHeizmannia is confinedto the Oriental previouslydescribed species in the subgenusCarrollia. At Region and contains30 speciesin 2 subgenera(Mattingly least21 speciesare nowrecognized (3 remain unnamed)in 1970, Knight and Stone 1977, Tanaka et al. 1979). The the Neotropicalsubgenus Carrollia (Valencia 1973,Knight subgenusMattinglyia has5 speciesthat do not have Mpn 1978,Ward 1984). Althoughpossessing the classiccharac- setaeor scales(Reinert 1973). An examinationof achae- ters of the genusCr.&q members of this subgenushave tae (Leicester), catesi Lien, discrepansEdwards, and thel- divergedfrom the classicCulex habitus in also possessing mae Mattingly, in this subgenusrevealed no Mpn setae. conspicuousmetallic ornamentation, as is more com- The remaining 25 species,in the subgenusHeizmannia, monly seen in the tribes, Aedini and Sabethini. During can be recognizedby the presenceof Mpn setae,and also routine identification of South American specimenswe scaleson some species. These speciesare: aurea Brug, discoveredthat bonnei Dyar, typicallypossesses a group of aureochaeta (Leicester), chandi Edwards, chengi Lien, 4 lower Mpn setae(Fig. lA). Examinationof an additional

Stm

Figure 1. Mesopostnotalsetae and/or scalesand scutellarscales on: (A) Cule~(Car.) bonnei, (B) Galindomyia leei, (C) Culex (Mel.) batesi, and (D) Trichoprosopon digitatum. 136 MOSQUITOSY~MATICS VOL. 22, No. 2

40 specimensof bonnei from Brazil, Colombiaand French Deinocetites (as Dinomimetes) epitedeus was noted by Guiana revealedthat thisis not an anomaly,but probably Snodgrass(1912: 60) to have one or 2 coarse spinelike a fixed characterfor thisspecies. We alsoexamined adults “bristles” near the posterior margin of the Mpn. In the of an additional14 speciesof thissubgenus, and Mpn setae PseudesGroup, Adames noted that one species,mcdonaMi were generallyabsent, with the followingexceptions. Five Belkin and Hogue, usuallyhas one middle Mpn seta. The of 35 specimensofmetempsytus Dyar, from CostaRica and remainingspecies apparently do not exhibitthis character. Panamahad from 2 to 4 short,curved, dark setae,and 3 of (8) Galindomyia is a monotypic genus of crab hole 40 specimensof urichii Coquillett, from Colombia and inhabitingspecies known only from Colombia and Ecua- Brazil, had 1 and 4 setae, respectively. Valencia (1973) dor (Adames and Arzube 1975, Adames and Galindo includedmetenzpsytus in his BihaicolusGroup, which he 1975). The single species,Zeei Stone and Barreto, is consideredmore primitive, while bonnei and urichii were describedas occasionallyhaving a small posteriortuft of included in the Iridescens Group, which he considered Mpn setae near the midline (Stone and Barreto 1969). more derived. In preservedCarrolla specimens,the hal- Three specimensof Zeeiwere examinedfrom Esmeraldas teresfrequently are shriveledand coverthe posteriorpart Province,Ecuador and Mpn setae(2) were presenton only of the mesopostnotum;thus, the Mpn setaeare difficult to one specimen,while all 3 specimenspossessed from 1 to 4 see. Mpn scales(Fig. 1B). Sirivanakarn(1983) recognized149 valid speciesin the (9) Publicationson the tribe Sabethinipertaining to the Western Hemisphere subgenusMelanoconion, and sev- Western Hemisphere normally list the presenceof Mpn eral new specieshave been describedsince, thus, over 150 setaeand/or scalesas a differentiatingcharacter for spe- speciesare now recognized.Rozeboom and Komp (1948) cies in that tribe (e.g., Lane 1953, Belkin et al. 1970). describeda new species,batesi, with 10 or fewer Mpn Apparently, the presenceof Mpn setae/scalesis a good setae. This hasbeen overlookedby most C&X workers, characterfor identifyingWestern Hemisphere sabethines. exceptSirivanakarn (1983). Lane (1953: 309) described However, mostEastern Hemisphere sabethineslack Mpn the mesopostnotumof C’uZa as,“ postnotumnude ”,but on setae/scales.Approximately 377 valid speciesare recog- page491 he quotedthe originaldescription of batesi, which nized in the tribe and slightlyover half (54.9%) are known clearlymentions 10 or lessMpn setae. Sirivanakarn(1983) in the8 WesternHemisphere genera: Johnbelkinia Zavortink, usedthe presenceof Mpn setaeto differentiatehis Evan- Limatus Theobald, Phoniomyia Theobald, Runchomyia saeGroup in a key, and listed 4 speciesin this group,viz., Theobald,Sabethes Robineau-Desvoidy, Shannoniana Lane evansaeRoot, batesi Rozeboomand Komp,johnnyi Duret, and Cerqueira, Trichoprosopon Theobald, and Wyeomyia and changuinolae Galindo and Blanton. We examined8 Theobald. In the Eastern Hemisphere, the remaining specimensof batesi from Ecuador,7 had palegolden Mpn 42.5% of the valid speciesoccur in 4 genera: MaZaya setae (Fig. lC), while one specimenapparently had the Leicester,Maorigoeldia Edwards,Topomyia Leicesterand setaerubbed off. Two specimensalso exhibited 2 or 3 pale Tripteroides Giles. The only Eastern Hemisphere sabe- scales. The lectotype and one female paralectotypeof thinesexhibiting Mpn setae(no scales)are certainspecies evansae were examinedand the lectotype,although badly in Tripteroides. Accordingly, authors should consider damagedand rubbed, appearsto have 1 Mpn alveolus. speciesfrom both hemisphereswhen makinggeneralized The paralectotype has the Mpn bare. The following statementsabout sabethines.This is particularlytrue as additionalspecimens in the NMNH labelled as Evansae thisis probably the leastknown and understood tribe in the Complexor nr. evansae had Mpn setaeor alveoli: 3 males, Culicidae. Most speciesprobably remain undiscovered 2 females (Brazil); 1 male (Ecuador); 17 males (French becauseof their affinity for phytotelmic( = inhabitingsmall Guiana); 1 male,3 females(Panama); and 1 female (Peru). poolsof water within or upon plants) ovipositionsites. Usually from 2 to 6 dark setae were present on these (10) Johnbelkinia is a Neotropical genusdescribed by specimens,however, 2 males from French Guiana each Zavortink (1979a) for 6 nominal speciespreviously in- possessedone alveolus,much like the lectotypeof evansae. dudedin theold compositegenus Trichopmsqwn. Zavortink We did not examine specimensof the other 2 speciesin recognized3 of these6 nominalspecies as valid: leucopus Sirivanakarn’sEvansae Group. (Dyar and Knab), longipes (Fabricius) and ulopus (Dyar (7) Deinocetites, a Neotropicalgenus of crabhole inhab- andKnab). These3 speciespossess numerous Mpn setae, iting mosquitoes,currently contains18 species,of which7 and Zeucopusoften has dark iridescentMpn scales,while havebeen recorded with Mpn setae(Adames 1971). Adames long&s and ulopus usuallylack scaleson the Mpn. (p. 36) listeddyatiBelkin andHogue, barretoi Adames,and (11) The genusLimatus contains8 valid species,all nicoyae Adames and Hogue, the 3 membersof the Dyari restrictedto the neotropics.The speciesin Limatus are all Group, asusually having a few lower Mpn setae. He also supposedto haveMpn setaeand somepossess Mpn scales. listedepitedeus (Knab), panamensis Adames,and colom- However, Lane (1953: 1045) possiblycaused confusion on bianus Adames, 3 of the 5 members of the Epitedeus this point by describingLimatus as having,“Postnotum Group as usually having 2 or more lower Mpn setae. nude, median portion with hyaline scaleswhich easily SEPTEMBERWO 137

becomedetached. ” The 8 valid speciesare: andinus Levi- Trichoprosopon, Zavortink (1979a,1979b, 1981) indicated Castillo,asulleptus (Theobald), durhamii Theobald,$!av- that 21 specieswere recognized(several undescribed) that isetosusDe Oliveira Castro,guayasi Levi-Castillo, hoffmani would remain in the Neotropical genus Trichoprosopon, Root, martiali Senevetand Abonnenc, andpseudomethys- and he listed20 availablespecies names he wasplacing in ticus (Bonne-Wepster and Bonne). Belkin et al. (1970) thisgenus. Ward (1984), with Zavortink’s approval,listed describedthe female of hoffinani ashaving a medianpatch 15 of the availablenames as valid, with 13 asspecies and 2 of lo-20 lightgolden setae preceded by an equalnumber of asvarieties, they are: andinum Levi-Castillo,brevipes (Da broad golden scales. Clastrier and Claustre (1978) de- Costa Lima), castroi Lane and Cerqueira, compressum scribedthe female of matiiali as having a tuft of 5 or 6 Lutz, compressumvar. mogilasium (Dyar and Knab), digi- golden setae on the posterior part of the Mpn, and as tatum (Rondani), digitatum var. townsendi Stone,evansae lackingscales on the Mpn. Antunes, Zampropus (Howard, Dyar and Knab), Zanei (12) The Neotropical genus,Phoniomyia, contains23 (Antunes), obscurum Lane and Cerqueira, paZZidiventer species,all of which are supposedto possessMpn setae (Lutz), simile Lane and Cerqueira,soaresi Lane and Cer- (Theobald 1903,Correa and Ramalho 1956). Also, many queira, and vonplesseni (Dyar and Knab). It is generally of the speciesmay exhibit scaleson the Mpn. Bonne- acceptedthat all of thesespecies and varieties exhibit a tuft Wepster and Bonne (1921) describedZassaZZi as lacking of Mpn setae,and some, e.g., compressum and digitatum Mpn scales.Lourenco-de-Oliveira (1983) describeda new (Fig. lD), alsoexhibit Mpn scales.Lane (1953) indicated species,deanei, as having some setae on the median the Mpn of obscurum was“ nude.” portion of the Mpn. (17) The genusTrzpteroides Giles, is an Oriental/Aus- (13) Runchomyia, another Neotropical taxon, was re- tralasian/ South Pacific taxon that is still relatively poorly stored to generic statusby Zavortink (1979a) in his pre- known. In Knightand Stone (1977), 113species were listed liminaryreclassification of the compositegenus Ttichopms~ in 3 subgenera.However, Mattingly (1980,198l) begana pon (Zavortink1979b). In thisnew classification,Zavortink reclassificationof thisgenus and recognized5 subgenera. recognized15 availablespecies names in Runchomyia, of At present, there are approximately 119 valid species whichhe considered11 asvalid species.These 11 species recognized. As noted below, most of these specieslack were listedby Ward (1984) accordingto their subgeneric Mpn setae and/or scales. The subgenusPolylepidomyia designation.The 11 speciesare: cerqueirai (Stone),espini Theobald, which is confinedto the AustralasianRegion, (Martini), frontosa Theobald, humboldti (Lane and contains18 speciesthat havethe Mpn bare. The subgenus Cerqueira), Zunata (Theobald), magna (Theobald), para- Rachionotomyia Theobald, contains13 Indian, Southeast nensis Brethes,perturbans (Williston),reversum (Lane and Asian and Oriental-Palaearcticspecies, of whichonly one, Cerqueira), theobaldi (Lane and Cerqueira), and walcotti edwardsi (Barraud) in India and SoutheastAsia, hasa tuft (Lane and Cerqueira). All of thesespecies are presumed of Mpn setae (Barraud 1934). The third subgenus,Ra- to exhibit Mpn setae and somemay possessMpn scales. chisoura Theobald,contains 27 speciesthat are confinedto (14) The Genus Sabethes, another Neotropical taxon, the Australasianand South Pacific regions. Knight and containsapproximately 30 valid species.Edwards (1932) Stone (1977) transferred mabinii Baisas and Ubaldo- stated that speciesin this genushave Mpn setae. Lane Pagayonto the subgenusTtipteroides. Three speciesof (1953) consideredall Sabethesto haveMpn setae,but gave Rachisoura definitely lackMpn setaeor scales,i.e., exnebu- very incompletedescriptions and only mentioned2 species Zis Bonne-Wepster, fuscipZeura Lee, and Zatisquama as also possessingMpn scales,they are: belisatioi Neiva (Edwards). Five species,i.e., confusus Lee, felicitatis and tarsopus Dyar and Knab. Lane (1953) specifically Bonne-WepsterJiZipes(Walker), ZeeiPeters, and Zongipal- mentioned that amazonicus Gordon and Evans, has the patus Lee, definitely have Mpn setae, but apparentlyno “postnotumnude ”, i.e. without scales.Sabethes forattinii scales. Belkin (1962) describedmathesoni Belkin, stonei Cerqueira, was described as having the Mpn without Belkin and torokinae Belkin, as having 2-4 minute setae scales,but with a group of setae(Cerqueira 1961). (frequently absent)and no scales.One species,flabeZZiger (15) The genusShannoniana was also elevated to ge- Bdnne-Wepster,has Mpn setaeand white scales(Van den neric statusby Zavortink (1979a,1979b). Zavortink recog- Assam1959). Sloof (1961) describedtityae as havingonly nized 4 availablespecies names in thisNeotropical genus, 5-8 white Mpn scales.Authors of the remaining14 species and he considered3 asvalid species.He indicatedthat he failed to mention Mpn setaeand/or scales,viz., adentata was aware of an additional 12 new undescribedspecies, Van den Assam, bisquamatus Lee, brevirhynchus Brug, and that “scores more” await discovery. The 3 valid cuttsi Van den Assam,fuZiginosus Lee, king* Lee, pallidus specieslisted by Ward (1984) are: fluviatilis (Theobald), Lee, Papua Brug, pilosus Lee, plumiger Bonne-Wepster, moralesi (Dyar and Knab), and schedocyclia (Dyar and simplex Brug, syzvestis(Theobald), szechwanensis Hsu, Knab). These 3 speciespossess Mpn setae. Accordingto and vanleeuweni (Edwards). Lane (1953), j7uviatilis doesnot exhibit Mpn scales. The fourth subgenus,Tricholeptomyia Dyar and Shan- (16) In his preliminary reclassificationof the genus non, contains9 speciesthat are restricted to the Philip- 138 MOSQUITO SYSTEMATKS VOL. 22, No. 2 pines in the SoutheastAsian subregion. Eight of the 9 thii. Only about 5% of the currently known Aedini speciesexhibit a smalltuft of Mpn setae,they are: apoensis speciespossess the character,and of those,83.6% occurin Baisasand Ubaldo-Pagayan,bekini Baisasand Ubaldo- the Eastern Hemisphere (Table 2). In the Western Pagayan,christophersi Baisas and Ubaldo-Pagayan,delpi- Hemisphere the only members of the Aedini possessing lari Baisasand Ubaldo-Pagayan, microcala (Dyar), nepen- the characterare 9 speciesof Haemagogus, all belongingto thicola (Banks), roxasi Baisasand Ubaldo-Pagayan,and the subgenusHaemagogus. In the Eastern Hemisphere wemeri Baisasand Ubaldo-Pagayan. The singlespecies mostof the speciesof Eretmapodites (Afrotropical) proba- having the Mpn bare is barraudi Baisas and Ubaldo- bly possessthe character, however, we were unable to Pagayan. confirm this because of limited material in the Smith- The fifth and largestsubgenus, Tripteroides Giles, con- soniancollection and becauseof inadequatedescriptions tainsabout 51 specieswhich are distributedthrough north- for many of the species.The remainingspecies of Aedini easternIndia eastwardto Japanand souththrough Malay- with thischaracter occur in the Oriental and/or Australa- siaand Indonesia,New Guinea and northernAustralia to sianregions. GenusHeizmannia, often mentionedas the the Solomonsand Santa CruzIsland, Fiji (Mattingly 1981). EasternHemisphere equivalent of Haemagogus, hasMpn Mattingly (1981: 79) describedthe membersof thissubge- setae/scaleson the 25 speciesin the subgenusHeizmannia, nus as havingthe Mpn bare. but lacksthis characteron the 5 speciesin the subgenus (18) There are at least 116 valid speciesof Wyeomyia. Mattinglyia. One speciesof Heizmannia in Korea, hi Wu, As membersof the tribe Sabethini,all of thesespecies are is the northernmost (35”N) speciesthat possessesthis supposedto possessa tuft of Mpn setae (Lane 1953). characterin the EasternHemisphere. The Oriental genus However, Belkm et al. (1970) reported that 2 species, Armigeres only has one specieswith Mpn setae/scales. albosquamata Bonne-Wepsterand Bonne, andsurinamen- This genus apparently is most closely related to the sis Bruijning,have scales,but no setae. Zavortink (1985) AfrotropicalAedes subgenus,Pseudamigeres, whosespe- describedz&ala as havinglong stronglydeveloped Mpn ciesdo not possessthis character(Edwards 1941). Aedes setaeand “sometimesa few dark scales.”This genus,like is cosmopolitanand is the largest culicid genuswith 41 most of the other sabethinegenera is very poorly known, subgenerathat utilize a wide variety of immature habitats. andbecause of the associationof mostspecies with brome- The presenceof Mpn setaeon only 2 speciesofAedes in liads in the Neotropical forests,there must be numerous the subgenusVerrallina from the Oriental and Australa- undescribedspecies. E. L. Peyton (pers. comm., 1990) sianregions was a surprise. suggeststhat there are at least 160 speciesin the genus, In the tribe Culicini there are only 3 genera and all 3 based on the specimensdeposited in the Smithsonian possessspecies with Mpn setae/scales(Table 2). How- Institution. We havenot encounteredspecimens of Wyeo- ever, only about0.5% of the speciesin C&x are knownto myia that have the Mpn entirely bare. possessthis character and they occur in the subgenera Carrollia and Melanoconion. Cuk is also cosmopolitan DISCUSSION and is the secondlargest culicid genus, with 20 subgenera. The presenceof Mpn setae/scalesonly in the monotypic Referencesto mesopostnotalsetae and/or scaleswere Neotropicalgenus Galindomyia, and in certain Neotropi- scattered throughout the mosquito literature, and then cal speciesof C&x andDeinocetites wasalso unexpected. only in the adult descriptionsor group discussions.One In the EasternHemisphere the presenceof Mpn setae/ obviousresult of thissurvey is findingthe highfrequency in scalesin the tribe Sabethiniis unusual(Table 3). Of the 4 which the mesopostnotumhas been overlooked or ig- sabethmegenera in the Eastern Hemisphere, only the nored, even to the present, and not includedin descrip- genusTripteroides has specieswith thischaracter, and then tions. Recent descriptionsof new mosquitospecies have only in 3 of 5 subgenera,i.e., Rachionotomyia, Rachisoura becomeincreasingly lengthy and complex,causing sugges- and Tricholeptomyia (Mattingly 1981). The 3 Eastern tionsthat shorterdiagnostic descriptions would suffice. A Hemisphere sabethinegenera without speciesexhibiting surveysuch as this one highlightsthe value and need for the character are Malaya, Maorigoeldia and Topomyia. thoroughdescriptions. Diagnostic descriptions may meet The genusMaZaya is Oriental, Afrotropical andAustrala- the requirementsof the International Code of Zoological sianin distribution,and in the Afrotropical region it is the Nomenclature (ICZN) to validate speciesin pure alpha only representativeof the Sabethini.The genusMaorigoel- level taxonomic works, however, the taxonomy of the dia is an isolated monotypic genus found only in New Culicidaehas long sincereached a level requiring highly Zealand (South Pacific). Belkin (1962) hypothesizedthat detaileddescriptions for beta and gammalevel considera- Maorigoeldia probablyarose through a primitive Ttipteroi- tions. des ancestor. This ancestormust have lacked the Mpn The distribution of Mpn setae/scalesin the family character as do a majority of Tripteroides to this day, Culicidae is restricted to only 3 of the 10 tribes in the includingmembers of subgenusPoi ylepidomyia’ in Austra- subfamilyCulicinae, namely, Aedini, Culicini and Sabe- lia andTasmania. The third genus, Topomyia, is Oriental SEPTEMBERW~O I39

andin manyways resembles species found in the Neotropi- strictedto genusgroups and speciesconfined to the sub- calgenera related to Trichoprosopon. AlthoughTopomyia tropicsand tropics between 30 ”N and30 ”s latitudes,except speciesdo not possessMpn setae/scales,they have a for Heizmannia Zii in the Oriental Palearctic, Wyeomyia similar unique characternot found in any other genusin smithii in the Nearcticand Phoniomyia muehlensi (Petroc- the Eastern Hemisphereexcept Toxorhynchites, i.e., setae chi), Runchomyia (Run.) paranensis (Brethes), and Wyeo- and scaleson the metepisternumthat are usuallyventral myia (Men.) ZeucostigmaLutz, which extend southto ap- and caudal to the metathoracic spiracle (Marks 1971). proximately35 ”slatitude in Argentina(Mitchell andDarsie There are 38 recognized speciesof Topomyia, with 28 1985). No attempt was made to define the elevation definitely having this character (usually as setae), 4 de- limitationsfor the Mpn character. scribed as lacking this character, and the statuson the The mostobvious result of thisstudy is the clear associa- remaining6 is unknown. tion of Mpn setae/scaleswith the subtropicsand tropics In the WesternHemisphere all of the describedspecies where most plantsevolved, particularly epiphyticplants. of Sabethini apparently exhibit Mpn setae/scales,and Furthermore, most specieshaving the Mpn characterare both setae and scalesare present on many species. On alsophytotelmic in their immature stages(Table 4). The certain species,e.g., Trichoprosopon digitatum, the scales Neotropical sabethines,all of which apparently possess are translucentand difficult to see (Fig. 1D). On others, Mpn setae/scales,are definitely correlatedto the success the pale to black color of the Mpn setaeserves as a useful andwide distributionof phytotelmatasuch as bromeliads, taxonomiccharacter (e.g., Sabethes). The location of the aroids,bamboo, etc. In the Eastern Hemispherethere are setaeand/or scaleson the mesopostnotummay alsobe of no bromeliadsor an equivalentof these plantsexcept for taxonomicvalue. The northernmostspecies possessing one species,Pitcamia feliciana (A. Chevalier) Harms and the Mpn setae/scalesin the WesternHemisphere is Wyeo- MiIdbraed, in West Africa (Richards1973, Thorne 1973). myia smithii (Coquillett) which extendsto approximately The nearly completeabsence of this plant group mustbe 55”N latitude in Canada. considereda major factor in the reduced successof the Geographically,mesopostnotal setae/scales have been sabethinesin the Eastern Hemisphere,particularly in the foundon speciesin 7 of the major faunalregions; Nearctic, Afrotropical Region. The restrictionof the Mpn character Neotropical,Afrotropical, Palearctic,Oriental, Australa- betweenthe 3O”Nand 30’S latitudes(5 exceptions)would sian and South Pacific. This character generally is re- indicate the character evolved in subtropical/tropical

Table 4. Primary ovipositionhabitats of taxa with Mpn setae/scaIes.

Taxa Primary ovipositionhabitats

Aedini Aedes (Verrallina) Shadedground pools Annigeres (Leicesteria) Phytotelmata Eretmapodites Phytotelmata Haemagogus (Haemagogus) Phytotelmata Heizmannia (Heizmannia) Phytotelmata

culiciIli Culex (Carrollia) Phytotelmata C&x (Melanoconion) Freshwatermarshes, swamps and pools Deinocetites Crab holesin brackishwater Galindomyia Crab holesin brackishwater Sabethini Johnbelkinia Phytotelmata Limatus Phytotelmata Phoniomyia Phytotelmata Runchomyia Phytotelmata Sabethes Phytotehnata Shannoniana Phytotelmata Trichoprosopon Phytotehnata Ttipteroides Phytotelmata Wyeomyia Phytotelmata 140 MOSQUITOSY~MATICS VOL. 22, No. 2 ancestors.Accordingly, the exceptionsare best explained portionof their taxawiththis character. This isparticularly as: (1) Wy. smithii, asan extensionfrom the subtropicsinto true of genera and speciesin the Simuliidae and Chi- the temperate-boreal zones of the Nearctic through its ronomidae where the character is rare. There is no symbioticrelationship with Sarraceniapurpurea L., pitcher evidenceof Mpn setae/scalesin the more primitive fami- plants(Ross 1964); and (2) Hz hi, a speciestrapped on the lies of superfamily Culicoidea, namely, Chaoboridae, Korean Peninsulafrom an earlier warm period when the Corethrellidaeand Dixidae, or in selectCulicinae isolated peninsula enjoyed subtropical or tropical weather. on the New Zealand block (Belkin 1968). However, only Apparently Hz. Zii is uncommon in Korea, and found a few speciesof chaoborids,corethrellids, dixids and primi- primarily in tree holes(Tanaka et al. 1979). Heizmannia hi tive culicidsare still extant. Thus, the absenceof this also occurs in the People’s Republicof China,but southof characterin thesetaxa may be a reflection of the declineof 3O”Nlatitude in Zhejiang Province (Tanaka et al. 1979). these taxa. This may also be true for certain Culicinae The remaining3 SouthernHemisphere (Neotropical) species tribes that are currently consideredmore ancestral(Cu- probably extend south of 30”s latitude becauseof their lisetini, Ficalbiini, etc. - see Belkin 1962: 117). We feel associationwith a particular phytotelmichabitat. Mpn setae and/or scalesare an ancestral character in A possiblefunction of the Mpn setae could be tactile nematocerousDiptera that has been retained in only 3 (sensory)monitoring of the dorsalmovement of the abdo- families of the infraorder Culicomorpha:Culicidae, Chi- men by contact with the tergum of the first abdominal ronomidaeand Simuliidae. Their occurrenceon members segmentduring mating and oviposition. Certain sabethine of only 3 tribes of Culicinae in family Culicidae is a specieswith this character, e.g., Sabethes chloroptems reflection of their lossin current, more derived taxa. (Von Humboldt), have highlyspecialized adaptations for The current distributionof this characterin Culicidae projectingova into tiny holesin bamboo (Galindo 1957). probablyarose throughcertain protoaedines,protoculic- However, there are other Western Hemisphere sabe- inesand protosabethines located in warm tropicalareas of thineswith Mpn setae that do not have suchadaptations Gondwana,the southernpart of Pangea. With the splitup (e.g., 7”. digitatum, see Lounibos 1983, Lounibos and of Gondwana,taxa possessingthis characterwere present Machado-Allison 1983). In Aedini, the only speciesin on the Indian-SoutheastAsian, Antarctica-Australian, Armigeres with Mpn setaeis Ar. (Lei.) flavus whichalso has African and South American blocks, where the present an unusualoviposition behavior pattern in which ova are speciesevolved and spreadto their current distributions. depositedon the hindlegsand dippedin the water to hatch The variousculicid components on the aboveblocks met (Macdonald 1960). To assistin this specializedbehavior, with varyingdegrees of success:(1) SouthAmerica, with the hindfemur of Ar. j?avus is much shorter than in the the evolving bromeliads, remained isolated for a long other speciesof the genus. However, the closelyrelated period with the sabethines(and the Mpn character)be- Ar. (Lei.) annulitarsis (Leicester), which does not have cominga dominantfaunal element; (2) Africa, with prac- Mpn setae/scales,has a similar type of ovipositionbehav- tically no bromeliadsand severeclimatic changes(Raven ior (Macdonald 1960, Bailey et al. 1975). In view of the andAxelrod 1975),lost most of the originalsabethines and abovedisparities and the presenceof Mpn setae/scaleson the Mpn character survived only in the unique aedine Deinocerites and Galindomyia, whose speciesoviposit in genus,Eretmapodites; (3) Antarctica,extinction; (4) India- crabholes, and on certainspecies of C&x (Melanoconion) SoutheastAsia, separatedfrom Gondwanaearly, possibly andAedes (VerraZZina)which typicallyoviposit in marshes at the same time (Ridd 1971). On the Indian block the andswamps, the function(if any) of the Mpn setaeremains aedinegenus Heizmannia wasthe mostsuccessful line with unclear. the Mpn character. The accompanyingSoutheast Asian block alsocarried the successfulHeizmannia aswell asthe PHYLOGENETIC CONSIDERATIONS ancestralTopomyia line. Otherwise,only Annigeresflavus and Tripteroidesedwardsi in both of theseareas, plus those A search of the literature for nematocerousDiptera Tnpteroidesin the Philippineshave the Mpn character;and other than Culicidae that possessMpn setae/scalesre- (5) Australia, of the successfulsabethines, most species vealedthat certainspecies in the followingfamilies possess lackedthe Mpn character,however, with the collisionwith this character: Blephariceridae (Hogue 1981), Myceto- the SoutheastAsian block, the sabethineswere rejuve- philidae (Vockeroth 1981), Sciaridae(Steffan 1981),Ani- natedby an abundanceof bambooand aroid phytotelmata. sopididae(Peterson 198la), Synneuridae(Peterson and We feel the presenceof Mpn setaeon Ar. fravus and the 2 Cook 1981), Simuliidae (Freeman and de Meillon 1954, Aedes (VerraZZina) speciesrepresent the presence of a Peterson1981b), and Chironomidae(Oliver 1981). Using relict characterlost by most of the other speciesin those the evolutionaryranking of thesefamilies (McAlpine et al. genera. Ae&s (VerraZZina) is recognized as having an 1981),only the mostprimitive family, Blephariceridae,has ancestraltype of male genitalia,i.e., possessingan opist- the characterthroughout the family. Like the Culicidae, hophallus(Reinert 1974). We alsointerpret the presence the remaining families listed above have only a small of Mpn setae/scalesin the Neotropical Culicini as relict, SEPTJZMBER1990 141 with mostof the modern specieshaving lost the character. Bonne-Wepster,J. and C. Bonne. 1921. Notes on South The species in many of the other tribes, genera and American mosquitoesin the British Museum (Diptera, subgeneraof Culicinaepossibly never possessedthe Mpn Cuhcidae). Inset. Inscit. Menst. 9:1-26. characterbecause they evolved in the cooler temperate Carpenter, SJ. and WJ. LaCasse. 1955. Mosquitoesof and boreal regions or on Laurasia, the northern part of North America (North of Mexico). Univ. Cahf. Press, Pangea.For thosewithout the Mpn characterthat evolved Berkeley and Los Angeles. 360 pp. + 127 pl. in the tropicsor on Gondwana,their non-associationwith Cerqueira, N.L. 1961. Cinco novossabetinos da Ama- phytotehnataprobably has had somerole in the absenceof zonia (Diptera, Cuhcidae). Rev. Bras. Entomol. 10:37- the Mpn/scaIe character. 52. Clastrier, J. and J. Claustre. 1978. Limatus martiali ACKNOWLEDGMENTS Senevetet Abonnenc,1939 et Wyeomyia nigricephala n. sp.(Diptera, CuIicidae) de la Guyane Franc&e. Ann. We thankthe followingpersonnel of the Department of Parasitol.Hum. Comp. 53:539-546. Entomology, Walter Reed Army Institute of Research Correa, R.R. and G.R. RamaIho. 1956. Revisao de (WRAIR): Ralph Harbach,E.L. Peytonand Ronald Ward Phoniomyia Theobald, 1903 (Diptera, Cuhcidae,Sabe- for criticallyreviewing the manuscript;Rosetta Trite and thini). FoIia CIin. Biol. 25:1-176. JamesPecor for typingand final preparationof the manu- Edwards,F.W. 1932. Genera InsectorumDiptera, Fam. script;and Taina Litwak for preparingthe illustrations. Culicidae.Fast. 194. Desmet-Verteneuil,Brussels. 258 PP. LITERATURE CITED Edwards,F.W. 1941.Mosquitoes of the EthiopianRegion III. Cuhcineadults and pupae. Brit. Mus. (Nat. Hist.), Adames, A.J. 1971. Mosquito studies (Diptera, Cu- London. 499 pp. licidae)XXIV. A revisionof the crabholemosquitoes of Ferrara, L. and J.P. Eouzan 1974. Description de deux nouveauEretmapodites, du groupeoedipodius Graham, the genusDeinocetites. Contrib. Am. Entomol. Inst. (Ann Arbor) 7(2):1-154. 1909 (Diptera: Culicidae) decouverts au Cameroun Adames, AJ. and M.E. Arzube R. 1975. Geographical Aed& (Dicemm@) m.t#uensik n. sp. Cah.O.R.S.T.O.M. extensionof Galindomyia Zeei Stone and Barreto to Entomol. Med. Parasitol.12:3-5. Ecuador (Diptera: Cuhcidae). Mosq. Syst.7:113-114. Freeman, P. and B. de Meillon. 1954. Simuhidaeof the Adames,AJ. and P. Gahndo. 1975. Description of the Ethiopian Region. Brit. Mus. (Nat. Hist.) Publ. 194:1- immature stagesof Galindomyia leei Stone and Bar- 224. reto, 1969. Mosq. Syst.7:132-136. Gaffigan,T.V. and R.A. Ward. 1985. Index to the second Arnell, J.H. 1973. Mosquito studies(Diptera, Culicidae) supplementto “A Catalog of the Mosquitoes of the XXXII. A revisionof the genusHaemagogus. Contrib. World”, with correctionsand additions(Diptera: Cu- Am. Entomol. Inst. (Ann Arbor) 10(2):1-174. Iicidae). Mosq. Syst.17:52-63. Bailey, C.L., S. Vongpradistand K. Mongkolpanya. 1975. GaIindo, P. 1957. A note on the ovipositionbehavior of Observationsof an unusualoviposition habit in Amzig- Sabethes (Sabethoides) chloroptenrs (Von Humboldt). eres (Leicesteria) annulitarxis (Leicester). Mosq. News Proc. Entomol. Sot. Wash. 59:287-288. 35:564-565. Galindo, P., SJ. Carpenter and H. Trapido. 1951. De- Baisas,F.E. and P. Feliciano. 1953. Notes on Philippine scriptionof two new speciesof Wjeomyia and the male mosquitoes,XIII. Four new speciesof Zeugnomyia and of Sabethes tanopus Dyar and Knab. Proc. Entomol. Topomyia. Fieldiana Zool. 33:161-179. Sot. Wash. 53:86-%. Barraud, PJ. 1934. The fauna of British India, including Harbach, R.E. and K.L. Knight. 1980. Taxonomists’ Ceylonand Burma. Diptera. Vol. V. Family Cuhcidae. glossaryof mosquitoanatomy. PlexusPublishing, Inc. Tribes Megarhinini and Cuhcini. Taylor and Francis, Marlton, NJ. xi + 415 pp. London, 463 pp. Hogue, C.L. 1981. Chapter 8. Blephariceridae,pp. 191- Belkin, J.N. 1962. The mosquitoesof the South Pacific 197. In: J.F. McAIpine, B.V. Peterson,G.E. ShewelI, (Diptera: Cuhcidae). Univ. Cahf. Press,Berkeley and HJ. Teskey, J.R. Vockeroth and D.M. Wood (ed.). Los Angeles. 2 ~01s.608 and 412 pp. Manual of Nearctic Diptera. Vol. 1. Agric. Canada Belkin, J.N. 1968. Mosquito studies(Diptera, Cuhcidae) Monogr., Ottawa. 674 pp. VII. The Culicidae of New Zealand. Contrib. Am. Knight, K.L. 1978. Supplement to a catalog of the Entomol. Inst. (Ann Arbor) 3(1):1-182. mosquitoesof the world (Diptera: Culicidae). Thomas BeIkin,J.N., S J. Heinemannand WA Page. 1970.Mosqui- Say Found. Entomol. Sot. Am. 6(Suppl.):l-107. to studies(Diptera, Culicidae). XXI. The Culicidaeof Knight,K.L. andA. Stone. 1977. A catalogof the mosqui- Jamaica. Contrib. Am. Entomol. Inst. (Ann Arbor) toesof the world (Diptera: Culicidae). SecondEdition. 6( l):l-458. Thomas Say Found. Entomol. Sot. Am. 6:1-611. 142 MO~OUKO Symmmcs VOL. 22, No. 2

Lane, J. 1953. NeotropicaI Cuhcidae. Univ. Sao Paula, H.J. Teskey, J.R. Vockeroth and D.M. Wood (ed.). Sao PauIo, Brazil, 2 vols., 1112 pp. Manual of Nearctic Diptera. Vol. 1. Agric. Canada Lee,D.J.,M.M.Hicks,M. Griffiths,M.L.Debenham, J.H. Monogr., Ottawa. 674 pp. Bryan, R.C. Russell,M. Gear-yand E.N. Marks. 1987. Owen, W.B. 1977. Morphology of the thoracic skeleton The Culicidae of the Australasian Region, Vol. 4. and musclesof the mosquito,Culiseta inomata (Wii- Commonw.Dept. Hlth. andSch. Pub. Hlth. Trop. Med., ton), (Diptera: Cuhcidae). J. Morphol. l53:427-460. Univ. of Sidney, Monogr. Ser., Entomol. Monogr. 2. Patton, W.S. and A.M. Evans 1929. Insects,ticks, mites Austr. Gov. Publ. Serv., Canberra. 324 pp. and venomousanimals of medical and veterinary im- Lounibos, L.P. 1983. Behavioral convergencesamong portance.Part 1. Medical. H.R. Grubb. Ltd., Croydon, fruit-huskmosquitoes. FIa. Entomol. 66:32-41. Great Britain. 786 pp. Lounibos,L.P. andC.E. Machado-Allison.1983. Oviposi- Peterson,B.V. 1981a.Chapter 19. Anisopodidae,pp. 305- tion and eggbrooding by the mosquitoTrichoprosopon 312. In: J.F. McAIpine, B.V. Peterson,G.E. ShewelI, digitatum in cacao husks. Ecol. Entomol. 8:475-478. H.J. Teskey, J.R. Vockeroth and D.M. Wood (ed.). Lourenco-de-Ohveira,R. 1983. Sobre uma nova especie Manual of Nearctic Diptera. Vol. 1. Agric. Canada de Phoniomyia (Diptera: Culicidae)do Rio de Janeiro, Monogr., Ottawa. 674 pp. Brasil. Mem. Inst. OS. Cruz 78:501-505. Peterson,B.V. 1981b. Chapter 27. Simuhidae,pp. 355- Macdonald,W.W. 1960. Malaysianparasites XXXVIII. 391. In: J.F. McAIpine, B.V. Peterson,G.E. SheweIl, On the systematicsand ecologyof Armigeres subgenus H.J. Teskey, J.R. Vockeroth and D.M. Wood (ed.). Leicesteria (Diptera, Culicidae). Stud.Inst. Med. Res. Manual of Nearctic Diptera. Vol. 1. Agric. Canada Malaya 29:110-153. Monogr., Ottawa. 674 pp. Marks, E.N. 1971. A metapleural generic characterin Peterson,B.V. andE.F. Cook. 1981. Chapter21. Synneuri- Topomyia Leicester. Mosq. Syst.News Letter 34-6. dae, pp. 321-324. In: J.F. McAIpine, B.V. Peterson, Mattingly, P.F. 1958. The cuhcine mosquitoesof the G.E. Shewell,HJ. Teskey, J.R. Vockeroth and D.M. Indomalayan area. Part III. Genus Aedes Meigen, Wood (ed.). Manual of NearcticDiptera. Vol. 1. Agric. subgenera,Paraedes Edwards,Rhino&sea Edwards CanadaMonogr., Ottawa. 674 pp. and Cancraedes Edwards. Brit. Mus. (Nat. Hist.), Raven, P.H. and D.I. Axehod. 1975. History of the flora London. 61 pp. and fauna of Latin America. Am. Sci. 63:420-429. Mattingly, P.F. 1970. Contributionto the mosquitofauna Reinert, J.F. 1973. Notes on the genusHeizmannia and of SoutheastAsia. VI. The genusHeizmannia Ludlow the inclusion in subgenusMattingyia of two species in SoutheastAsia. Contrib. Am. Entomol. Inst. (Ann previouslyplaced tie&s subgenusuncertain (Diptera: Arbor) 5(7):1-104. Culicidae). Mosq. Syst .5: 14-23. Mattingly, P.F. 1980. An interim reclassificationof the Reinert,J.F. 1974. Medical entomologystudies - I. A new genus Tripteroides with particular reference to the intepretation of the subgenusVerraZZina of the genus Australasiansubgenera. Mosq. Syst.12164-171. Aedes (Diptera: Cuhcidae). Contrib. Am. Entomol. Mattingly, P.F. 1981. Medical entomologystudies - XIV. Inst. (Ann. Arbor) ll(l):l-249. The subgeneraRachionotomyia, Tricholeptomyia and Richards,P.W. 1973.Africa, the “Odd Man Out.“, pp. 21- Ttipteroides (Mabinii Group) of genusTtipteroides in 26. In: B J. Meggers,E.S. Ayensuand W.D. Duckworth the Oriental Region (Diptera: Culicidae). Contrib. (ed.). Tropical forest ecosystemsin Africa and South Am. Entomol. Inst. (Ann Arbor) 17(5):1-147. America:a comparativereview. SmithsonianInst. Press, McAlpine, J.F. 1981. Chapter 2. Morphology and termi- Washington. 350 pp. nology - adults, pp. 9-63. In: J.F. McAIpine, B.V. Rickenbach,A. and J.P. Eouzan. 1970. Description de Peterson, G.E. SheweIl, H.J. Teskey, J.R. Vockeroth quatre Eretmapodites nouveauxdu groupe plioleucus and D.M. Wood (ed.). Manual of Nearctic Diptera. Edwards 1941, capturesau Cameroun (Dip&a: Cu- Vol. 1. Agric. CanadaMonogr., Ottawa. 674 pp., licidae). Cah. O.R.S.T.O.M. Entomol. Med. Parasitol. McAlpine, J.F., B.V. Peterson,G.E. SheweII,H.J. Teskey, 8:131-135. J.R. Vockeroth and D.M. Wood. 1981. Chapter 1. Rickenbach,A., J.P. Eouzan, L. Ferrara and H. Bailly- Introduction,pp. l-7. In: J.F. McAlpine, B.V. Peterson, Choumara. 1976. Donnees nouvellessur la presence, G.E. ShewelI, H.J. Teskey, J.R. Vockeroth and D.M. la frequence et la repartition des Toxorhynchites et Wood (ed.). Manual of NearcticDiptera. Vol. 1. Agric. Culicinae (Diptera, Culicidae) au Cameroun. 2. Genres CanadaMonogr., Ottawa. 674 pp. Eretmapodites et CuZex. Cah. O.R.S.T.O.M. Entomol. Mitchell, CJ. and R.F. Darsie, Jr. 1985. Mosquitoesof Med. Parasitol.1493-100. ArgentinaPart II. Geographicdistribution and bibliog- Rickenbach,A. andG. Lombrici. 1975. LesEretmapodites raphy (Diptera: Cuhcidae). Mosq. Syst.17~279-360. du groupeZeucopus Graham, 1909. Position systema- Oliver, D.R. 1981. Chapter 29. Chironomidae,pp. 423- tique des deux sous-especesactuallement connues et 458. In: J.F. M&pine, B.V. Peterson,G.E. Shewell, descriptionde deux especesnouvehes (Diptera: Cu- SEPTEMBER1990 143

licidae). Bull. Sot. Entomol. France 80:39-42. cal Africa and tropical America, pp. 27-47. In: B.J. Ridd, M.F. 1971. South-eastAsia asa part of Gondwanal- Meggers, EA. Ayensu and W.D. Duckworth (ed.). and. Nature 234:531-533. Tropical forest ecosystemsin Africa and SouthAmer- Ross, H.H. 1964. The colonizationof temperate North ica: a comparative review. Smithsonian Institution America by mosquitoesand man. Mosq. News24:103- Press,Washington. 350 pp. 118. Thurman, E.B. 1959. A contributionto a revisionof the Rozeboom, L.E. and W.H.W. Komp. 1948. Three new Culicidaeof northern Thailand. Univ. Md. Agric. Exp. speciesof Culex(Diptera: Culicidae)from Colombia. J. Sta. Bull. A-100:1-177. Parasitol.34:396-406. Valencia, J.D. 1973. Mosquito studies (Diptera, Cu- Sirivanakarn,S. 1983. A review of the systematicsand a licidae) XXXI. A revisionof the subgenusCarroZZia of proposedscheme of internal classificationof the New CuZex.Contrib. Am. Entomol. Inst. (AnnArbor) 9(4):1- World subgenusMelanoconion of C&X (Diptera, Cu- 134. licidae). Mosq. Syst.14265333. Van den Assam,J. 1959. Notes on New Guinean species Sloof, R. 1961. Tripteroides (Rachisoura) tityae (Diptera: of Tripteroides, SubgenusRachisoura (Diptera, Cu- Culicidae),a new speciesfrom Netherlands,New Guinea. licidae) with descriptionsof two new species.Tijdsch. Entomol. Ber. Amst. 21:22-23. Entomol. 102:35-55. Snodgrass,R.E. 1912. The thorax, pp. 55-60. In: L.O. Vockeroth, J.R. 1981. Chapter 14. Mycetophilidae,pp. Howard, H.G. Dyar and F. Knab. The Mosquitoesof 223-246. In: J.F. McAlpine, B.V. Peterson,G.E. Shew- North and Central America and the West Indies. Car- ell, H.J. Teskey,J.R. Vockeroth and D.M. Wood (ed.). negie Inst. Wash. Publ. 159. Manual of Nearctic Diptera. Vol. 1. Agric. Canada Steffan, WA. 1981. Chapter 15. Sciaridae,pp. 247-255. Monogr., Ottawa. 674 pp. In: J.F. McAlpine, B.V. Peterson,G.E. Shewell,H.J. Ward, R.A. 1984.Second supplement to “A Catalogof the Teskey,J.R. Vockeroth and D.M. Wood (ed.). Manual Mosquitoesof the World” (Diptera: Culicidae). Mosq. of Nearctic Diptera. Vol. 1. Agric. CanadaMonogr., Syst.16:227-270. Ottawa. 674 pp. White, G.B. 1980. Family Culicidae, pp. 114-148. In: Stone,A. and P. Barreto. 1969. A new genusand species R.W. Crosskey,(ed.). Catalogueof the Diptera of the of mosquitofrom Colombia,Galindomyia Zeei(Diptera, Afrotropical region. Brit. Mus. (Nat. Mus.), London. Culicidae,Culicini). J. Med. Entomol. 6:143-146. 1437 pp. Stone,A., K.L. Knight and H. Starcke. 1959. A synoptic Zavortink, T.J. 1972. Mosquito studies (Diptera, Cu- catalog of the mosquitoesof the world (Diptera, Cu- licidae) XXVIII. The New World speciesformerly licidae). Thomas Say Found. Entomol. Sot. Am. 6:1- placedinAedes (Finlaya). Contrib.Am. Entomol. Inst. 358. (Ann Arbor) 8(2):1-206. Tanaka, K., K. Mizusawa and E.S. Saugstad. 1979. A Zavortink, T.J. 1979a. Mosquito studies(Diptera, Cu- revision of the adult and larval mosquitoesof Japan licidae)XXXV. The new sabethinegenus JohnbeZkinia (includingthe RyukyuArchipelago and the Ogasawara and a preliminary reclassificationof the composite Islands)and Korea (Diptera: Culicidae). Contrib.Am. genus Trichoprosopon. Contrib. Am. Entomol. Inst. Entomol. Inst. (Ann Arbor) 16:l-987. (Ann Arbor) 17(l):l-61. Theobald,F.V. 19Ola. The classificationof mosquitoes.J. Zavortink, T.J. 1979b. A reclassificationof the sabethine Trop. Med. 4:229-235. genusTrichoprosopon. Mosq. Syst.11:255-257. Theobald, F.V. 19Olb. A monographof the Culicidaeor Zavortink, T.J. 1981. Speciescomplexes in the genus mosquitoes. Vol. 1. Brit. Mus. (Nat. Hist.), London. Trichoprosopon. Mosq. Syst.13182-85. 424 pp. Zavortink, T.J. 1985. Zinzala, a new subgenusof Wyeo- Theobald, F.V. 1903. A monographof the Culicidae or myia with two new speciesfrom pitcher-plantsin Vene- mosquitoes. Vol. 3. Brit. Mus. (Nat. Hist.), London. zuela (Diptera, Culicidae,Sabethini). Wasmann J. Biol. 359 pp. 43:46-59. Thorne, R.F. 1973. Floristicrelationships between tropi-