Memórias do Instituto Oswaldo Cruz Fundação Oswaldo Cruz, Fiocruz ISSN: 1678-8060 EISSN: 1678-8060 Vol. 90, Num. 3, 1995, pp. 375-385

Wyeomyia luteoventrafis Theobald, the Type Species of the Subgenus Dendrornyia Theobald (Diptera: Culicidae)

Monitlue Albuquerque Motta, Ricardo Lourenco-de-Oliveira

Laboratorio de Transmissores de Hematozoanos, Instituto Oswaldo Cruz, Av. Brasil 4365, 21045-900 Rio de Janeiro, RJ, Brasil

Code Number: OC95074
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Key words: Dendromyia - Wyeomyia luteoventralis - Wyeotnyia ypsipola - mosquito taxonomy

Wyeomyia luteoventralis was described by Theobald (1901), from three females collected in Para (-Belem, State of Pars, see Belkin et al. 1971), Brazil. Only one female is remaining in the collection of The Natural History Museum (NHM), marked as type by Theobald and selected as lectotype by Belkin (1968). Wyeomyia luteoventralis is the type species of Dendromyia Theobald (1903) by subsequent designation (Blanchard 1905). Nearly 50 species of Wyeomyia have been grouped in the Neotropical subgenus Dendromyia, under an obviously unnatural classification. Most species presently regarded as belonging to Dendromyia were described superficially from only a few specimens and, in general, their immature stages and the morphological characters of both males and females are poorly known or unknown. Because of this, the subgenus certainly contains species whose phylogenetie relationships are unknown. Even the type species is poorly known: its larva and pupa, behavior and geographical distribution have never been described. In order to fix the concept of the subgenus Dendromyia we are redescribing Wy. luteoventralis and studying the related species.

Wyeomyia (Dendrotnyia) luteoventralis Theobald luteoventralis Theobald, 1901: 348. 'Lectotype female (designated by Belkin, 1968): Belem, Para, Brazil (NHM). Belkin et al. 1971 (type locality restriction). quasiluteoventralis Theobald 1903:317 (Den- dromyia).

Dendromyia luteoventralis of Theobald, 1903:318.

FEMALE - Head: vertex covered with darkish broad scales with blue and green reflections; a spot of white scales on median area of vertex and occiput; this spot generally triangular in shape, but sometimes resembles an irregular longitudinal stripe; side of eyes and postgena with white broad scales; ocular line with white scales and many darkish setae; two dark, strong and long interocular setae. Proboscis covered by darkish brown scales, length 0.8-0.9 of forefemur, expanded distally; basal labial setae long, brown. Maxillary palpus same color as proboscis, about l/7 proboscis length. Clypeus ovate, blackish, pruinose, nude. Antenna: pedicel brownish to blackish, pruinose, with some bronzy scales. Flagellum slightly plumose, a little shorter than pro- boscis, with 13 flagellomeres. Thorax: integument pale brown. Antepronotum largely covered with darkish scales with bluish reflections, sometimes with few white scales ventrally; setae dark. Postpronotum densely covered with whitish scales. Scutal integument pale brown, much darker than pleura, covered with darkish broad scales with blue and green reflections (same as antepronotal scales), except for a group of few white scales on anterior promontory; median anterior promontory and median scutal fossal setae dark, numerous; supraalar setae long, strong, dark. Scutellum darkish-scaled, same color as scutum; 3 or 4 developed setae on each lobe. Mesopostnotum pale brown, with about 10 (8-14) slightly dark setae of different sizes; the longest and turned upward mesopostnotal setae reach or cross the median scutellar setae. Pleural sclerites with very pale, yellowish integument, densely covered with whitish scales; pleural setae pale, mostly yellowish, or light tan: 3 or 4 tan, prespiracular setae; 5(3-7) very pale, yellowish, elongate lower mesokateepisternal setae inserted below and above upper margin of mesomeron; 4 or 5 tan upper mesokatepisternal setae; 9 (9-11) very pale, yellowish upper mesepimeral setae. Wing: upper calypter without seta. Scales of veins brown, broad. Rs, R2+3, R2, R3, MI*2, MI and M2 with both lateral anterior and appressed scales broad and spatulate; M with broad appressed scales; Cu with narrow, long spatulate scales; IA with spatulate scales. Distal third of all veins with broad appressed scales dorsally. Halter: scabellum yellowish; pedicel yellowish, dark scaled on one side; capitellum with dark scales. Legs: coxae and trochanters with very pale integument, partially covered by whitish scales, setae pale. Femora, tibiae and tarsi largely dark-scaled. Femora and mid- and hind tibiae with a stripe of white scales ventrally. Foretibia with a tenuous and incomplete stripe of white scales ventrally, very indistinct distally. Mid- and hind tarsomeres I with a few white scales ventrally. Ungues simple. Abdomen: densely covered with brownish broad scales with blue and green reflections above, whitish scaled bellow, the colors separated on sides in nearly straight line. Terga I and VIII with numerous long brownish setae. Genitalia (Fig. IA-C): tergum IX spiculose with a median lobe bearing 8 setae; postgenital lobe short, evenly rounded, spiculose, with a median triangular area densely covered with minute setae arising from conspicuous alveoli; cercus digiti- fonn, spiculose, with elongate setae apically.

MALE - Similar to female. The white spot on vertex and occiput generally resembles an irregular longitudinal stripe. Antenna slightly more plumose than in female, with 14 flagellomeres; pedicel darkish brown, pruinose, with small bronzy scales. Proboscis 0.8 - 0.9 of forefemur length. Maxillary palpus darkish, 0.15 proboscis length. Legs similar to female, except naidtarsomeres II-V white-scaled. Genitalia (Figs ID, E and 2): tergum VIII covered with broad scales and minute setae, strong and long setae distally. Tergum IX with broad interlobar space; each lobe bearing 2 stout, strongly sclerotized setae with round apices, inner seta stronger than outer. Paraproct sclerotized at apex, with a single strong tooth or sometimes with a less developed second one; 3 or 4 small setae. Gonocoxite short; inner surface covered with minute setae; stemal surface with numerous setae; tergal surface with minute setae and 3 long, strong tergomesal setae. Basal mesal lobe slightly sclerotized, with a row of about 15 small setae and 2 prominent distal setae. Gonostylus short, essentially without stem, divided into 4 lobes: one narrow, long, laterally bent lobe, fringed at apex (homologous in part with lobe C[?] of Belkin et al. 1970); a slightly flat, nude, bifurcated lobe, broadened to apex and forming a gutter on mesal side (homologous with lobe C[?]); a cylindrical lobe (lobe M), acornshaped at apex, with numerous short, spiniform, and longer setae mainly on distal mesal and tergal sides; a digitiform slender tergal lobe (probably homologous with lobe A,E), arising from the cylindrical lobe, with about 10 setae, most at apex. Aedeagus roughly round in tergal view; apical tergal arms joined and forming a rather broad apical tergal bridge, with a small process on each side of apex; submedian tergal arms not joined; median sternal plate simple, membranous, rugose laterally.

PUPA (Fig. 3) - Number of branches of setae in Table I. Cephalothorax: pale; seta 5-CT nearly as long as I-CT, with 3-5 (3) aciculate branches. Trumpet: slightly and evenly tanned; short, cylindrical; index about 4 (3.7 - 5.6) (width measured at midlength). Abdomen: pale; anterior margin of stema darker. Seta 1-I strongly developed; 2-11VII very near posterior margin of tergum, 2-11 between and nearly in line with setae 1 and 3, 2-III mesad of seta 1 which in turn is mesad of seta 3, 2-IV-VII far mesad of seta 1; 6-11 usually single, long; (FIll-VII short, weak; 3-I slightly aciculate, 3-IV well anterior to seta 1, 3-V-VI/only slightly anterior to seta 1; 5-IV-VI long, single, about 1.5 length of following tergum, slightly aciculate; 9VII,VIII aciculate, 9-VII strongly developed, about length of paddle; 9-VIII considerably longer than paddle. Genital lobe: moderately tanned; same length or slightly longer than segment VIII in male. Paddle: pale, long, narrowed and strongly produced beyond midlength, spiculose at margins; spicules better developed distally; inner margin only slightly curved compared to outer margin.

LARVA (Figs 4, 5) - Number of branches of setae in Table II. Head: wider than long, pale. Maxilla (Fig. 5C, D): elongate, with a prominent apical tooth (AT); dorsomesal surface normally with 8 teeth (laciniarastrum, LR), progressively longer apicaIly; dorsal surface with numerous long and moderatIy long setae on mesal and apicoIateraI margins; maxillary brush represented by 1 or 2 more strongly developed (stouter) setae bome dorsolateraly at base of AT. Setae 2-4,6Mx single; seta l-Mx spiniform, bifid at tip. Mandible as figured (Fig. 5 A, B); 3 or 4 developed mandibular sweeper setae in 2 groups. Dorsomentum generally with 8 teeth on each side of slightly more prominent median tooth. Hypos- tomal suture complete; occipital foramen with dorsolateral slitlike extension on either side, margins heavily pigmented laterally. Seta I-C stout, curved; 4-(FC single; 7-C single or double, all simple; 11,14-C long, aciculate, 14-C posterior to 15-C. Antenna: short, cylindrical: Setae single; IA very close to apex. Thorax: integument smooth. Setae 4,7-P and 7,13-T on individual pigmented basal plates; plural groups 9-12-P,M,T on common basal plates; 1 I-P,M,T spiniform.

Fig. 1: Wyeomyia (Dendromyia) luteoventralis A-C female genitalia. A: tergum IX. B: cerci. C: postgenital lobe. D-E male genitalia. D: sternum VIII. E: tergum VIII.

TAXONOMY- The three females used by Theobald (1901) for the original description of Wy. luteoventralts were collected by Dr Herbert Edward Durham during a yellow fever expedition to Para, Brazil, in 1900-1 (Durham 1902). These specimeus were included with about 90 Brazilian mosquitoes taken with Durham to Theobald, in London, in August, 1901 (Theobald 1901:34950). The original specimens of Wy. luteoventralts were definitely collected in Bel6m (the capital of the State of Para, in the Brazilian Amazon), sometimes erroneously confused with the name of the State, and called "Para" by Durham (1902:46,49,58). We have read all available literature about the mosquito collections performed by Durham in Brazil as well as all correspondence between him and Theobald presently in the NHM library. We are convinced that the type specimens of Wy. luteoventralis were caught on human bait by Durham at some of the sites he visited within the city or in the surrounding forests of Belem (Durham 1902: 5'1,52,55,57; Theobald 1901: 346). This indicated that Wy. luteoventraits is somewhat anthropophilic, so we performed some captures on human bait in forests in Belem and collected females indistinguishable from the lectotype specimen. We were surprised to learn that these specimens were different from those described as Wy. luteoventralis by Lane and Cerqueira (1942: 590).

The original description of Wy. luteoventralis by Theobald (1901) was very superficial, and could not be used to identify a single mosquito species. Because of this, Wy. luteoventralts has rarely been reported in biological, ecological and taxonomic studies carried out on South American mosquitoes. Dyar (1924), discussing the identity of Wy. luteoventralts, stated that "No species with these characters [referring to the broad scales on the wing veins] is known to me, and it may be that luteoventralis remains to be rediscovered. Until this is done and the male made known, the proper application of Dendromyia perhaps cannot be made".

TABLE I

Ranges of numbers of branches for setae of pupa of Wyeomyia Luteoventralis. Mode in parenthesis

------------------------------------------------------------
                                   Abdominal segments
Seta  Cephalothorax               I        II     III
------------------------------------------------------------
0       -                         -         -       -
1      1-3(2)                     d        1,2(1)   1-5(4) 
2      1-3(2)                     1,2(1)   1        1 
3      2,3(2)                     1,3(1)   1,2(1)   1-3(1) 
4      24(2)                      3-7(5)   2-5(2,4) 14(2) 
5      3-5(3)                     1,2(1)   1-3(2)   14(1) 
6      1,2(1)                     1        1,2(1)   14(2) 
7      1-3(3)                     24(3)    24(2)    2-5(3) 
8      1,2(1)                     -        -        2-5(4) 
9      1-3(2)                     1        1        1 
10     1                          -        1-3(2)   1,2(2) 
11     1                          -        1,2(1)   1,2(1)
12     1-3(2)                     -        -        -
14     -                          -        -        -
-----------------------------------------------------------
d-dendritic
-----------------------------------------------------------
Seta               IV      V       VI       VII    VIII 
----------------------------------------------------------- 
0                  -       -        -        -       -
1                 1-3    1-3(2)   14(2)    1,2(1)    - 
2                 1      1,2(1)   1        1         -
3                 24(3)  1-4(3)   1-3(2)   1-3(1)    - 
4                 1-3(2) 24(4)    1-3(2)   1         1,2(1) 
5                 1      1        1        14(1)     -
6                 14(2)  1-4(2)   1-5(2)   1,2(1)    - 
7                 24(2)  2-6(4)   1-3(2)   1-3(1)    -
8                 24(4)  1-5(3)   2-5(3)   4-7(6)    -    
9                 1      1        1        12-21     16-28 
1                 01,2(2)1-3(2)   1,2(2)   1,2(1)    -  
1                 11-3(1)1,2(1)   1-3(1)   14(1)     -
12                -      -        -         -        - 
14                -      -        -         -        1
-------------------------------------------------------------

Fig. 2: Wyeomyia (Dendromyia) luteoventralis. Male genitalia. A: gonocoxopodite (mesal aspect). B: gonostylus (lateral aspect of lobes AE, C and M). C: aedeagus (stemal aspect). D: tergum IX. E: paraproct (lateral aspect).

Lane and Cerqueira (1942) described the female and male of a species identified as Wy. luteoventralis from reared specimens collected as larvae in the axils of Montrichardia arborescens (aninga plant) on Marajo Island (Curralinho and Canaticu river), Santarem and S. Isabel, State of Para, but outside the type locality (Belem). We compared the lectotype of Wy. luteoventralis with the specimens used by Lane and Cerqueira, as well as numerous examples identical to those described by these authors, which we recently collected in aninga plants in Belem, and noted that they are unmistakably different.

Fig. 3: Wyeomyia (Dendromyia) luteoventralis. Pupa. A: cephalothorax. B: metanotum and abdomen.

Fig. 4: Wyeomyia (Dendromyia) luteoventralis. Larva. CS: comb scales.

Fig. 5: Wyeomyia (Dendromyia) luteoventralis. Larval mouthparis. A-B: mandible. C-D: maxilla (A and C: ventral; B and D: domal views). E: domomenlum.

Wyeomyia ypsipola Dyar is the species that appears to be most closely related to Wy. luteoventralis. Although the female, male and male genitalia are distinct from those of Wy. luteoventralis, the fourth-instar larva and the pupa are practically identical. The adult of Wy. ypsipola has been described and illustrated previously (Dyar 1922ab, Lane & Cerqueira 1942, Lane 1953, Bruijning 1959). The female of Wy. ypsipola has white scaling on midtarsomeres II-V, the male has a continuous stripe of white scales on one side of midtarsomeres I-V (tarsomeres IV,V may be whitish on both sides), the male genitalia are distinct and very different from those of Wy. luteoventralis. The larva of Wy. ypsipola has the maxilla developed like that of Wy. luteoventralis (Harbach & Peyton 1993). It differs from Wy. luteoventralis in having seta 4-P generally with .more than 10 branches (1014[13]), 4-M 2,3(3) branched, 13-T 12-18 branched, 3,4,5-11 3-5, 1,2 and double, respectively, 6-111 double, 6-IV 3-5(4), 8-V 3,4, 4-VIII single and 5-VIII triple. The pupa of Wy. ypsipola has seta 7-V generally triple (1-4), 1 I-VII 2-4(3) branched and 9- VIII with 24-30 branches.

TABLE II

Ranges of branches for setae of the fourth-instar Larva of Wyeomyia Luteoventralis. Mode in parenthesis

-------------------------------------------------------------
                  Toracic segments       Abdominal segments
seta Head Antena   P         M       T       I      II     III 
-------------------------------------------------------------
0   1      -       6-12(10)  -       -       -      -      - 
1   1      1       1        1,2(2)  14(2)  1-3(3)  24(3)   2
2   -      1       1,2(1)   1,2(1)  1-3(2)   1      1      1
3   1      1       1,2(1)   1,2(1)  2-5(5) 3,4(3)  2,3(3)  2
4   1      1       6-14(8)  1-3(2)  1-4(2) 4-7(6)  3-6     1
5   1      1       1,2(1)   1       1    1,2(2) 1-3(2) 2,3(2)
6   1      -       1,2    1,2(1) 24(3) 7-11(11)8-12(11)1-3(2)
7   1,2    -       7-14(11) 24(3)9-14(9)5-8(8) 4-8(5)  4-7(7)
8   1-3(2) -       4-8(5)   4-7(5)3-5(4) -     1,2(1)  1-3 
9   4-6(4)  -      6-10(8)  11-13(11)  1     1      1       1
10  1-4(3) -       1-3(2)   1      1,2(1) 2,3(3) 2,3(2) 2,3(2)
11  3-5(3) -       1        1          1  3-5(4) 24(3)  24(3)
12  2-5(3) -       1,2(1)   1          1   -     3-6(4)     1
13  2,3,5(2)       -        4-7(6) 9-14(13)4-6(4)1,2(1) 2,3(2)
14  2,3(2) -       24(2)    4-6(5)     -    -      -      -
15  3-6(3) -       -         -         -    -      -      -
--------------------------------------------------------------
--------------------------------------------------------------
                      Abdominal segments

           IV       V        VI       VII       VIII      X
--------------------------------------------------------------
0         -        -        -         -         -        -    
1        2,3(2)  2-6(4)   3-5(4)   3,4(4)    4-6(5)    2,3(2) 
2         1        1        1         1         1      5-7(6) 
3        2-4(3)    1      1-3      1,2(1)    4-6       3,4(4) 
4         1      5-7(5)   2,3(2)      1      1-3(2)    4-6(6)
5        2,3(2)  2,3(2)   2,3(2)   2-5(4)    1,2(1)      -
6        2-4(2)  2-4(2)   2-4(3)    5-8          -       -
7        3-5(4)  4-7(5)   4-6(6)    3-5(3)       -       -
8        2-2(3)  1-3(2)   3-6(6)    5-9(6)       -       - 
9        1       1        1         24(2)        -       -
10       2       3-5(4)   5,6(5)    2,3(3)       -       -
11       2,3(3)  2,3(2)   2-4(4)    2-4(4)       -       -
12       1       1        3-5(3)    4-6(4)       -       -
13       2,3(2)  2,3(2)   3-5(5)    2,3(2)       -       -
14         -       -        -         -          -       -
15         -       -        -         -          -       -
-------------------------------------------------------------- 

Wyeomyia trifurcata Clastrier is very similar to both Wy. luteoventralis and Wy. testei in fe- male, larval and pupal characters, but has distinct male genitalia. According to the original decription (Clastrier 1973), the larva of Wy. trifurcata has pecten scales restricted to the distal third of siphon and seta 2a-S composed of 2 laceolate and one hairlike seta; the pupa has a short paddle and seta 9-VIII with strongly developed branches.

Based on morphological characters of the adults, male genitalia (as redescribed by Lane 1953) and larva (EL Peyton, personal communication), Wy. complosa (Dyar) and Wy. jocosa seems to belong to the same group of Wy. luteoventralis.

Wyeomyia luteoventralis, Wy. ypsipola, Wy. testei, Wy. trifurcata, Wy. complosa and Wy. jocosa apparently form a monophyletic group, the subgenus Dendromyia. We hereby recognize and restrict this subgenus to include only these five species, because of their affinities in the adult, larval and pupal stages. These species have adults generally with white scales on the ocular line and scales forming a spot on the vertex and occiput; wing veins densely covered with broad spatulate scales; mesopostnotal setae long and numerous; antepronotum dark-scaled, rarely with a few whitish scales ventrally; and the abdomen with dark dorsal and pale lateral scallig s,eparated in almost a straight line. The larvae have a prominent apical tooth on the maxilla; seta 14-C developed, posterior to 15-C; seta I-P placed far mesad or caudomesad of 2,3-P; a short siphon with a single midventral row of pecten spines, and seta 2a-S chiefly or completely composed of dark lanceolate setae. The paddles of the pupae are elongate and densely spiculose on the margins.

Since the type species of Dendromyia is now properly defined, the correct interpretation and application of the subgeneric name is hereby established. The many unrelated Wyeomyia species previously included in Dendromyia are hereby excluded from the subgenus, and are retained in Wyeomyia without subgeneric placement.

We have been collecting and studying all life stages of those Wyeomyia species now excluded from the subgenus, with a view of understanding their relationships and establishing a more natural system of classification. The subgenus Dendromyia as now defined seems to be related to the subgenus Caenomyiella Harbach and Peyton and the Cleobonnea and Prosopolepis Series of Lane and Cerqueira (1942). The non currently recognized subgenus Shropshirea Dyar is now the only junior synonym of Dendromyia.

Bionomics: Wyeomyia luteoventralis may be collected on human bait in forest during the daytime. Many more females of this species were caught during our survey at the type locality, Belem, than those included in the list of material examined. Several blood fed wild-caught females died without laying eggs, even when traumatized by removing one wing. These specimens were damaged and were excluded from the morphological analysis. Larvae and pupae of Wy. lute- oventralis were found only in the axils of Heliconia sp. and Calathea sp. The eggs are curiously elongate and shaped like grains of rice.

Distribution: Wyeomyia luteoventralis is known only from the type locality, Belem, State of Para, Brazil.

Material examined: Wyeomyia luteoventralis: Lectotype female, Para, Brazil, H. Durham, 1901, NHM, London; 1 male Le Pe G and 1 female Le Pc, Estrada do Mosqueiro, DER (1 15'S 48 13'W), near Paricatuba River, Belem, Para, Brazil, XI-1992, reared from larvae collected in Heliconia sp., respectively deposited at NHM and Instituto Oswaldo Cruz (IOC), Rio de Janeiro, det. MA Motta and R Lourenqo-deOliveira, 1993; col. MA Motta and OV Silva; 5 females Le Pc, same data as above; 1 male Le Pe G, same data as above except 28-111-1993; reared from larvae collected in Calathea sp.; 3 females Le Pe, 1 female Le Pe 6, same data as preceding male, collected in Heliconia sp. and Calathea sp.; 6 females and 1 female G, same data as above, except 1-VII-1992, caught on human bait, col. RNL Lacerda.

Wyeomyia ypsipola: 1 male Le Pe G, Cumutu, Trinidad, 20-VII-1942-4, from Heliconia sp. flower, n. 3938, slide 1078, Faculdade de Saude P6blica (FSP), Sao Paulo, der. J Lane; col. DG Dow; 2 females Le Pe G and 1 female Le Pc, Estrada do Mosqueiro, DER, Belem, Para, Brazil, XI-1992, from Heliconia sp. IOC, MA Motta and R Lourenco-de-Oliveira der.; col. MA Motta and OV Silva; 1 male 12 Pe G, same data as above except 111-1993, col. OV Silva, from Calathea sp.; 1 male 12 Pe G, 1 female 12 Pc, same date as above except from Heliconia sp.; 2 males Pe G, 1 male G, 1 female Pe and 1 female Le Pc, Machadinho (MA-32), Rondbnia, Brazil,. III1987, from Aracea, IOC, der. MA Motta, MG Castro and R Lourenco- de-Oliveira, 1989, col. R Lourenco-de-Oliveira; 2 females 12 Pc, Estrada Velha da Mibrasa, Itapoa do Oeste, Rondbnia, Brazil, IV- 1987, other data as above; 2 females Pe Vila Marechal Rondon. Ariouemes, Rondonia, VII-1987, other data as above; 1 male and 1 female Le Pe, Labelle Park, Vinhais, Sao Luiz, Maranhao, Brazil, VII- 1994 from Heliconia sp. IOC, det. Ma Motta col. R Lorenco-deOliveira; 2 females Pe, 1 male Pe G and 3 males Le Pe, from Dieffenbachia picta (?), other data as above.

Wyeomyia sp. (misidentified as Wy. luteoventralis by Lane and Cerqueira 1942): Brazil: 1 female, Curralinho, Para, n. 2539, FSP, der. J Lane and N Cerqueira, 1940; 1 female, Sao Paulo, NILM, det. J Lane and N C, erqueira, 1940; F Lane col.; 2 males G, Curralinho, Para, XI-1935, one n. 9800 and another without number; slides n. 2292, 2294; from larvae collected in aninga plant (Montrichardia arborescens), deposited at Centro de pesquisas Rene Rachou (CPRR), Belo Horizonte, der. N Cerqueira; col. Servio de Fe bre Amarela; 2 females and 2 males, same data as above (females numbered 9800); 1 male (23, Muri queira, Bahia, n. 2538, FSP, der. Lane and Cer queira, 1940 (Wy. howardi Lane & Cerqueira 1942). 10 males 12 Pe G, 15 males Le Pc, 30 fe males Le Pc, from 13 individual rearings, Estrada do Mosqueiro, DER, Belem, Para, Brazil, X1 1992, 10C, der. MA Motta; col. MA Motta and OV Silva. 12 males 12 Pe G, 12 females Le Pc, Catu, Utinga, Belem, Para, Brazil, XI-91, 10C, der. MA Motta, 1993, col. R Lourenco-de- Oliveira and MG Castro.

ACKNOWLEDGEMENTS

To Dr Ralph E Harbach (NIlM) for his suggestions and review of the manuscript; to Drs Amelia PA Travassos da Rosa and Nicola' Degalier and all techificians from Instituto Evandro Chagas - Arbovirus research group, particularly to Orlando V Silva as well as to Dr Amiraldo Pinheiro from Fundacao Nacional de Saude, for the help in the field work in Belem; to EL Peyton (Waiter Reed Amy Inst. Research) for examining larvae of Wy. complosa and Wy. jocosa; to Drs Oswaldo P Forattini and Mafia Anice M Sallum (FSP), Angelica Oliveira (CPRR) and AJ Shelley (NHM), for the loan of specimens; to Main Lemos for the help in the illustrations.

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