COMPARATIVE ANATOMY OF THE FEMALE GENITALIA OF GENERA AND SUBGENERA IN TRIBE AEDINI (DIPTERA: CULICIDAE). PART XI. GENUS WILLISTON

John F. Reineft I

Centerfor Medical, Agriculturaland VeterinaryEntomology (CMAVE), United StatesDepartment of Agriculture,Agricultural Research Service, 1600/1700 S.W. 23rd Drive, Gainesville,Florida 32608-1067 USA

Abstract. A comparative,morphological analysis of the femalegenitalia of speciesof the genus HaemagogusWilliston was conducted. Based on theanalysis, the femalegenitalia of the genusare characterized,a key to the includedsubgenera (i.e., ConopostegusDyar andHaemagogus) is given, anda comparisonwith othertaxa is provided.The femalegenitalia of thetwo currentlyrecognized subgeneraare described. Treatmentof the genital morphologyof each subgenusincludes a compositedescription, description ofsubcategories, detailed description and illustration of thetype species,list of thespecies examined, list of publishedillustrations of specieswith theircitations, and a discussion. The discussion section contains a list of the most distinctive features that characterize speciesof eachsubgenus, a comparisonwith othersubgenera, and otherpertinent information.

TABLE OF CONTENTS

ABSTRACT ...... 17 INTRODUCTION ...... 17 MATERIALS AND METHODS ...... 18 FEMALE GENITALIA OF GENUS HAEMAGOGUS WILLISTON ...... 19 KEY TO SUBGENERA AND ASSEMBLAGES BASED ON FEMALE GENITALIA ...... 21 FEMALE GENITALIA OF SUBGENERA OF HAEMAGOGUS ...... 21 SUBGENUS CONOPOSTEGUS DYAR ...... 21 SUBGENUS HAEMAGOGUS WILLISTON ...... 22 ACKNOWLEDGMENTS ...... 24 LITERATURE CITED ...... 25 LIST OF FIGURES ...... 27 LIST OF FIGURE ABBREVIATIONS USED IN THE TEXT AND/OR FIGURES ...... 27 FIGURES ...... 28 SYSTEMATIC INDEX ...... 30

INTRODUCTION

This is the eleventhin a seriesof papersby the authorthat describesthe female genitalia of the generaand subgeneraincluded in tribe Aedini of family Culicidae. Part I of the series (Reinert2000b) includeda brief historicalbackground of publishedpapers dealing with the subject

•Also collaborator,Walter Reed BiosystematicsUnit (WRBU), National Museum of Natural History, SmithsonianInstitution, Washington, DC 20560-0165. 18 Contributionsof the AmericanEntomological Institute 32 (5) andprovided an introductionto the series,part II (Reinert2000c) dealt with genusPsorophora Robineau-Desvoidy,part III (Reinert2000d) with genus Udaya Thurman, part IV (Reinert2000e) with genusZeugnomyia Leicester, part V (Reinert2000f) with genusAedes Meigen, part VI (Reinert2001a) with genusAyurakitia Thurman, part VII (Reinert200lb) with genusOpifex Hutton,part VIII (Reinert2001c) with genusI/errallina Theobald, part IX (Reinert2001d) with genusEretmapodites Theobald, and Part X (Reineft2002) with genusHeizmannia Ludlow. This papercovers genus Haemagogus Williston.

MATERIALS AND METHODS

A comparative,morphological analysis of the femalegenitalia of the subgeneraof Haemagogus(i.e., ConopostegusDyar andHaemagogus) was conducted, a characterizationof the femalegenitalia of thegenus is given,a keyto theincluded subgenera is provided, and a discussion that includesa comparisonwith othertaxa is included.The genitaliaof the two currentlyrec- ognizedsubgenera of the genusare described.The formatused for eachsubgenus includes a compositedescription, a detailed description and illustration of thetype species, a listof thespecies examined,a listof publishedillustrations of specieswith theircitations, a discussionincluding the most distinctivefeatures, and other items of note. Withinthe subgenusHaemagogus, the term Assemblage is usedto avoidconfusion with otherrecognized informal categories (e.g., Section, Group). Assemblage, as used here, refers to a collectionof specieswith similarmorphological features of the femalegenitalia. The term is not meantto be a designatedtaxon; however, the groupingof speciesin the subgenuscorrelates with the Albomaculatusand SplendensSections of Arnell (1973). Femalegenitalia of genusHaemagogus are considered here to includeall structurescaudad of abdominalsegment VII. SegmentVIII is includedsince its tergum and sternumare often modifiedin developmentand shape, and possess specialized setae. Terminologyused in the descriptionsand illustrationsfollows Reinert (2000b) and the abbreviationsused are foundin the "List of AbbreviationsUsed in the Text and/orFigures" that precedesthe figures. Morphologicaldescriptions are basedon slide-mountedgenitalia that were dissectedfrom nonliving,dried females. Measurementsand descriptions of femalegenital struc- turesare basedon specimensthat were cleared,dissected, arranged in a dorsoventrallyflattened position,and mounted in Canadabalsam under glass cover slips on microscopeslides. Rangesof measurementsand setalcounts given in the key anddescriptions are basedon the species(listed under"species examined" section of each subgenus)and the specimensthat I have examined, thereforefor a specificsubgenus some variation may occurin speciesnot seen. A phasecontrast microscopewas used becausethis was usuallynecessary to determinesome structuresof the spermathecaleminence on the roof of the vagina. Measurementsof structures(e.g., lengthand width oftergaVIII andIX, sternumVII1, cercus,etc.) include only the pigmentedand sclerotized areasand were made at 400X magnificationusing an ocularmicrometer having a linearscale of 100 divisionsthat hadbeen calibrated using a stagemicrometer. The scalesused in the illustrationsare in millimeters. The methodof preparationof specimensfollows Reinert (2000b). Duringdissection of the genitaliaextra care was taken when separatingthe insulaand lowervaginal lip from sternum VIII sincethe distalportion of the insulacontaining the setaeoften breaks off andremains attached to the apical intersegmentalmembrane of sternumVIII. Thesestructures tend to fold beneath sternumVIII and becomeobscured. Also, the setaecan become detached and lost during the Reineft:Female genitalia of Aedini - genusHaemagogus 19 dissection.These conditions were noted in a numberof specimensthat were used in Arnell's(1973) study.

FEMALE GENITALIA OF GENUS HAEMAGOGUS WILLISTON

Genus description. Segments VII and VIII. Laterally compressed. Tergum VIII. Proximal 0.20-0.60 retractedinto segmentVII; heavily pigmented;moderately long; wide; apex convex,with severalmoderately long and few shorter,stout setae; base wider thanapex, very gently andbroadly concave but with lateralcomers slightly produced cephalad; numerous broad, spatulate scalesdensely covering distal 0.61-0.92; setaeon distal0.32-0.77; basallateral seta present (rarely on only one side);VIII-Te index 0.56-0.94; VIII-Te/IX-Te index 1.46-2.38; length0.23-0.42 mm; width 0.31-0.57 mm. Sternum VIII. Heavily pigmented;width normally slightly greaterthan length;apex very gentlysloping caudally from apicolateralcomers to minute(0.01-0.03 of dorsal VIII-S length),median emargination separating very small lobe on eachside of midline or apex nearly flat, numerousshort, mostly straight,slender setae on apical margin; apex and base approximatelyequal in width; basevery gentlyconvex; spatulate scales few to numerous,forming elongatepatch on each side of distal 0.66-0.99, patchnarrow distally and broaderproximally; numerousshort and few moderatelylong, slender setae on distal0.95-1.00; apical, intersegmental membranelightly pigmented but few speciesalso with moderatelypigmented spots; intersegmental membranebetween segments VII andVlli shortor moderatelylong; VIII-S index0.76-1.08; length 0.30-0.49 mm; width 0.35-0.50 mm. TergumIX. Heavily pigmented;more or lessV- or Y-shaped; apex wider than base,with moderatelylong to long, narrow,thumb-shaped lobe on each side of midline, apicalmargin of lobedarker and with setaeabsent or with 1-3 shortsetae on eachlobe, 2-5 total setae;IX-Te index 0.96-1.60; length0.12-0.25 mm; width 0.11-0.19 mm. lnsula. Long; tonguelike;with 2-5 moderatelylong to long, slendersetae. Lower vaginal lip. Moderately pigmented;narrow; lower vaginal sclerite absent. Upper vaginallip. Heavily pigmented;narrow; posteriormargin evenly rounded;hinge narrow; upper vaginal sclerite absent. Spermathecal eminence. Membranous;with few short,simple spermathecal eminence spicules. Postgenita!lobe. With short spiculesmostly arrangedin small groupsgiving surfacerough appearance;heavily pigmented; long; moderately wide to wide; apex broadly rounded or with small, median emargination;ventral surfacewith 8-25 setaedistally on eachside ofmidline, 18-49 total setae(1 pair of apicalsetae moderately long and remainder short); dorsal surface with 1 pair (very rarely 2 pairs) of very shortsetae on distal 0.40-0.50; ventral PGL/Ce index 0.75-0.95; dorsalPGL index 0.84-1.28; ventral?GL index 1.52-2.40;ventral length 0.12-0.21 mm. Cercus.Heavily pigmented; moderatelylong; moderately wide to wide; apextruncate; apical margin normally with 3 (rarely 4) long, very stout,somewhat flattened (especially inner and outer) setae and few short,curved setae; lateral margin normally with 1 long and 1 moderatelylong (only I setawhen 4 setaeon apical margin),stout setae on distalportion; scales normally absent but somespecies with 1 or few broad, spatularescales on one or both cerci;dorsal surface with severalshort and 1,2 long setaeon distal 0.40-0.80; mesalmargin with patchof short,curved setae on distal area;cercus index 1.68-2.93; Ce/dorsal PGL index 1.79-2.70; length 0.14-0.23 mm; width 0.06-0.10 mm. Spermathecal capsules'. One large and 2 medium;heavily pigmented;spherical; with few to severalsmall, spermathecalcapsule pores near orifice. Accessory gland duct. Basal,lightly to heavilypigmented area short. Discussion.Haemagogus was originally described by Williston in 1896. The mostrecent revisionsof the genusare by Zavortink(1972) who provideda detaileddescription of subgenus 20 Contributionsof the AmericanEntomological Institute 32 (5)

Conopostegus,resurrected the name,removed it from synonymywith subgenusFinlaya Theobald (formerlyin genusAedes, see Reinert 2000a), and included it asa subgenusof Haernagogus,and by Arnell (1973) who revisedthe genusHaernagogus and provideddetailed descriptions of the speciesincluded in subgenusHaemagogus. Similarity of the female genitalia,as shownby the present study, strongly supportthe conclusionsof Zavortink (1972) and Arnell (1973) that Conopostegusbelongs in the genusHaernagogus. All specieswithin eachsubgenus and amongthe two subgeneradisplay a greatamount of similarity. The followingcombination of featuresis mostdistinctive for the femalegenitalia of genus Haernagogus:tergum IX isV- or Y-shapedwith the apical margin darker; sternum VIII hasthe apex very gently slopingcaudally from the lateroapicalcorners to a minute,median emargination that separatesa pair of small lobes or the apex is fiat, setaeon the apical margin are short,mostly straight,and slender,numerous primarily short setae cover the scleritefrom the apexto the baseor nearlyto the base,and the baseis gentlyconvex; insula is long,tonguelike, and bears two to five moderatelylong to long,slender setae; cercus is moderatelylong and moderately wide to wide, the apex is moderatelyto stronglytruncate with normallythree (rarely four) long,very stout,somewhat flattenedsetae along the apical margin, and the lateral margin has normally one long and one moderatelylong (only onemoderately long seta when there are four setaeon the apex),stout setae distally;postgenital lobe is long,moderately wide to wide, is coveredwith smallgroups of short spiculesthat give it a roughappearance, the ventralsurface bears a pair of moderatelylong setae apicallyand 16-49 shortsetae distally, and the dorsalsurface has a pair (very rarely two pairs)of very shortsetae at aboutmidlength; and the uppervaginal sclerite is absent. A resemblanceof the adulthabitus of Haernagogusand Heizrnannia Ludlow was noted by Edwards(1922, 1932), Mattingly(1957, 1970), Arnell (1973), andothers. The femalegenitalia of the two genera,however, are very different. Thoseof Heizrnanniadiffer notablyin the following: upper vaginal scleriteis presentand moderatelylarge; insula bears three to six very short setae (subgenusHeizrnannia), or setaeare absentand one or two small tuberculiare usuallypresent (subgenusMattinglyia Lien); cercus is broad (however, cercus also is broad in subgenus Conopostegus)and usually bears four or five (rangefour to six) long,stout (but notvery stout) setae apically;tergum IX is shapeddifferently and bearstwo to six shortsetae apically (except setae absentin Hz. reidi Mattingly); and sternumVIII hassetae covering less of the surface(distal 0.75- 0.89) and hasa differentshape, especially the apicaland basal margins. Arnell(1973) alsoindicated "Members of theKochi Group ofAedes (Finlaya) of Southeast Asia and the South Pacific show remarkablesimilarities to Haernagogusin the male genitalia, especiallyin the aedeagusand proctiger, the sidepieceincluding the specializedscales on the distal sternomesalmargin, the clasperand the reductionoftergite IX. Howeverthere is little in the adults or immaturesstages to indicaterelationship to Haernagogus."The female genitaliaof the Kochi Group of Finlaya alsodiffer from thoseof Haernagogus,especially in the following: insulais liplike;upper vaginal sclerite is well developed;postgenital lobe is differently developed; and cercus hasthe apicalmargin broadly rounded and bears about five longand moderately long, stout setae, and the dorsalsurface has numerous broad, spatulate scales on the distal0.45-0.74. The presenceofsetae on the tonguelikeinsula of genusHaernagogus is unusualfor Aedini, but this conditionalso is found in generaAyurakitia, , and subgenusHeizrnannia of genusHeizrnannia; however, in the lasttaxon the setaeare very short. Reinert:Female genitalia of Aedini - genusHaemagogus 21

KEY TO SUBGENERA AND ASSEMBLAGES BASED ON FEMALE GENITALIA

1. Cercusshorter, length usually 0.14-0.16 mm (rarely 0.18 ram), index 1.68-2.00;sternum VIII with apicalmargin nearly flat; ventralpostgenital lobe index 1.52-1.69 .... subgenus Conopostegus

Cercuslonger, length usually 0.18-0.23 mm (rarely0.17 ram), index2.07-2.93; sternumVIII with apical margin convex; ventral postgenital lobe index 1.78-2.40 (subgenus Haemagogus)...... 2

2. TergumIX withoutsetae ...... Splendens Assemblage

Tergum IX with 1-3 setaeon eachapical lobe (rarely one of lobeswithout setae) ...... Equinus Assemblage

FEMALE GENITALIA OF SUBGENERA OF HAEMAGOGUS

SUBGENUS CONOPOSTIGUS DYAR (Fig. 1)

Subgenusdescription. Tergum VIII. Proximal0.40-0.60 retractedinto segmentVII; coveredwith minutespicules; apex convex, with severalmoderately long and few shorter,stout setae;numerous scales densely covering distal 0.75-0.87; few moderatelylong and severalshort setae(in additionto apicalsetae) on distal0.42-0.63 andmore numerous along margins ofsclerite; basallateral seta present; VIII-Te index0.61-0.94; VIII-Te/IX-Te index1.55-2.23; length 0.23-0.31 mm; width 0.31-0.45 ram. Sternum VIII. Coveredwith minutespicules; heavily pigmentedbut Hg. leucocelaenus(Dyar and Shannon)and Hg. leucotaeniatus(Komp) with narrow, lightly pigmented,median stripe; width greater than length; apex very gentlyconvex and nearly flat, minute (0.01-0.02 of dorsalVIII-S length),median emargination present or absent;lateral margins nearly straight;number of broadand few moderatelybroad, spatulate scales forming elongate patch on each sideon distal0.79-0.88; numeroussetae on distal0.95-0.98; basolateral seta normally near basal margin;setae 1-5-S present,1-3-S in more or lesssubmedian line and 4,5-S slopingdiagonally to apicolateralcomer, 1-S near basomesal margin; apical, intersegmental membrane lightly pigmented; intersegmentalmembrane between segments VII andVIII short;VIII-S index0.81-0.91; length 0.30- 0.39 ram;width 0.35-0.43 ram. TergumIX. Coveredwith minutespicules; more or lessV-shaped; apexwith long,narrow, thumb-shaped lobe on eachside ofmidline, lobeoften with few longitudinal wrinkles,apical margin of lobesomewhat truncate, more darkly pigmented and bearing few longer, stouterspicules (often on small tuberculi), setae absent; area proximal to lobesrelatively short, basal margin convex; IX-Te index 0.96-1.18; length 0.12-0.19 ram; width 0.12-0.18 ram. lnsula. Coveredwith minutespicules; moderately pigmented but some specimens with distal0.20-0.30 more lightlypigmented; 4 moderatelylong to long,slender setae on distal0.15-0.50. Lower vaginallip. Coveredwith shortspicules; ventral tuft present.Upper vaginal lip. Coveredwith shortspicules. $permathecaleminence. Posterior 0.75 somewhatovoid and basal area narrower; few short,simple spermathecaleminence spicules on basolateral areas. Postgenita!lobe. Covered with shortspicules, lateral marginswith longer, stouterspicules, those at approximatelymidventral length with basolateraldenticles; moderately long; wide; apexbroadly rounded or with small (0.03 of dorsal 22 Contributionsof the AmericanEntomological Institute 32 (5)

PGL length),median emargination; ventral surface with 11-25setae distally on each side ofmidline, 22-49 total setae(1 pair of apicalsetae moderately long and remainder short); dorsal surface with 1 pairof very shortsetae on distal0.50; basalmesal apodeme present or absent;ventral PGL/Ce index0.75-0.92; dorsal PGL index0.84-1.02; ventral PGL index1.52-1.69; ventral length 0.12-0.16 ram. Proctiger. With scatteredsmall spicules;membranous. Cercus. Covered with shortto moderatelylong spicules; relatively wide; apex wide, strongly truncate; apical margin with 3 long, stout,somewhat flattened, nearly straight setae and few short,curved setae; lateral margin with 1 longand 1 moderatelylong, stout setae on distalportion; base with basomesalextension; scales absentor only 1-4 broad,spatulate ones on one or bothcerci; dorsal surface with severalshort and 2 longsetae on distal0.52-0.67; ventral surface with few shortsetae distally near apical margin; mesalmargin slightlyconcave on basal0.40-0.50; cercusindex 1.68-2.00; Ce/dorsalPGL index 1.79-2.41;length 0.14-0.18 ram; width 0.08-0.10 mm. Sœermathecalcapsules. One largeand 2 medium;heavily pigmented; spherical; with few to severalsmall, spermathecal capsule pores near orifice. Accessorygland duct. Basal,moderately pigmented area short. Type speciesdescription (Hg. leucocelaenus,Fig. 1). TergumI/III. Scalescovering distal 0.75-0.83;setae on distal 0.42-0.61; VI II-Te index0.61-0.75; VII I-Te/IX-Te index1.55-2.05; length 0.25-0.31mm; width0.35-0.45 mm. SternumI/III. Numberof scalesin lateralpatches on distal 0.80-0.88;setae covering distal 0.93-0.98; VlII-S index0.81-0.91; length 0.32-0.39 ram; width 0.35- 0.43 mm. Tergum IX. IX-Te index 0.97-1.18; length 0.14-0.19 ram; width 0.13-0.18 ram. Postgenitallobe. With 14-26 setaedistally (combined setae on dorsaland ventral surfaces) on each side of midline, 30-51 total setae;ventral PGL/Ce index 0.79-0.92, dorsal PGL index 0.84-0.94; ventral PGL index 1.55-1.69; ventral length0.12-0.16 mm. Cercus. Scalesnormally absent(4 scaleson 1 cercusof I specimen);setae on distal0.56-0.66; cercus index 1.68-2.00; Ce/dorsal PGL index 1.79-2.41;length 0.14-0.18 mm; width 0.08-0.10 mm. Species examined. Haemagogusclarki (Galindo, Carpenter, and Trapido), Hg. leucocelaenus,and Hg. leucotaeniatus. Discussion. The following combinationof featuresis most distinctivefor the female genitaliaof subgenusConopostegus: cercus is moderatelylong (lengthis usually0.14-0.16 ram, rarely 0.18 mm) andwide, the apexis stronglytruncate and bearsthree long, very stout,somewhat flattened,nearly straight setae and severalshort, curved setae, the lateralmargin has one long and one moderatelylong, stoutsetae on the distalportion, and the cercusindex is 1.86-2.00;sternum VIII hasthe apexnormally nearly flat, the lateralmargins are nearlystraight, and numerous setae coverthe distal0.95-0.98; tergum IX is V-shaped,setae are absent,and the areaproximal to the lobesis relativelyshort; and postgenitallobe is long, wide, bearsa pair of moderatelylong setae apicallyand 22-49 shortsetae on the distalarea of the ventralsurface, and the ventralpostgenital lobe index is 1.52-1.69. Publishedillustration of female genitalia. Zavortink(1972), Hg. leucocelaenus.

SUBGENUS HAEMAGOGUS WILLISTON (Fig. 2)

Subgenusdescription. Tergum I/III. Proximal 0.20-0.45 retractedinto segmentVII; coveredwith minutespicules; apex convex,with severalmoderately long and few shorter,stout setae;numerous scales densely covering distal 0.61-0.92; few moderatelylong and shortsetae (in additionto apical setae)on distal0.32-0.77 and more numerousalong marginsof sclerite;basal lateralseta present (rarely on only one side);VIII-Te index0.56-0.82; VIII-Te/IX-Te index 1.46- 2.38; length 0.25-0.42 mm; width 0.37-0.57ram. Sternum I/1II. Coveredwith minutespicules; Reineft: Femalegenitalia of Aedini - genusHaemagogus 23 width slightlygreater than length (rarely slightly less than length); apex very gentlysloping caudally from apicolateralcomers to minute (0.01-0.03 of dorsal VIII-S length), median emargination separatingvery small lobe on eachside of midline; lateralmargins slightly bowed outward;basal marginvery gently convex; broad, spatulare scales few to numerous,forming elongate patch on each sideon distal0.68-0.99; numerousshort and few moderatelylong, slender setae on distal0.97-1.00; basolateralseta usually present;setae 1-4 (5)-S in more or less, irregular,diagonal line or in submedianline, I-S near basomesalmargin, 4- or 5-S apicolateralbut some distancefrom apicolateralcomer; apical, intersegmental membrane lightly pigmented but somespecies with few moderatelypigmented spots or entirelymoderately pigmented; intersegmental membrane between segmentsVII andVIII shortor moderatelylong; VIII-S index0.76-1.08; length 0.34-0.49 mm; width 0.37-0.50 mm. Tergum IX. Coveredwith minute spicules;more or lessY-shaped; apex with moderatelylong to long,narrow, thumb-shaped lobe on eachside ofmidline, apicalmargin of lobe somewhattruncate or pointed,more darkly pigmented and bearing few longer,stouter spicules (often on smalltuberculi), setae absent (Splendens Assemblage) or with 1-3 short,slightly curved setae on eachlobe (rarely one of lobeswithout setae) (Equinus Assemblage), 2-5 total setae;area proximal to lobesrelatively long, basal margin convex; IX-Te index 1.03-1.60;length 0.16-0.25 mm; width 0.11-0.19 mm; dorsalspheres present, small, darkly pigmented. Insula. Coveredwith minute spicules;moderately pigmented (some specieswith distal 0.25-0.40 lightly pigmented);long; tonguelike;2-5 moderatelylong to long, slendersetae on distal 0.35-0.60. Lower vaginal lip. Coveredwith shortspicules; ventral tuft present.Upper vaginal lip. Coveredwith shortspicules. Spermathecaleminence. Somewhatirregularly ovoid but with basalarea flattened;few short, simplespermathecal eminence spicules on basolateralareas. Postge nita!!obe. Coveredwith short spicules,lateral margins with longer,stouter spicules, those at approximatelymidventral length with basolateraldenticles; moderately long; moderately wide; apexnormally broadly rounded but few specieswith small(0.03-0.06 of dorsalPGL length),median emargination; ventral surface with 8-23 setaedistally on eachside of midline, 18-46 total setae(1 pair ofapical setaemoderately long and remaindershort); dorsal surface with normally 1 pair (very rarely 2 pairs) of very shortsetae on distal0.40-0.50; basalmesal apodeme present or absent;ventral PGL/Ce index 0.81-0.95; dorsal PGL index0.91-1.28; ventralPGL index 1.78-2.40;ventral length 0.14-0.21 mm. Proctiger. With scatteredsmall spicules; membranous. Cercus. Coveredwith shortto moderatelylong spicules; moderatelywide (somespecies with proximalportion wider); apexmoderately wide, moderately truncate;apical margin with normally 3 (rarely 4) long, very stout,somewhat flattened, nearly straightsetae and few short,curved setae; lateral margin with normally 1 longand 1 moderatelylong (only 1 moderatelylong seta when 4 setaeon apex),stout setae on distalportion; base with narrow emarginationnear mesal margin (point at which apex ofapical lobe oftergum IX is attached);scales normallyabsent (with 1 or few broad,spatulate scales on oneor bothcerci in Hg. boshelliOsorno- Mesa,Hg. janthinomysDyar, Hg. mesodentatusKomp andKumm, andHg. soperiLevi-Castillo); dorsalsurface with severalshort and 1 (rarely 2) longsetae on distal0.40-0.80; ventralsurface with few shortsetae near apical margin; mesal margin straight or slightlyconcave at midlength;cercus index 2.07-2.93; Ce/dorsal PGL index 2.09-2.70; length 0.17-0.23 mm; width 0.06-0.10 mm. Spermathecalcapsules.One largeand 2 medium;heavily pigmented; spherical; with severalsmall, spermathecalcapsule pores near orifice. Accessorygland duct. Basal,lightly to heavilypigmented area short. Type speciesdescription (Hg. splendensWilliston, Fig. 2). Tergum VIII. Proximal0.35- 0.40 retractedinto segmentVII; scalescovering distal 0.78-0.87; setaeon distal0.60-0.77; VIII-Te index 0.72-0.75; length0.29-0.33 mm; width 0.46 mm. Sternum VIII. Number of scalesin lateral patcheson distal 0.93-0.99; setaecovering distal 0.99; apical intersegmentalmembrane lightly pigmented;VIII-S index 0.87-0.91; length 0.38-0.39 mm; width 0.42-0.44 mm. TergumIX. 24 Contributionsof the AmericanEntomological Institute 32 (5)

Apical lobespointed, without setae; length 0.17-0,21 mm; width 0.13-0.14 mm. Insula. With 4,5 setae.Postgenital lobe. Apex broadlyrounded; 9-19 setaedistally (combined setae on dorsaland ventralsurfaces) on eachside of midline,20-34 total setae;basal mesal apodeme circular; ventral PGL/Ceindex 0.81-0.87; dorsal PGL index1.14-1.18; ventral PGL index2.21-2.32; ventral length 0.16-0.17 mm. Cercus. Apex with 3 long,very stoutsetae and few short,curved setae; lateral marginwith 1 longand 1 moderatelylong, stout setae on distalportion; scales absent; dorsal surface with shortsetae on distal0.40-0.56; cercus index 2.67-2.73; Ce/dorsalPGL index2.15-2.50; length 0.18-0.21 mm; width 0.07-0.08 mm. Accessorygland duct. Basalarea lightly pigmented. Speciesassemblages within the subgenusbased on morphological characters of the female gertitalia. I. Equinus Assemblage. TergumIX. With 1-3 short,curved setae on eachapical lobe (rarely one of lobeswithout setae), 2-5 total setae. II. SplendensAssemblage. Tergum IX. Without setae. Speciesexamined. I. Equinus Assemblage. Haemagogusacutisentis Arnell, Hg. anastasionisDyar, Hg. andinusOsorno-Mesa, Hg. baresi Cerqueira,Hg. capricorniiLutz, Hg. equinusTheobald, Hg. janthinomys,Hg. mesodentatus,Hg. panarchysDyar, andHg. soperi. ii. SplendensAssemblage. Haemagogus argyromeris Dyar andLudlow, Hg. boshelli,Hg. celesteDyar andNunez Tovar, Hg. chalcospilansDyar, Hg. iridicolorDyar, Hg. lucifer(Howard, Dyar, and Knab), Hg. regalisDyar and Knab,and Hg. splendens. Discussion. The following combinationof featuresis most distinctivefor the female genitaliaof subgenusHaemagogus: cercus is moderatelylong (lengthis usually0.18-0.23 mm, rarely0.17 mm) andmoderately wide, apexis moderatelytruncate and bears normally three (rarely four) long,very stout,somewhat flattened, nearly straight setae and several short, curved setae, the lateralmargin has one longand one moderately long (only onemoderately long seta when apex has four setae),stout setae on the distalportion, and the cercusindex is 2.07-2.93; sternumVIII hasthe apexvery gentlysloping caudally from the apicolateral corners to a minute,median emargination thatseparates a pair of smalllobes, the lateralmargins are bowed outward, and numerous setae cover the distal0.97-1.00; tergum IX is Y-shaped,has one to threeshort, slightly curved setae apically on eachlobe (rarely is oneof the lobeswithout setae) (Equinus Assemblage) or bothlobes are without setae(Splendens Assemblage), and the area proximal to thelobes is relatively long; and postgenital lobe is moderatelylong, moderately wide, bearsa pair of moderatelylong setae apically and 16-44 shortsetae on the distalarea of the ventralsurface, and the ventralpostgenital lobe index is 1.78- 2.40. Coher (1948) provideda brief written descriptionof the female genitalia of genus Haemagogusbased on his examinationof two species. Publishedillustrations of female genitalia. Cerqueira(1943), Hg. capricorniiand Hg. spegazziniiBrethes; Lane (1953), Hg. capricorniiand Hg. spegazzinii;Morhig (1967), Hg. albomaculatusTheobald, Hg. equinus,Hg. iridicolor, Hg. lucifer, and Hg. regalis; and Arnell (1973), Hg.janthinomys, Hg. panarchys,and Hg. splendens.

ACKNOWLEDGMENTS

Appreciationis expressedto DonaldR. Barnard(CMAVE) for providingfacilities to completethis study; to RichardC. Wilkerson(WRBU) for providingfunding for the illustrations and for reviewingthe manuscript;to RalphE. liarbach(The Natural liistory Museum,London, Reinert:Female genitalia of Aedini - genusHaemagogus 25

UnitedKingdom) for reviewingthe manuscript; to ThomasV. Gaffgan (WRBU) andGary J. Steck (Florida StateCollection of ,Division of Plant Industry,Gainesville, FL) for the loan of specimens;and to TainaR. Litwak (Litwak IllustrationStudio, Damestown, MD) for preparing the illustrations.

LITERATURE CITED

Arnell, J. H. 1973. studies(Diptera, Culicidae)XXXII. A revision of the genus Haemagogus[sic]. Contributionsof the American EntomologicalInstitute (Ann Arbor) 10(2):1-174. Cerqueira,N. L. 1943. Algumas especiesnovas da Bolivia, e referenciaa tres especiesde Haemagogus[sic] (Diptera,Culicidae). Memoriasdo InstitutoOswaldo Cruz 39(1):1-14 (in Portuguese). Coher, E. I. 1948. A study of the female genitalia of Culicidae: with particularreference to charactersof genericvalue. EntomologicaAmericana 28:75-112. Edwards,F. W. 1922. A synopsisof adultOriental culicine (including megarhine and sabethine) mosquitoes.Part II. Indian dourhal of Medical Research10:430-475. Edwards,F. W. 1932. Genera insectorum. Diptera, Fam. Culicidae. Fascicle 194. Desmet- Verteneuil,Imprimeur-Editeur. Bruxelles, Belgium, 258 pages+ 5 plates. Lane,J. 1953. NeotropicalCulicidae VolumeII-Tribe Culicini [sic], Deinocerites,Uranotaenia, Mansonia, Orthopodomyia,Aedomyia [sic], , Psorophora,Haemagogus, tribe Sabethini [sic], Trichoprosopon,Wyeomyia, Phoniomyia, Limatus and Sabethes. Universityof SaoPaulo, Brazil, pages549-1112. Mattingly, P. F. 1957. The culicine mosquitoesof the Indomalayan Area Part H: Genus HeizmanniaLudlow. BritishMuseum (Natural History), London,United Kingdom. 57 pages. Mattingly, P. F. 1970. Contributionsto the mosquitofauna of SoutheastAsia. - VI. The genus HeizmanniaLudlow in SoutheastAsia. Contributionsof the AmericanEntomological Institute(Ann Arbor) 5(7):1- 104. Morhig, W. 1967. The taxonomicimportance of the structureof the female genitalorgans in culicidtribe Aedini. AngewandteParasitologie 8:67-100 (in German). Reinert, J. F. 2000a. New classificationfor the compositegenus Aedes (Diptera: Culicidae: Aedini), elevationof subgenusOchlerotatus to genericrank, reclassificationof the other subgenera,and notes on certainsubgenera and species. dourhal of theAmerican Mosquito Control Association 16:175-188. Reinert,J. F. 2000b. Comparativeanatomy of thefemale genitalia of generaand subgenera in tribe Aedini (Diptera:Culicidae). PartI. Introduction,preparation techniques, and anatomical terminology.Contributions of theAmerican Entomological Institute (Gainesville) 32(2): 1- 18. Reinert,J. F. 2000c. Comparativeanatomy of thefemale genitalia of generaand subgenera in tribe Aedini (Diptera: Culicidae). Part II. GenusPsorophora Robineau-Desvoidy. Contribu- tionsof the AmericanEntomological Institute (Gainesville) 32(2):19-35. Reinert,J. F. 2000d. Comparativeanatomy of thefemale genitalia of generaand subgenera in tribe Aedini (Diptera:Culicidae). PartIll. GenusUdaya Thurman. ContributionsoftheAmer- ican EntomologicalInstitute (Gainesville) 32(2):37-44. 26 Contributionsof the AmericanEntomological Institute 32 (5)

Reinert,J. F. 2000e. Comparativeanatomy of thefemale genitalia of generaand subgenera in tribe Aedini (Diptera:Culicidae). Part IV. GenusZeugnomyia Leicester. Contributions of the AmericanEntomological Institute (Gainesville) 32(2):45-52. Reinert,J. F. 2000f. Comparativeanatomy of thefemale genitalia of generaand subgenera In tribe Aedini (Diptera:Culicidae). Part V. Genus/ledesMeigen. Contributionsof theAmerican EntomologicalInstitute (Gainesville) 32(3) 1- 102. Reinert,J. F. 2001a. Comparativeanatomy of thefemale genitalia of generaand subgenera in tribe Aedini (Diptera:Culicidae). Part VI. GenusAyurakitia Thurman. Contributionsof the AmericanEntomological Institute (Gainesville) 32(4):1-7. Reinert,J. F. 2001b. Comparativeanatomy of the femalegenitalia of generaand subgenera In tribe Aedini (Diptera:Culicidae). PartVII. GenusOp•fex Hutton. Contributionsof theAmer- ican EntomologicalInstitute (Gainesville) 32(4):9-15. Reinert,J. F. 2001c. Comparativeanatomy of thefemale genitalia of generaand subgenera in tribe Aedini (Diptera:Culicidae). Part VIII. Genusl•errallina Theobald. Contributions of the AmericanEntomological Institute (Gainesville) 32(4):17-39. Reinert,J. F. 2001d. Comparativeanatomy of thefemale genitalia of generaand subgenera in tribe Aedini (Diptera:Culicidae). PartIX. GenusEretmapodites Theobald. Contributionsof theAmerican Entomological Institute (Gainesville) 32(4):41-50. Reinert,J. F. 2002. Comparativeanatomy of the femalegenitalia of generaand subgenera In tribe Aedini (Diptera:Culicidae). PartX. GenusHeizmannia Ludlow. Contributionsof the AmericanEntomological Institute (Gainesville) 32(5): 1 - 15. Williston, S. W. 1896. On the Dipteraof St. Vincent(West Indies). Transactionsof the Royal EntomologicalSociety of London44:253-446. Zavortink, T. J. 1972. Mosquitostudies (Diptera, Culicidae)XXVIII. The New World species formerlyplaced in Aedes (Finlaya). Contributions of the American Entomological Institute (AnnArbor) 8(3):1-206. Reinerr:Female genitalia of Aedini - genusHaemagogus 27

LIST OF FIGURES

Figure1. Femalegenitalia ofHaemagogus (Conopostegus) leucocelaenus (Dyar and Shannon).

Figure2. Femalegenitalia of Haemagogus(Haemagogus) splendens Williston.

LIST OF ABBREVIATIONS USED IN THE TEXT AND/OR FIGURES

AGDB : accessorygland duct base BLS : basal lateral seta BMA = basalmesal apodeme Ce = cercus DPGL = line of attachment of Pr to dorsal surface of PGL DS : dorsalsphere H : hinge I : insula IX-Te = tergumIX LVL : lower vaginallip mm = millimeter ?GL = postgenitallobe Pr : proctiger SCa = spermathecalcapsule SCaP = spermathecalcapsule pore SE : spermathecaleminence SES : spermathecaleminence spicule UVL : uppervaginal lip UVS : uppervaginal sclerite VIII-S = sternum VIII VIII-Te = tergum VIII VT = ventral tuft 1-5-S = setae 1-5 of sternum VIII 28 Contributionsof the AmericanEntomological Institute 32 (5) Fig. 1

Ce

., • . AGDB

III -

ventral

SCoP

03

dorsa J

V III- Te

IX- Te Haemagogus (Oonopostegus) leucocelaenus Reinerr:Female genitalia of Aedini - genusHaemagogus 29 Fig. 2

DPGL

BMA

SES

ventral 0.3

SCaP

SCa

dorsal

i

•.....•. • •.•:•.. • ...,• ...... :/.,i •,

VIII-Te

IX-Te Haemagogus (Haem•gogus) splenclens 30 Contributionsof the AmericanEntomological Institute 32 (5)

SYSTEMATIC INDEX

Valid genericand specific taxa are italicized, synonyms and other taxa are in Romantype. Boldfacepage numbers are those which began the primarytreatment of the taxon. Figurenumbers are in parentheses.

acutisentis 24 Aedes 18, 20 Aedini 17, 20 albomaculatus 24 anastasionis 24 andinus 24 argyromeris 24 Ayurakitia 18, 20 baresi 24 boshelli 23, 24 capricornii 24 celeste 24 chalcospilans 24 clarki 22 Conopostegus 17, 18, 20, 21, 22 Culicidae 17 Diptera 17 equinus 24 18 Finlaya 20 Haemagogus 17, 18, 19, 20, 21, 22, 24 Heizmannia 18, 20 iridicolor 24 janthinomys 23, 24 leucocelaenus 21, 22, (1) [eucotaeniatus 21, 22 lucifer 24 Mattinglyia 20 mesodentatus 23, 24 18 panarchys 24 Psorophora 18, 20 regalis 24 reidi 20 soperi 23, 24 spegazzinii 24 splendens 23, 24, (2) Verrallina 18 Udclya 18 Zeugnomyia 18