Joumal of the American Control Association, l5(l):77-83, 1999 Copyright @ 1999 by the American Mosquito Control Association, Inc.

THE DORSAL APOTOME OF PUPAE AND FOURTH-INSTAR LARVAE OF CULICIDAE (DIPTERA), A STRUCTURE OF PHYLOGENETIC SIGNIFICANCE

JOHN R REINERT

Center for Medical, Agricultural and Veterinary Entomology, U.S. Department of Agriculture, Agriculture Research Service, 1600/1700 SW 23rd Drive, Gainesville, FL 32608

ABSTRACT, The development of the pupal dorsal apotome was examined and compared in representatives of the families Culicidae, , , and . Illustrations of the pupal dorsal apotome for 73 species, in 25 genera, are provided to show the differences and similarities between taxa. The 4th-instar larval dorsal apotome is compared and illustrated for the 3 subfamilies of Culicidae. The phylogenetic signifi- cance of the dorsal apotome of the pupae and 4th-instar larvae is discussed.

KEY WORDS Phylogenetics, dorsal apotome, mosquitoes, pupae, larvae

Based on a cladistic analysis of the anatomical most recent one (Culicidae) listed last. However, characteristics of larvae (22 features), pupae (2 Pawlowski et al. (1996), utilizing ribosomal RNA ". features), and adults (6 features), Wood and Bor- gene sequences, found . . the sister-group rela- kent (1989) included the 4 families Dixidae, Cor- tionship between Culicidae, Chaoboridae and Cor- ethrellidae, Chaoboridae, and Culicidae within the ethrellidae was clearly confirmed, the Dixidae, tra- infraorder , superfamily . ditionally considered as closely related to these Their listing of the families was with the phylo- two families, were not placed close to them on our genetically older one (Dixidae) listed first to the trees."

Table l. Taxa evaluated having pupal dorsal apotome with a single sclerite.r:

Family Chaoboridae Chaoborus sp. (S) Family Corethrellidae Corethre lla appendiculata Grabham (I) Corethrella longitubus Belkin, Heinemann, and Page (I) Corethrella brakeleyi (Coquillett) (S) Corethrella wlrrlui Stone (S) Corethrella librutta Belkin, Heinemann, and Page (I) Family Dixidae .scitula Belkin, Heinemann, and Page (I) Paradixa fuscinervi.s (Tonnoir) (I) Dixina .solomonlsBelkin (I) Paradixa fiarrisi (Tonnoir) (l) Mesodixa biambulacra Belkin, Heinemann, and Page (I) Paradira neoTelanaica (Tonnoir) (I) Nothodixa campbelli (Alexander) (I) Paradixa tonnoiri Belkin (I) Family Culicidae Subfamily Anophelinae An. (Ano.) atropos Dyar and Knab (S)x An. (Ker.) neivae Howard, Dyar, and Knab (I)* An. (Ano.) barberi Coquillett (S)* An. (Ker.) pholidotus Zavortink (I)* An. (Ano.) bradleyi King (S)* An. (Nys.) albimanus Wiedemann (S)* An. (Ano.) campestris Reid (I)* An. (Nys.) aquasalis Curry (I)* An. (Ano.) crucians s.l. Wiedemann (S)* An. (Nys.) nontestensis Galvao and Lane (I)* An. (Ano.) earlei Yargas (S)* An. (Ny.s.)oswaldoi (Peryassu) (I)x An. (Ano.) punctipennis Dyar and Knab (S)* An. (Nys.) srrzrzleiRoot (I)* An. (Ano.) quadrimaculatas s.s. Say (S)* An. (Ste.) ftonpi Edwards (S)* An. (Ano.) walkeri Theobald (S)* An. (Ste.) nimbus (Theobald) (S)* An. (Cel.) aconitu.s Doenitz (I)x Bi. (Bir.) gracilis Theobald (I)* An. (Cel.) culicifacies Giles (I)* Bi. (Bru.) hollandi Taylor (S)* An. (Cel.) minimus Theobald (I)* Bi. (Bru.) obscura Tenorio (I)* An. (Cel.) pampanai Buettiker and Beales (I)* Bi. (Nbi.) confustt Bonne-Wepster (I)* An. (.Cel.) varuna Iyengar (I)* Bi. (Nbi.) soesiloi (Strickland and Chowdhury) (I) An. (Ker.) bambusicolus Komp (I)* Ch. bathana (Dyar) (S)* An. (Ker.) bellator Dyar and Knab (I)* Ch. bonneae Root (S)* An. (Ker.) homunculus Komp (I)" Ch. fajardoi (Lutz) (S)E 'Abbreviations of genera and subgenera of Culicidae follow Reinen (1975, 1982, l99l). 'I, published illustration of pupal exuviae examined; S, specimen of pupal exuviae examined; x, condition of larval dorsal apotome confirmed on sDecimen and./or oublished illustration. 78 JounNx- op rxe AuenrceN Moseurro CoNrnor_ AssocrerroN Vor-. 15, No. I

Table 2. Taxa evaluated having pupal dorsal apotome with 2 sclerites.',2 Family Culicidae Subfamily Tribe Aedeomyiini (Ady.) Ad. catasticta Knab (I)x Ad. (Ady.) squamipennis (Lynch Anibalzaga) (S)* Tribe Aedini Ae. (Abr.) papago Zavortink (S)* Ae. (Muc.) quasiferinus Mattingly (S)* Ae. (Adm.) alboscutellatus (Theobald) (S)x Ae. (Neo.) lineatopennis (Ludlow) (S)* Ae. (Adm.) senyavinensis Knight and Hurlbut (S)* Ae. (Not.) chathamicus Dumbleton (I)* Ae. (Adm.) veran.r (Meigen) (S)* Ae. (Och.) canadensis (Theobald) (S)* Ae. (Aed.) cinereus Meigen (S)* Ae. (Och.) dupreei (Coquillett) (S)* Ae. (Aed.) esoensis Yamada (S)+ Ae. (Och.) inJirmatus Dyar and Knab (S)* Ae. (Ala.) brevitibia (Edwards) (S)x Ae. (Och.) mitchellae (Dyar) (S)* Ae. (Alb.) ngong van Someren (S)* Ae. (Och.) muelleri Dyar (S)x Ae. (Ayu.) peytoni Reinert (S)* Ae. (Och.) taeniorhynchus (Wiedemann) (S)* Ae. (Azt.) ramirezi Vargas and Downs (I)* Ae. (Och.) tormentor Dyar and Knab (S)* Ae. (Blk.) aurotaeniatus Edwards (I)* Ae. (Och.) uigilax (Skuse) (S)x Ae. (Bot.) helenae Reinert (S)* Ae. (Par.) ostentatio (Leicester) (S)* Ae. (Can.) indonesiae Mattingly (S)* Ae. (Par.) thailandensis Reinert (S)* Ae. (Can.) masculinus Mattingly (S)* Ae. (Pro.) triseriatus (Say) (S)* Ae. (Cha.) tulliae Taylor (I)+ Ae. (Rhi.) longirostris (Leicester) (S)* Ae. (Chr.) thomsoni (Theobald) (S)* Ae. (.Scu.) albolineatus (Theobald) (S)* Ae. (Dic.) franciscol Mattingly (S)x Ae. (Sku.) pembaensis Theobald (I)* Ae. (Dic.) whartoni Mattingly (S)* Ae- (StS.) aegypti (Linnaeus) (S)+ Ae. (Etlw.) imprimens (Walker) (S)* Ae. (StS.) albopictus (Skuse) (S)* Ae. (Fin.) albotaeniatus (Leicester) (S)* Ae. (Ver.) atrias Barraud (S)* Ae. (Fin.) chrysolineatus (Theobald) (S)* Ae. (Ver.) bzrleri Theobald (S)* Ae. (Fin.) cros.si Lein (S)* Ae. (Ver.) carmenti Edwards (S)* Ae. (Fin.) fusclpal2ls Belkin (S)* Ae. (Ver.) yzsal Barraud (S)* Ae. (Fin.) geniculatu.s (Olivier) (S)* Ar. (Arm.) alkatirii Toma, Miyagi, and Syafruddin (I)* Ae. (Fin.) leonls Colless (S)* Ar. (Arm.) barsasl Stone and Thurman (S)x Ae. (Fin.) longipalpis (Grunberg) (S)* Ar. (Arm.) subalbatus (Coquillett) (S)* Ae. (.Fin.) macfarlanei (Edwards) (S)* Ar. (l,ei.) magnus (Theobald) (S)* Ae. (Fin.) oreophilus (Edwards) (S)* Er. inornatus Newstead (S)* Ae. (Fin.) papuensis (Taylor) (S)* Er. Ieut'opous Graham tS.1* Ae. (Fin.) poicilius (Theobald) (g)x Er. penicillatus Edwards (S)* Ae. (Fin.) reinerti Rattanarithikul and Harrison (S)* Er. silvestris Ingram and De Meillon (S)* Ae. (Fin.) rogoi (Theobald) (S)* Hg. (Con.) leucocelaenus Dyar and Knab (I)* Ae. (Geo.) kabaensis Brug (S)* Hg. (Hag.) janthinomys Dyar (S)* Ae. (Gym.) mediovittatus (Coquillett) (S)x Hg. (Hag.) spegazzinii Brethes (S)* Ae. (Hal.) australis (Erickson) (I)* Hz. (.Hez..)complex (Theobald) (S)* Ae. (How.) walkeri (Theobald) (I)* Hz. (Hez..) reiuli Mattingly (S)+ Ae. (Hua.) wauensis Huang (S)x Hz. (Hez..) persimilis Mattingly (S)* Ae. (lso.) cavaticus Reinert (S)* Op. fuscus Hutton (S)* Ae. (Ken.) dissimilis (Leicester) (S)x Ps. (Gra.) columbiae (Dyar and Knab) (S)* Ae. (.Ken.) harbachi Reinert (S)+ Ps. (Jan.) cyanescens (Coquillett) (S)* Ae. (Kom.) purpureipes Aitken (S)* Ps. (.Jan.) ferox (Von Humboldt) (S)* Ae. (Lep.) aurimargo Edwards (S)* Ps. (Pso.) ciliata (Fabricius) (S)* Ae. (Lev.) geoskusea Amos (I)* Ps. (Pso.) howardii Coquillett (S)* Ae. (br.) aze.sii (Ludlow) (S)* Ud. argyrurus (Edwards) (S)* Ae. (Mac.) tremulus (Theobald) (S)* Ze. gracilis Leicester (S)* Ae. (Mol.) pecuniosus Edwards (S)* Ze. Iawtoni Baisas (S)* Ae. (Muc.) laniger (Wiedemann) (S)* Tribe Culicini Cx. (Acl.) belkini Stone and Penn (I)* Cx. (Ads.) amazonensis (Lutz) (I)* Cx. (Lut.) halifaxii Theobald (I)* Cx. (And.) conservator Dyar and Knab (I)* Cx. (Mca.) bisulcatus (Coquillett) (I)* Cx. (Bel.) eldridgei Admes md Galindo (I)* Cx. (Mel.) enaticus (Dyar and Knab) (S)* Cx. (Car.) iridenscens (Lutz) (I)* Cx. (Mel.) pilosus (Dyar and Knab) (S)x Cx. (Cui.) nigropunctatus Edwards (S)* Cx. (Ncx.) leonardi Belkin (I)* Cx. (Cux.) quinquefasciatus Say (S)* Cx. (Ncx.) territan.s Walker (S)* Cx. (Cux.) restuans Theobald (S)* Cx. (Tha.) dispectus Bram (I)* Cx. (Cux.) salinarius Coquillett (S)* Cx. (Tin.) latisquama (Coquiuett) (I)* Cx. (Cux.) tarsalis Coquillett (S)* De. cancer Theobald (S)* Cx. (Eum.) brevipalpis (Giles) (S)* De. pseudes Dyar and Knab (S)* Cx. (Lop.) peytoni Bram and Rattanarithikul (S)* Ga. leei Stone and Barreto (D+ Me.ncs 1999 ScrcNrrnc Norrs

'Iable 2. Continued.

Tribe Culisetini Cs. (All.) Iongiareolata (Macquart) (I)x Cs. (Cus.) incidens (Thomson) (I)* Cs. (CU.) melanura (Coquillett) (S)x Cs. (Cus.) inornata (Williston) (S)* Tribe Ficalbiini Fi. minima (Theobald) (S)* Mi. (lng.) milloti Grjebtne (l)* Fi. uniformis (Theobald) (I)* Mi. (Mim.) chamberlaini Ludlow (S)* ML (Ins.) bernardi (Doucet) (I)* Mi. (Mim.) gurneyi (Belkin) (I)* Mi. (Ing.) beytouti (Doucet) (I)* ML (Mim.) hispida (Theobald) (I)* Mi. (Ing.) grjebinei Brunhes (I)* ML (Mim.) splendens Theobald (I)+ Tribe Hodgesiini Ho. malayi Leicester (S)* Ho. solomonis Belkin (I)* Tiibe Mansoniini Cq. (Aus.) tenuipalpis (Edwards) (I) Ma. (Man.) titillans (Walker) (I)* Cq. (Coq.) latea (Belkin) (I)* Ma. (Man.) uniformis (Theobald) (S)* Cq. (Rhy.) nigricans (Coquillett) (I)* Ma. (Mnd.) bonneae Edwards (S)* Ma. (Man.) leberi Boreham (l)+ Tribe Orthopodomyiini Or. anopheloide^s (Giles) (I)+ Or. siamensis Zavortink (I)* Or. fascipes (Coquillett) (I)* Or. signifura (Coquillett) (S)* Or. phylloToa (Dyar and Knab) (I)* On wilsoni Macdonald (S)x Tribe Sabethini Jb. longipes (Fabricius) (I)x Tp. (Rac.)sronei Belkin (I)* Jb. ulopus (Dyar and Knab) (I)* Tp. (Rah.) coheni Belkin (l)* Li. durhamii Theobald (S)* Tp. (Tri.) ro.rasl Baisas and Ubaldo-Pagayon (I)* Mg. argyropus (Walker) (I)* Tp. (Trp.) purpuratus (Edwards) (I)* Ml. solomonis (Wharton) (I)* Tr. compressum Lutz (I)* Ru. (lso.) lunatum (Theobald) (I)x Tr. pallidiventer (Lutz) (I)* Ru. (Run.) fronrasrm (Theobald) (I)* Wy. (Cae.) fernandezyepezi Cova Garcia, Sutil Ru. \Run-l reversum Lane and Cerqueira (I)* Oramas, and Pulido F (I)* Sa. (Dav-) petrochiae (Shannon and del Ponte) (I)* Wy. (Den.) forcipenis Lourenco de Oliviera and da Sa. (Pey.) aurescens (Lutz) (I)* Dilva (I) Sa. (Sab.) gymnothorax Harbach and Petersen (I)* Wy. (Exm.) arborea Galindo, Carpenter, and Trapido Sa. (Sbn.) xhyphydes Harbach (I)* (I)* To. (Sua.) sparhulirostris Edwards (S)* Wy. (Wyo.) haynei Dodge (S)* To. (Top.) javaensrs Miyagi and Toma (I)* Wy. (Wyo.) mitchellii (Theobald) (S)* To. (Top.) lindsayi Thurman (I)* Wy. (Wyo.) vanduzeei Dyar and Knab (S)* To. (Top.) spathulirostris Edwards (S)* Wy. (Zin.) fishi Zavortink (I)* Tribe Uranotaeniini Ur. (Pfc.) anhydor Dyar (I)* Ur. (Ura.) lowll Theobald (S)* Ur. (Pfc.) bimaculata Leicester (I)* Ur. (Ura.) sapphirinia (Osten Sacken) (S)x Subfamily Toxorhynchitinae Tx. (Lyn.) portoricensis (Roeder) (I)* Tx. (Lyn.) rutilus (Coqulllett) (S)* Tx. (Tox.) brevipalpis Theobald (I)* Tx. (Tox.) amboinensis (Doleschall) (I)* Tx. (Tox.) splendens (Wiedemann) (I)*

'Abbreviations of genera md subgenera of Culicidae follow Reinert (1975, 1982, l99l). 'I, published illustration of pupal exuviae examined; S, specimen of pupal exuviae examined; *, condition of lrval dorsal apotome confimed on specimen and/or published illustration.

While conducting a morphologic taxonomic bounded laterally by the frontal ecdysial lines and study of the Quadrimaculatus Complex of the ge- apically by an imaginary line between the most api- nus Anopheles Meigen (Reinert et al. 1997),I noted cal parts of the frontal ecdysial lines . . . . In mos- and illustrated differences in the development of the quito pupae, perhaps homologous with the dorsal pupal dorsal apotome. Crawford (1938) previously apotome of mosquito larvae." In slide-mounted illustrated the dorsal apotome (as vertical plate) of specimens of pupal exuviae (positioned in a flat- 15 Malayan species of Anopheles and suggested tened aspect [Reinert et al. 1997]) the dorsal apo- "appeared that the shape of this structure to vary tome projects from the median anterior margin of to some extent specifically." Also, most authors the cephalothorax in a manner appropriate for com- who have illustrated the pupal cephalothorax have parative study. included the dorsal apotome, but without any writ- After noting differences in the structure of the ten mention or description of it. Harbach and dorsal apotome in the pupae of the Quadrimacula- Knight (1980) provided the following definition of tus Complex (5 sibling species), this structure was ". the dorsal apotome of pupae . . the cranial area compared to species in other mosquito genera and JounNel oR rne AvenrcaN Mosqurro CoNrnol AssocrlrroN Vol. 15. No

T I I /-l /-\ 0,3 (/ a-) - \./ /t i\ It al \/ I t--J /--J U'\J V..r/ 1 2 5 6 7 nn /lh m\ / l\ \ //l\ o al..\r1rl [l (tU \^/ LJ \-rV \17 u, UU 8 9 10 11 12 13 14 /^n ( /\ l \tt)tf\ r-x1 nn rT) ff) t/\)\ /\ ) /-lf-\ \/\I w VYVUVV VY VV UTJ 15 16 17 18 19 20 21 an aY^\ [lD fl|) \il | \ /\ I nr).A) ffi u1/ U\] VV VV \-l u u/ 22 23 24 25 26 27 28 m\ A rn rln ,n \ /\ I t/\)/.n t/\l 1 i\ \ I /\ | \/\t o0Ll u U\] t/v L/V u\l VV 30 31 32 33 34 35 ax-) no(A),-af-\ CN \ /\ | |) \ t) I vt/ \t/ \l v \-/v OD37 38 39 41 42

fiD43 QD45 Fig. l. Dorsal apotome (DAp) of pupae; drawn from pupal exuviae; all DAps drawn to same scale,0.3 mm. 1, Corethrella brakeleyi, d, Florida, USA; 2, Corethrella wirthi, 6, Florida, USA; 3, Bironella hollandi, d, Solomon Islands; 4, Anopheles quadrimaculatas s.s., d, Minnesota, USA; 5, An. atropos, d, Florida, USA; 6, An. earlei, 6, Wisconsin, USA;7, An. barberi, d, Florida, USA;8, An. bradleyi, d, Louisiana, USA;9, An. punctipennis, 6, Wisconsin, USA; 10, An. crucians s.1., d, Louisiana, USA; ll,An. albimanus,9, Florida, USA; 12, Orthopodomyia sign{era, d, Florida, USA; 13, Limatus durfuimii, 9, Costa Rica; 14, Wyeomyia vanduzeei, d, Florida, USA; 15, Aedes aurimargo, d, Papua New Guinea; 16, Ae. wauensls, 9, Papua New Guinea; 17, Ae. tremulzs, d, Australia; Ml.ncH 1999 ScrBNrrrrcNores 81 major differences were observed that seem to be of by a very narrow U-shaped band and located im- phylogenetic signiflcance. In specimens and/or pub- mediately ventral to the spiracular apparatus, Nut- lished illustrations (e.g., Knight and Chamberlain tal and Shipley's organ is present dorsally on the 1948, Belkin 1962, Belkin et al. 1970, Zavortink prothorax, a tergal plate is situated anteromesally 1972; publications of the Mosquitoes of Middle on each of abdominal segments I-VII, and pal- America Project, Southeast Asian Mosquito Pro- mate setae are present (except 10 of 12 species of ject, Medical Entomology Project, Walter Reed the Umbrosus Group of Anopheles); and the eggs Biosystematics Unit, and others) the dorsal apoto- have floats (except absent in a few species, for me appears as a single more or less oblong sclerite example, An. barianensls James, An. dthali Patton, in species of the subfamily Anophelinae (genera An. multicolor Cambouliu, An. pessoai Golvao Anopheles, Bironella Theobald, and Chagasia and Lane : An. braziliensis (Chagas), An. sawyeri Cruz) and as a structure consisting of 2 longitudinal Causey, Deane, Deane and Sampaio, An. super- sclerites connected by a membrane mesally in the pictus Grassi, and An. turkhudi Liston; see Chris- subfamilies Culicinae (i.e., tribes Uranotaeniini, tophers 1933, Causey et al. 1944). In comparison Culicini, Aedeomyiini, Hodgesiini, Culisetini, Fi- the 4th-instar larvae of Culicinae and Toxorhyn- calbiini, Mansoniini, Orthopodomyiini, Aedini, and chitinae have the siphon developed as an elongate Sabethini) and Toxorhynchitinae (Knight and Stone sclerotized tube with the spiracular apparatus sit- 1977). Tables I and 2 list the broad range of taxa uated apically, Nuttal and Shipley's organ is ab- evaluated during this study (4 families, 3 subfami- sent, tergal plates are absent on segments I-V (ter- lies, l0 tibes, 42 genera, 102 subgenera, and 233 gal plate present on one or more of segments VI- species). VIII in Orthopodomyia), and palmate setae are ab- The pupal dorsal apotome in species of Dixidae, sent; and the eggs are without floats. Larvae ofthe Corethrellidae, and Chaoboridae is represented as subfamily Toxorhynchitinae (Fig. 2C) have seta 8- a single sclerite, suggesting that this is the plesio- C displaced far cephalad on the dorsal apotome, morphic state and the divided condition is the apo- situated between setae 6-C and 7-C, and more or morphic state. Therefore, the development of the less in a line with setae 5-7-C, and antennal setae pupal dorsal apotome supports the contention of 2,3-A are inserted basad of seta l-A, which dis- most authors that the Anophelinae is the most tinctively separate them from larvae of the other primitive subfamily of the Culicidae. From the ex- 2 subfamilies of the Culicidae. In larvae of Ano- amination of published specimens and/or illustra- phelinae (Fig. 2A) and Culicinae (Fig. 2B) seta 8- tions it seems that the shape and development of C is inserted noticeably caudad to seta 7-C on the the dorsal apotome of pupae also may be of value dorsal apotome and is not located in line with se- at other taxonomic (e.g., gen- levels separation of tae 5-7-C, and antennal setae 2,3-A are inserted era, subgenera, species groups, and some species). distal to seta 1-A. Terminology used follows Har- The dorsal apotomes of various pupae of families bach and Knight (1980). (2)" subfamilies (3), genera (25), subgenera (46), and species (73) are illustrated (Figs. 1 and 2) to show the differences or similarities observed be- ACKNOWLEDGMENTS tween taxa. The posterior portion of the dorsal apotome is Special acknowledgment and appreciation is ex- nirrrow in 4th-instar larvae of species of Anophel- pressed to: Ralph E. Harbach (The Natural History inae (Fig. 2A) and broad in the Culicinae (Fig. 2B) Museum, London, United Kingdom) for examining and Toxorhynchitinae (Fig. 2C) (some Uranotaen- the pupal dorsal apotome in several species and for ia Lynch Arrlbalzaga?). As pointed out by others, reviewing the manuscript; and to Bruce A. Harrison the Anophelinae also is distinguished as follows: (North Carolina Department of Environment and in the 4th-instar larvae the siphon (see discussion Natural Resources, Winston-Salem, NC) and E. L in Belkin 1951,1962; Harbach 1978) consists only Peyton (Walter Reed Biosystematics Unit, Smith- of a pair of sclerotized pecten plates, one on each sonian Institution, Washington, DC) for reviewing side dorsolaterally, that are connected posteriorly the manuscript. (- 82 JounNlr- or rgr ArventcANMoseuno CoNTRoLAssocteuoN VoL. 15, No. I nn r/1) / l\ \ /\ 03T n11 ffl 1/l/ \il ) fl-) \/\t I v\.,/ \ly U\/ \JV UU t/v 47 48 49 so 51 52

/\ /\ nf\ /)fl t /\ I tt\l rYl .lt) VV ur/ \]U I7 VV s3 v)/54 55 56 58 57 /N /\ /\ /ln VY UU d\) VV .tnVV 59 ffi 60 61 62 63 64 /.n\ nn IA\/-----).'(----\ |\ ) [K) VV \]V VV 65 66 CDCD 69 70

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Fig. 2. Dorsal apotome (DAp) of pupae; drawn from pupal exuviae; all DAps drawn to same scale, 0.3 mm. 47, Culiseta melanura, d, Florida, USA;48, Cs. inornata, 9, Florida, USA;49, Aetleomyia squamipennis, g, panama; 5O, , I, Malaysia; 51, Ficatbia minima, d, Malaysia; 52, Topomyia spithulirostris, g, Ma- laysia; 53, Culex brevipalpi^1 I, Malaysia:54, Cx. nigropunctatus, 9, Malaysia; 55, Cx. quinquefasciatus, g, Florida, USA;56, Cx. salinarius, d, Florida, USA; 57, Cx. restuans, d, Florida, USA;58, Cx. eriaticis, d, Florida, USA:59. Cx. territans, 9, Florida, USA; 60, Deinocerites cancer, 6, Florida, USA; 61, Hodgesia matayi, ?, ;62, Uranotaenia lowii, 9 , Florida, USA; 63, IJr. sapphirina, d, Florida, USA; 64, Ae. butleii, d, Malaysia; 65, Ae. esoensts, d, Japan; 66, Ae. dissimilis, 9, Malaysia;67, Armigeres magnus, d, Thailand; 68, Eretmapoditis silvestris, d, South Africa;69, Psorophoracolumbiae, d, Florida, USA;70, Ps. cyanescens, d, Florida, USA;71, Ps. ciliata, d, Florida, USA; 72, Toxorhynchites rutilus, 6 , Florida, USA; 73, Opifux fuscus, d , New Zealand. Cranium of Culicidae showing dorsal apotome (DAp) and antenna (A); drawn from larvae; not drawn to scale. A,Anopheles crucians s.l.;B, Cutex q uinquefa sc iat us ; C, Toxo rhync hit es rut ilus.