Research on the mosquitoes of Angola - Systematic Catalog of ...

!NTA DE INVESTIOAQOE8 

IENTIFICAS DO ULTRAMAR 

ESTUDOS. ENSAIOS 

DOCUMENTOS 

134 

<strong>Research</strong> on the Mosquitoes of 

Angola (Diptera, Culicidae) 

WITH NEW RECORDS, KEYS TO FEMALES AND 

LARVAE, DISTRIBUTION, AND TAXONOMIC AND 

BIOECOLOGICAL NOTES. 

H. RIBEIRO 

and 

HELENA DA CUNHA RAMOS 

LISBOA- 1980

JUNTA DE INVESTIGACOES CIENTiFICAS DO ULTRAMAR 

Rua da Junqueira, 86--1300 LISBOA Pre,co' $00500

ESTUDOS, ENSAIOS 

E 

DOCUMENTOS 

N. ø 134 


Angola (Diptera, Culicidae) 

X THE GENUS CULEX L., 1758. CHECK-LIST 



BIOECOLOGICAL NOTES.

Recebido para publicacCo em 21/2/1978

VOLUMES J/ PUBLICADOS NA COLEC(3,O 

DE 

1 --Contribuico para o Conhecimento da Flora de Mocambique. I- por F. Ascens, 

o Mendonna. 

2- ,,lspectos do Problema da Eroso do Solo em ,d[rica- por Botelho da Costa. 

3- ,,lcerca da Casa e do Povoctmento na Guing por Francisco Tenreiro. 

4--Explorac6es Botdnicas em Timor--por Ruy Cinatti Vaz Monteiro Gomes. 

5--Reconhecimento Preliminar das Formac6es Florestais no Timor Portugus- 

por Ruy Cinatti Vaz Monteiro Gomes. 

6- Madeiras Coloniais- por Luis de Seabra e Manuel P. Ferreirinha. 

7- Contribution la Connaissance Lithologique de l'Archipel du Cap-Vert- 

L. Berthais. 

por 

8--Notas de Zoogeogra[ia e de Histdria das Explorac6es Faunisticas da Guing 

Portuguesa-- por F. Frade. 

9- Resenha Geogri[ica do Distrito da Beira por Egberto Rodrigues Pedro e 

Alfredo Esteves de Sousa. 

10--,,1 Prop6sito da Cultura do ,,lmendoim no Plano de Valorizaco EconOmica 

de Mocambique--por F. Monteiro Grilo. 

11- Estudo do Sistema Raydist e das Redes Hiperb6licas- por Joaquim B. V. 

Soeiro de Brito. 

12--Contribuic6es para o Conhecimento da Flora de Mocambique. II--por F. 

Ascenso Mendon9a. 

13 Os Barn bus na Indgstria da Celulose--por Luis de Seabra. 

14 Os Moluscos de ,dgua Doce do Ultramar Portugus. I--Introduco. Generalidades- 

por J. Fraga de Azevedo e Lidia do Carmo M. de Medeiros. 

15- O Carneiro do Fei]o por A. F. Teixeira Constantino.

16--0s Insectos do Tabaco Armazenado--por A. Antunes de Almeida. 

17--Contribuio para o Estudo da Defesa FitossaniuJria da Copra do Ultramar 

Portugus--por J. M. Cardoso da Costa. 

18--A Entomofauna dos Produtos Armazenados. Os Tribolium spp. (Coleopetera, 

Tenebrionidae)--por F. L. de Faria Estficio. 

19 -- Contribuio para o Estudo da Ecologia de Pachymerus acaciae Gyll. (ColcoF- 

tera, Bruchidae)--pot Jorge Cancela da Fortseca. 

20--Bracystegia spp. de Mocambique--por Manuel Nogueira Ramos. 

21- Cretaceous and Tertiary Nautiloids from Angola--by A. K. Miller and Lee 

B. Carpenter. 

22--Estudos sobre a Cultura do Chri em Mocambique- pot Hillder Lains c Silva. 

23- Ensaios sobre a Titulao de Vacinas contra a Peripneumonia Contagiosa dos 

Bovinos--por Antbnio Martins Mendes. 

24--Les Roches Phosphatdes d'Angola--por Edmond Dartvelle. 

25- Primeiro Reconhecimento Petrogrfico da Circunscri½io do Barut --por Ale- 

xandre Borges e A. V. Pinto Coelho. 

26- Acerca do Equilibrio BioevohSgico dos Povoamentos de Cibes Borassus spp. 

na Guin Portuguesa--por J. F. Castel-Branco e G. C. Tordo. 

27--Movimentos Associativos na ,4frica Negra - pot Silva Cunha. 

28 -- Contribui(So para o Estudo do Microplncton Marinho de Mo('arnbique -- Ior 

Estela de Sousa e Silva. 

29- A Entomofauna dos Produtos A rmazenados. Dermestes maculatus Deg. e Der- 

mestes ater Deg. (Coleoptera, Dermestidae)- por J. Monteiro Guimar.es. 

30- Contribuio para o Estudo do Problema FIorestal da Guind Portuguesa -- l;or 

J. A. Tarares de Carvalho e J. S. de F. Pereira Nunes. 

31 --Os Moluscos de ,,[gua Doce do Ultramar Portugus. I1- Moluscos do Sul do 

Save (Moqamique)--por J. Fraga de Azevedo, Lidia do Cartoo M. de 

Medeiros e Manuel M. da Costa Faro. 

32--Sero-Antropologia das llhas de Cabo Verde. Mesa-Redonda sobre o Homem 

Cabo-Verdiano- por Almerindo Lessa e Jacques Ruffi& 

33- A ExperimentacCo no Poslo de Culturas Regadas do Vale do Limpopo- For 

Ant6nio Henriques de Sousa Falc.o. 

34- O Clima e o Solo de Titnor. Suas Rela½i3es coma Agricultura- por Firmino 

Ant6nio Soares. 

35- A Entomofauna dos Produtos Armazenados. Oryzaephilus mercator (Fauv.) e 

Oryzaephilus surinamensis (L.) (Colcoptera, Cucujidac)- por M. M. Cor- 

delto.

36--A Entomofauta dos Produtos Armazenados. Tenebroides mauritanicus L. 

(Coleoptera, Ostomidae)--por F. S. Nevcs Evaristo. 

37--1mpressSes Digitais nos lndlgenas da Gitinc Portuguesa--pot Leopoldina 

Ferreira Paulo. 

38--Contribuiq3o para o Estudo da Fertilidade da Mulher lndigena no Ultramar 

Portugus--por F. Figueira Henriques, A. Sarmento, J. J. Pals Morals, 

N. Alves Morgado e Eduino de Brito. 

39--Vida do EritrScito Humano Estudada corn o Uso do CrSmio Radioactivo-- 

por F. A. Carvfio Gomes e F. M. Braganga Gil. 

40--Aspectos da Defesa FitossanittJria dos Produtos Armazenados em Anola-- 

por J. P. Amaro e A. J. Soares de Gouveia. 

41--Subsidios para o Estudo do Regime HidrogrtJfico do Porto da Beira--pot 

J. A. Barahona Fernandes. 

42--Primeiro Reconhecimento Petrogrti.[ico da Serra da Gorongosa (Moqambi- 

que)--por A. Vasconcelos Pinto Coelho. 

43---Pastas Celuldsicas de Gramineas. Estudo Laboratorial de Algumas Esptcies da 

MetrSpole e do Ultramar- por Luis de Seabra e Manuel Lopes da Silva. 

44--Etudes stir les Mallophages. Observations sur les Cuclotogaster (lschnocera, 

Philopteridae) Parasites des Galliformes des Genres Francolinus et Pternistis 

-- por JoSo, Tendeiro. 

45--Estudos do Sistema Tellurometer--por Joaquim B. V. Soeiro de Brito. 

46- Glosstrio Internacional dos Termos Usados em Atatomia de Madeira--por 

Manuel P. Ferreirinha. 

47- CianOfitas de S. Tom e Principe--por Joaquim Sampaio. 

48- Contributiot it la Connaissance de la Ggologie de la Province Portugaise de 

Timor--por Robert Gageonnet et Marcel Lemoine. 

49--O Fei]o de Angola. Panorama Actual da Sua Cultura, Comr:rcio c Armaze- 

namento--por A. T. Constantino. 

50--Agrupamento e Caracterizaco P2tnica dos lnd,;genas de Mocambique--por 

Ant6nio Rita-Ferreira. 

51 -- A cerca de Uma Classificacio FitossatffttJria do A rmazenamento -- por Amilcar 

Lopes Cabral 

52--Minerais de Fracc3o Argilosa de Solos de Angola. I--Curvas de Desidrataqho--por 

J. M. Bastos de Macedo, E. P. Cardoso Franco e J. C. Soveral 

Dias. 

53- Caracteriza½io das Principals Utidades Pedoldgicas do --por Jos Carvalho Cardoso.

54--Subsldio para o Estudo da Peripneumonia Contagiosa dos Bovinos em An- 

gola-pot Ant6nio Martins Mendes. 

55--A Entomofauna dos Produtos Armazenados. Corcyra cephMonica (Staint) 

(Lepidoptera, Pyralidae)--por Maria Manuela Carmona. 

56- A Arte Crist na india Portuguesa- por Carlos de Azevedo. 

57 -- Mahamba-- Tentativa de InterpretacCo Artlstica e Psicol6gica de Documentos 

da Arte dos Negros Africanos--por Ant6nio de Oliveira. 

58- Possibilidades de Aplica½5o dos Espcies Ultramarinas nas Diversas lndt$strias 

da Madeira--pot Luis de Seabra. 

59--lnflugncia dos Tratamentos lnsecticidos no Poder Germinativo das Sementes 

(0 Caso Particular do Trigo)--por Ant6nio Henriques Pinto, de Matos. 

60- O Comrcio Mundial de Madeiras Tropicais Africanas- por Manuel P. Fer- 

reirinha. 

61- O Ferro em Medicina--I parte--por Carlos Trinc5o et alii. 

62- O Ferro em Medicina- II parte- pot Carlos Trinc5o et alii. 

63- O Fei]o de Angola Alteraches das Suas Qualidades Culindrias Durante o 

Armazenamento--por A. Teixeira Constantino. 

64--Estudos de Hidrobiologia no Ultramar Portugugs--Contactos corn Labora- 

t6rios Estrangeiros--por Rui Monteiro. 

65- ltudes sur les Mallophages Africains- por Jo5o Tendeiro. 

66- Para a Caracteriza½o das CondieSes Fitossanitrias do Armazenamento- por 

A. L. Cabral, M. I. S. Moreira, A. G. Costa e A. S. de Carvalho. 

67--Foraminiferos da Costa de Mocambique--por J. M. Braga. 

68--CondieSes Fitossanitgrias de Produtos Ultramarinos em Armazgn do Porto 

de Lisboa (Alcdntara-Norte)--por Amilcar Lopes Cabral e A. J. Soares 

de Gouveia. 

69 -- Le Thon Patudo Parathunus obesus (Lowe) et sa Pgche--por Fernando Frade. 

70- Prospec½es e Ensaios Experimentais Apcolas em Angola- pot J. F. Rosrio 

Nunes e G. C. Tordo, com uma > de F. Frade. 

71- Agricultores e Pescadores Portugueses na Cidade do Rio de Janeiro (Estudo 

Comparativo)--por Raquel Soeiro de Brito. 

72--Contribuicgo para o Estudo das Diatomgceas do Lago Niassa (Mocambi- 

que)--por Maria Ins Monteiro. 

73--Contribution to lhe Theory of Certain Non-linear Differential Equations-- 

por Rui Pacheco de Figueiredo. 

74- Aspectos do Povoamento Branco de Angola- por Ilidio do Amaral.

75 -- Notas sobre a Criaqo de Gado Bovino em Angola -- por J. B. Vieira da Silva. 

76--Prospecqo Parasitoldgica em Timor--Subsidios para o Estudo da Fauna 

Parasitoldgica dos Seus Animais Domdsticos--por H. R. B. de Cabrier 

da Silva. 

77- Contribuiqo para o Conhecimento da Flora da Guind Portuguesa- por Ester 

Pereira de Sousa. 

78--Contribuiqo para o Estudo da Tineola bissellie!la (Hummel) e Seu Corn- 

bate-pot M. de Lourdes N. Baptista Peteira. 

79--A Lagarta dos Coqueiros (Nephantis serinopa Meyrick) na india Portu- 

guesa--por A. J. F. Castel-branco. 

80--Sobre a rtMedicina>> e a Magia dos Quiocos--por Eduardo dos Santos. 

81--Estudos sobre a Etnologia do Ultramar Portugus (vol. I)--por Ant6nio de 

Almeida, Eduardo dos Santos, M,Srio Milheiros, Ant6nio da Silva Rego 

e Ant6nio Scarpa. 

82--Reconhecimento Orizicola do Distrito de Goa--por M. S. Portela Feijao. 

83 -- Conspectus da Entomo[auna Cabo-Verdiana--1. a parte--por A. Coutinho 

Saraiva. 

84- Estudos sobre a Etnologia do Ultramar Portuguis (vol. II)--por E. dos San- 

tos, Santa Rita, I!idio Lopes, F. Frade, Diogo d'Orey, A. da Silva Rego, 

G. I. Janz e Ant6nio de Almeida. 

85- Contribuiqo para o Estudo da Gdnese dos Minerais da Argila--por J. M. 

Bastos de Macedo e M. A. Monteiro de Lemos. 

86--Contribuiqo para o Estudo das Espdcies Angolanas do Gdnero Culicoides 

Latreille, 1909 (Diptera: Ceratopogonidae)--por Vitor Manuel Pais 

Caeiro. 

87--Caracterœsticos dos .4cidos Htmidos de Alguns Solos de Angola--por Rui 

Pinto Ricardo. 

88- Os Moluscos de Agua Doce do Ultramar Portugus. IH--Moluscos de Mo- 

cambique--por $. Fraga de Azevedo, Lidia do C. M. de Medeiros, M. 

M. da Cos.ta Faro, Maria de Lurdes Xavier, A. Franco Ghndara e Tito 

de Morais. 

89--O Reaparecimento da Glossina palpalis palpalis na llha do Principe--pot 

J. Fraga de Azevedo, J. Tendeiro, L. T. de A!meida Franco, M. da Costa 

Mour.o e J. M. de Castro Salazar. 

90--Introduqo ao Estudo Tcnico-EconSmico da Criaqo de Gado Bovino em 

Angola (A Economia da Carne)--por I. Lima Pereira. 

91 -- A Erradicaqo da Glossina palpalis palpalis da Ilha do Principe (1956-1960) -- 

por I. Fraga de Azevedo, M. da Costa Mouro e J. M. de Castro Salazar. 

92--Os Fendmenos 

de Metamor[ismo e Granitizao no Distrito de Goapor 

J. Avila Martins.

93- Contribution fi I'llrude de la Famille du Pa TM par des Corrt;latious Angulaires 

de Quelques Cascades .(..( et ..( --par Fernando Braganqa Gil. 

94--Os Solos da llha de S. Nicolau (Arquipdlago de Cabo l'erde)--por Mateus 

Nunes. 

95--1ntroduco Antropologia do Ultramar--por A. Athaide, M. E. de Castro 

e Almeida e A. A. Mendes Correia. 

96- Sobre a Religio dos Quiocos- por Eduardo dos Santos. 

97- Ensaio de um Estudo Geogrd[ico da Rede Urbana de Angola--por Ilidio do 

Amaral. 

98--Lhe Thon Rouge et le Germon Thunnus thynnus L. et Germon alalunga 

(Bonn.)--par F. Frade et. H. Vilela. 

99--Petrologia das Lavas dos Libombos (Subsidios para o Seu Conhecimento)-- 

pot Torre de Assungo, A. V. T. Pinto Coeiho e A. Tarares Rocha. 

100--Os Solos da Guin Portuguesa--por Silva Teixeira. 

101--O Problema de S. Nicolau por Mateus Nunes. 

102--Estudos sobre a Etnologia do Ultramar PortuguSs (vol. III)--por M. E. de 

Castro e Alineida, Miguel Vieira, Ant6nio Carreira, Jos6 Lampreia e W. 

A, A. wilson. 

103--Mineralogia dos Jazigos de Ferro e de Mangangs de Goa--por Luis Aires 

Barcos. 

104- Nomes Verndculos de ,41gumas Plantas da Guind Portuguesa- por J. do Espirito 

Santo. 

lOS--Inheritance and Relationships ,4mong Growth Characters of Young Cacao 

Seedlings--by Josa Crespo Ascenso. 

106- Segunda Contribuigo para o Estudo das DiatomJceas do Lago Niassa (Moambique)--por 

Maria lnCs Monteiro. 

107--Entomofauna de S. Tomd (lnseclos de Cacueiro)--pot A. J. Castel-branco. 

108- Ciandfitas de S. Tomd e Principe (3. Sdrie)--por Joaquim Sampaio. 

109--,4 Entomofauna dos Produtos ,4rmazenados. Contribuio do Mt;todo Radiogrdfico 

para o Estudo da Sitotroga ceralella (Oily.) (Lepidoptera: Gelechiidae)--por 

Passos de Carvalho. 

110,4 Vegetao do Extremo Sul da Provincia de Mocambique (Contribui½o 

para o Seu Estudo)--pot Mrio Myre. 

111- Gramineas Novas da Guind Portuguesa--por Maria Margarida Pinto de 

Castro e Jos6 Vicente C. Malato Beliz. 

112 

Identification of Metamict Minerals by X-Ray Powder Pholographs- por J. 

Lima de Facia.

113---Reconhecimento Glossinico de Uma ,,[rea do Concelho do Dondo e Esquema 

para a Recupera½Oo de Parte da Mesma Mosca Tsr-Tsd--por J. A. 

Travassos Santos Dias. 

114- O Hornera nos Tr6picos. Aspectos Bioecoldgicos- por J. Fraga de Azevedo, 

com um prefficio do Prof. Adriano Pond. 

115--Reserva Mineral e Minerais da Areia de Alguns Solos de Cela (Angola)- por 

Carlos A.M. Portas e A. F. Sanches Furtado. 

116- Israel--Curso lnternacional de Rega--por Raul Wahnon Correia Pinto. 

__ 

117 -- Vulcanismo das Ilhas de Cabo Verde e das Outras llhas Atlntidas -- lor Frederico 

Machado. 

118- Mineralogia dos Solos de S. Tom e Prbcipe- por J. Bailim Pissarra, J. Car- 

valho Cardoso e J. Sacadura Garcia. 

119-- Elementos de Vulcanologia-- por Frederico Machado. 

120 -- Estudo da lntrluncia de Alguns Factores Bioecolgicos e Toxicoldgtcos na Sus- 

ceptibilidade do ,,fcaro da Farinha, Acarus siro L. (Acarina, Acaridae)- 

por Joaquim Pedro Ferreira Amaro. 

121 --Contribui½Oo para o Estudo do Decllnio do 277 Th- por Fernando Monteiro 

de Braganna Gil. 

122- Contribui½Oo para o Estudo do Grupo Tamanduri da Srie Rio das Velhas 

Minas Gerais, Brasil)--por Joo da Rocha Hirson. 

123--A Ovinicultura de Li em Regi6es Tropicais (Bases para o Fomento Zootc- 

nico da Criaco de Ovinos de LO em Angola)--pot J. Lima Pereira. 

124--Os Solos da llha de Santiago (Arquipdlago de Cabo Verde)--por Fernando 

Xavier de Faria. 

125- Curso de Sismologia--por Frederico Machado. 

126- Contribui½Oo para o Estudo das Caractersticas dos Ca)ds de Angola--por A. 

Bai.o Esteves e J. Santos Oliveira. 

127- A Evolu½Oo da Geodesia e a Ocupa½Oo Geoddsica do Ultramar Portugu& em 

,,[1'fica- por Jos Farinha da Concei.o. 

128- Estudo das Caractersticas Fecnol6gicas das Pastas Produzidas lndustrialmente 

corn Madeiras de Eucalipto--por J. L. Ferreira da Silva Dias. 

129--Os Solos da Ilha do Fogo (Arquiplago de Cabo Verde)--por Fernando 

Xavier de Faria. 

130--S. Tom e Prlncipe sob o Potno de Vista Agricola--por F. M. de Carvalho 

Rodrigues. 

130-A- S. Tomd e Prlncipe sob o Ponto de Vista Agricola (Cartas Agricolas)--por 

F. M. de Carvalho Rodrigues 

131- Caracteristicas de Ernpilhamento e Modelos Condensados das Micas e Filos- 

silicatos A)%s--por Maria Ondina Dionisio Figueiredo.

132--Flora de S. Tomd e Pr[ncipe (,,[cidos Gordos e Protelnas de AIguma Semen- 

res) por J. E. Mendes Ferr5o. 

133- F, studos sobre os Goniodideos (Mallophaga, Ischnocera) dos Columbiformes 

Gdnero Physconelloides Ewing--por Joao Tendeiro

CDU 576.895.711 (673) 

JUNTA DE INVESTIOAGOES CIENTiFICAS DO ULTRAMAR 


Angola (Diptera,, Culicida 0 

X--THE GENUS CULEX L., 1758. CHECK-LIST 



B1OECOLOGICAL NOTES. 

by 

H. RIBEIRO 

and 

HELENA DA CUNHA RAMOS 

LISBOA- 1980

CONTENTS 

I -- INTRODUCTION ............... 17 

2--HISTORICAL OUTLINE ..... 19 

3--GAZETTEER OF COLLECTING LOCALITIES 

4--ON THE ZOOGEOGRAPHY OF ANGOLA ... 27 

5--L1ST OF THE ANGOLAN SPECIES, SUBSPECIES AND FORMS 29 

6--KEYS TO FEMALES AND LARVAE OF THE ANGOLAN MOS- 

QUITOES .................................. 

7 -- SYSTEMATIC ACCOUNT .......... 51 

7.1 --Subgenus Lutzia Theobald, 1903 ..... 

7.2-- Subgenus Maillotia Theobald, 1907 

7.3 -- Subgenus Eumelanomyia Theobald, 1909 

7.4 -- Subgenus Culiciomyia Theobald, 1007 ... 

7.5-- Subgenus Culex Linnaeus, 1758 . . 

8--RELATION TO DISEASE ........... 125 

ACKNOWLEDGEMENTS ............... ... 127 

SUMMARY .................................. 129 

RESUME .................................... 131 

RESUMO ........................... 133 

REFERENCES ................................... 137 

51 

54 

_55 

62 

65

I --INTRODUCTION 

With this paper on the genus Culex L., the writers conclude 

a main series of articles revising the mosquito fauna of Angola. 

The present paper is based upon the examination of more than 

8,400 Culex specimens (near 3,100 adults and more than 5,300 larvae) 

collected throughout Angola during the period from 1963 to 1971, 

and is intended to include all the previously published distributional 

data concerning the Angolan representatives of the genus. 

Estud., Ens. e Doc.- 134 17

2 -- HISTORICAL OUTLINE 

Culex pipiens was the first species of genus Culex to be recorded 

from Angola (Mog/tmedes), in 1903, by B. Roque (1). In 1905, Giles 

(2) recorded C. poicilipes among material sent by Dr. Yale Massey 

from Benguela. F. C. Wellman (3) (4) and F. C. Wellman & W. E. 

Fay (5), in 1905 and 1907, added C. guiarti, C. theileri, C. duttoni and 

C. univittat,ts to the Angolan mosquitoes in the Bifi-Bailundo area. 

The next new C,dex records for Angola were C. annulloris by Theobald 

(6), in 1910, from Catema (Bi) and, in 1914, C. invidiosus, C. 

decens, C. rima and C. tigripes by M. Gamble (7), from So Salvador. 

With the new records C. annulioris annulioris and C. pipiens pipiens, 

in 1914, by Edwards (8) and C. argenteopunctatus, C. horridus, C. 

nebulosus, C. pipiens quinque[asciatus and C. thalassius, in 1952, by 

A. T. F. Colato (9), 15 Culex species and subspecies became known 

from Angola. In 1956, our knowledge on the Angolan mosquitoes was 

definitely improved with the revision carried out by A. F. G/tndara 

(0) (1) who added, besides much new distributional data, 12 

new Culex records to the previous list: C. annulioris consimilis, C. antennatus, 

C. argenteopunctatus kingi, C. cinerellus, C. cinereus, C. 

grahami, C. inconspicuosus, C. rubinotus, C. simpsoni, C. telesilla, 

C. watti and C. :ombaensis. In 1961, C. B. Worth & H. E. Paterson (12) 

added C. ingrami, C. insignis, C. invidiosus vexillatus, C. ornatothoracis, 

C. tri[ilatus and C. tri[oliatus to the previous list of Angolan 

species and subspecies, extending also considerably our knowledge on 

their geographical distribution. In 1966, one of us (3) recorded C. 

bitaeniorhynchux, C. tritaeniorhynchus and C. univittatus neavei as 


RIBIglRO, H. & RAMOS, H. Cunha -- <strong>Research</strong> on the mosquitoes of Angola X 

new for Angola, besides some new distributional data. Finally, in 1967, 

Brunhes et al. (14) described C. quintetti on the base of material from 

Man (Ivory Coast) and recorded it from Dundo (North-eastern Angola), 

increasing to a total of 35 the number of species and subspecies of 

Culex presumed to be known from Angola. 

20 Estud., Ens. e Doc.- 134

3 m GAZETTEER OF COLLECTING LOCALITIES 

In Map 1 are shown the collecting localities concerned in this 

paper, which approximate coordinates and altitude are given below. 

137 out of the 152 assigned localities were surveyed by the writers, 

while those marked with an asterisk were only surveyed by other 

Authors. 

1 -- Alto Catumbela ............... 12 ø 58' S 14 ø 45' E 1,250 m 

2 -- Ambrizete ........................ 7 ø 14' S 12 ø 52' E sea coast 

3-- Baa dos Tigres ............... 16 ø 36' S 11 ø 43' E sea coast 

4 -- Baia Farta ........................ 12 ø 37' S 13 ø 13' E sea coast 

5--Bailundo (-Teixeira da Silva) 

* ................................ 12 ø ITS 

6 --Barra do Cuanza ............... 9 ø 08'S 

7 --Barra do Dande ............... 8 ø 29'S 

8 -- Belize .............................. 4 ø 38' S 12 ø 42'E 240m 

9 Benguela ............................. 12 ø 35' S 13 ø 25'E sea coast 

10-- Bero River (N of Virei) ...... 15 ø 37' S 12 ø 51'E 350m 

11-- Bibala (= Vila Arriaga) ...... 14 ø 46' S 13 ø 21'E 920m 

12-- Bocoio (= Sousa Lara) ...... 12 ø 29' S 14 ø11'E 1,000m 

13-- Bruco .............................. 15 ø 08' S 13 ø10'E 700m 

14-- 

15-- 

16-- 

17-- 

18-- 

19-- 

15 ø 52' E 1,650 m 

13 ø 10' E sea coast 

13 ø 24' E sea coast 

Buco Zau ........................ 4 ø 45' S 12 ø 31'E 350m 

Bundas (=Gago Coutinho) * 14 ø 06' S 21 ø 26'E 1,130m 

Caala (= R. Williams) ......... 12 ø 51' S 15 ø 35'E 1,700 m 

Cabinda ........................... 5 ø 33' S 12 ø 11'E sea coast 

Cabuta ............................ 9 ø 50' S 14 ø 52'E 750m 

Caconda* ......................... 13 ø 43' S 15 ø 05'E 1,650m 

Estud.. Ens. e Doc.- 134 21

RIBgIRO. H. & RAMOS, H. Cunha -- <strong>Research</strong> ou the mosquitoes o/ Angola -- X 

20-- Cacuaco .......................... 8 ø 46' S 130 20' E sea coast 

21 -- Cafima ............................ 16" 39' S 16 ø 37' E 1,150 m 

22 -- Cainde ............................ 15 ø 27' S 130 22' E 900 m 

23 -- Calai ............................... 17" 51' S 19" 27' E 1,050 m 

24 -- Calulo .............................. 10 ø 00' S 140 54' E 950 m 

25 -- Cangandala ....................... 9" 50' S 16 ø 20' E 1,050 m 

26--Cangandala (24km SE) ...... 9"55'S 16"36'E 1,100m 

27 -- Capelongo (=Folgares)* ...... 140 53' S 15 ø 04' E 1,270 m 

28 -- Capunda .......................... 10" 41' S 17" 22' E 1,130 m 

29 -- Caraculo .......................... 15 ø 01' S 120 40' E 440 m 

30-- Cariango .......................... 10 ø 30' S 15" 25' E 1,500 m 

31 --- Carlaongo ........................ l0 ø 45' S 14" 15' E 200 m 

32 -- Cassoalala ........................ 9" 28' S ld" 25' E 200 m 

33 -- Ca[rna* .......................... 12 ø12'S 16ø47'E 1,750m 

34 -- Catete ............................. 9" 06' S 13" 42' E 120 m 

35 -- Catumbela ........................ 12" 18' S 13" 45' E 100 m 

36 -- Caxito ............................. 8" 35' S 13" 41' E 20 m 

37--Ceilunga (=circa Bifi=S. S. 

Porto) .............................. 12 ' 24'S 16 ø 57'E 1,700m 

38 --Chfio de Chela .................. 15 ø 08' S 13" 14' E 1,120 m 

39-- Cherugema ...................... 16" 18'S 12 ø 27'E 600m 

40-- Chicala ............................ 12" 48' S 17 ø 03' E 1,500 m 

41-- Chiede ............................. 170 09'S 15ø 59' E 1,110m 

42 -- Chinguar .......................... 12" 33' S 16 ø 20' E 1,800 m 

43 -- Chiquite ........................... 13 ø 46'S 130 07'E 700m 

44 -- Chissamba (=Nova Sintra)* 120 08'S 17"17'E 1,550m 

45 -- Chitado ........................... 170 19' S 13 ø 55'E 1,000 m 

46 --- Chitembo ......................... 130 32' S 160 46' E 1,500 m 

47 Cuando ........................... 120 47'S 15 ø 55'E 1,700m 

48-- Cuangar ........................... 17 ø 35'S 18 ø 39' E 1,050 m 

49-- Cuango ............................ 9 ø 06'S 180 05'E 850m 

50-- Cuanza River (N of Callulo) 90 41' S 140 48' E 700 m 

51-- Cubal .............................. 13 ø 01'S 14 ø 17'E 900m 

52-- 

53-- 

54-- 

Cuchi* ............................. 14 ø 38'S 160 58'E 1,430m 

Cui (= Sumbo) .................. 

Cunene River (S of Espi- 

13 ø 39' S 140 12'E 1,800 m 

nheira) ............................. 17 ø 09' S 120 Off E 200 m 

22 Eslud., Ens. e Doc.- 134

RIBgIRO, H. & RAMOS, H. Cunha -- <strong>Research</strong> on the mosquitoes of Angola -- X 

55 1 Curoca River (N of p.o Ale- 

xandre) ............................ 15 ø 45' S 11ø 55' E 50 rn 

56 -- Dala Tando ...................... 90 19' S 14 ø 57' E 690 rn 

57 -- Damba ............................ 6 ø 41' S 15 ø 08' E 1,160 m 

58--Dando ............................. 11 ø15'S 17ø26'E 1,150m 

59 --- Dange-Ia-Menha ............... 90 31' S 14 ø 38' E 250 m 

60--- Dilolo (=Teixeira de Sousa)* 10 ø 43' S 12 ø 15' E 1,100 rn 

61 -- Dinde .............................. 14 ø 19' S 13 ø 42' E 900 m 

62 -- Dirico .............................. 17 ø 58' S 20 ø 47' E 1,200 m 

63 -- Dundo ............................. 7 ø 22' S 20 ø 50' E 730 m 

64 -- Duque de Braganqa ............ 90 06' S 15 ø 58' E 1,060 m 

65 -- Ebanga ............................ 12 ø 42' S 14 ø 45' E 2,000 m 

66--Ebo ................................. 11 ø02'S 14 ø41'E 1,400m 

67 -- Equirnina ......................... 13 ø 11' S 12 (' 48' E sea coast 

68 -- Foz do Cuncne .................. 17 ø 16' S 11 ø 45' E sea coast 

69-- Foz do Queve .................. 10 ø 50' S 13 ø 49' E sea coast 

70--Ganda (--Mariano Macha- 

do)* ................................ 13 ø 01' S 14 ø 40' E 1,750 rn 

71 -- Giraul River ..................... 15 ø 05' S 12 ø 17' E 100 rn 

72--Gungo ............................. 11ø48'S 14 ø11'E 900m 

73-- Hanha ............................. 12 ø 15' S 13 ø 40' E sea coast 

74 -- Huambo (:Nova Lisboa) ... 12 ø 48' S 15 ø 45' E 1,700 m 

75 -- Hunguria ........................ 15 ø 20' S 13 ø 31' E 1,400 m 

76 -- Impulo ............................ 13 ø 53' S 130 39' E 800 m 

77 -- Inhuca River .................... 4 ø 45' S 12 ø 25' E 80 rn 

78 -- Iona ................................ 16 ø 53' S 12 ø 34' E 1,000 rn 

79 Jamba .............................. 15 ø 05' S 16 ø 05' E 1,300 m 

80--Jangada do Cungar 1,100m 

(50 km W) ........................ 17 ø 52' S 17 ø 59' E 

81 -- Lagoa Banda .................... 8 ø 50' S 13 ø 34' E 

100 m 

82 -- Lagoa dos Paralelos ........... 15 ø 50' S 12 ø 45'E 

420 m 

83--L.ndana (= Guilherme Ca- 

sea coast 

pelo) ............................... 5 ø 13' S 12o 08'E 

84 -- Lifune ............................. 8 ø 23' S 13 ø 25' E 

50 rn 

85 -- Lobito ............................. 12 ø 22' S 

86 -- Lombe River (W of Malanje) 9 ø 24' S 

87 -- Lonjava ........................... 12 ø 49' S 

88 -- Luanda ............................ 80 49' S 

13 ø 32' E 

16 ø 12' E 

14 ø 40' E 

13 ø 13' E 

sea coast 

1,100 m 

1,200 rn 

sea coast 


RIBEI}O, H. & R.MOS, H. Cunha -- <strong>Research</strong> on the mosquitoes of Atgola -- X 

89-- Luanda River ................... 10 ø 55' S 17 ø 38' E 

90 -- Luele River (Source of) ...... 9" 55' S 20 ø 05' E 

91 -- Lubiri .............................. 12 ø 08' S 150 27' E 

92-- Lucala ............................. 9 ø 18' S 15 ø 13' E 

93 -- Lucala River .................... 9 ø 27' S 14 ø 45' E 

94-- Luce River (S of Quimbango) 11 ø 17' S 17 ø 36' E 

95 -- Lucira ............................. 130 52' S 12 ø 31' E 

96-- Luinga ............................. 8 ø 27' S 15 ø 38' E 

97--Luso (=Moxico) ............... 11ø47'S 19"55'E 

98 -- Maculama (NE of Iona) ...... 16 ø 44' S 12 ø 38' E 

99 -- Malanje ........................... 9 ø 33' S 16 ø 22' E 

100-- Maquela do Zombo * ......... 6" 03' S 15 ø 06' E 

101 -- Marco de Canavezes ......... 12 ø 59' S 14" 23' E 

102 -- Mavinga * ........................ 15" 50' S 20" 21' E 

103-- Mazozo ............................ 9 ø 07' S 13 ø 37' E 

104 -- Mogfimedes ...................... 15" 12' S 12 ' 09' E 

105 -- Mona Quimbundo ............. 9" 49' S 19" 59' E 

106--M'pupa ............................ 17" 35'S 200 01'E 

107-- Mucuio ............................ 14 ø 54' S 12 ') 12' E 

108 -- Mulondo * . ....................... 15" 38' S 15 ø 12' E 

109-- Mumbondo ....................... 10" 10' S 14" 09' E 

ll0--Munhino .......................... 14 ø 58'S 12 ø58'E 

111--Munhino (25km W) ......... 14 ø 53'S 12 ø 47'E 

112 -- Mussende ........................ 10" 30' S 16 ø 02' E 

113 -- Mussulo ........................... 8 ø 55' S 13 ø 06' E 

114 -- Mutangc .......................... 170 51' S 19 ø 53' E 

115 -- Muxima ........................... 9" 32' S 13" 57' E 

116--Nangura (=Maiuvo) ......... 17ø49'S 19ø01'E 

117--N'giva (=Peteira d'Ega) ..... 17ø04'S 15ø4YE 

118 -- Novo Redondo .................. ll ø12'S 13ø51'E 

119-- Onc6cua .......................... 16" 39' S 13 ø 25' E 

120 -- Panda .............................. 17 ø 25' S 18 ø 30' E 

121 -- Pediva ............................. 16 ø 17' S 12" 34' E 

122 -- Pico do Azevedo ............... 15 ø 28' S 12 ø 25' E 

123 Porto Alexandre ............... 15 ø 48' S 11ø 51' E 

124 -- Porto Amboim .................. 10 ø 44' S 13 ø 46' E 

125--Quela .............................. 9 ø16'S 17ø05'E 

126-- Quicuco ........................... 14 ø 17' S 13 ø 51' E 

1,180m 

1,200 m 

1,500 m 

950 m 

850 m 

1,150 m 

sea coast 

1,200 m 

1,320 m 

620 m 

1,140 m 

925 m 

900 m 

1,190 m 

100 m 

sea coast 

1,150m 

1,150 m 

sea coast 

1,200 m 

1,150m 

480 m 

460 m 

1,370m 

sea coast 

1,050 m 

20 m 

1,100 m 

1,100 m 

sea coast 

1,080 m 

1,000 m 

350 m 

370 m 

sea coast 

sea coast 

1,200 m 

1,100 m 

24 Estud., Ens. e Doc. 134

RIBEIRO, H. & RAMOS, H. Cunha -- <strong>Research</strong> on the mosquitoes o[ Atgola -- X 

127 -- Quifangondo ..................... 8 ø 46' S 13 ø 27' E 80 m 

128 -- Quilengues ....................... 14 ø 06' s 14 ø 05' E 900 m 

129 -- Quimbango ....................... 10 ø 53' S 17 ø 32' E 1,200 m 

130 -- Quissol ............................ 9 ø 29' s 16 ø 30' E 1,100 m 

131 -- Quissongo ........................ 10 ø 02' s 15 ø 05' E 1,200 m 

132-- Quitexe ........................... 7" 45' s 15 ø 03' E 900 rn 

133 -- Rio Gango ....................... 10 ø 37' S 15 ø 57' E 1,600 m 

134-- Saco do Giraul .................. 12 ø 06' S 15 ø 07' E sea coast 

135 --- Sfi. da Bandeira (=Lubango) 14 ø 56' S 13 ø 34' E 1,760 rn 

136--Sfi. da Bandeira (20 km W) 14ø44'S 13ø27'E 1,600m 

137--Sambo* ........................... 13ø04'S 16ø08'E 1,710m 

138-- Silo Nicolau ..................... 14 ø 22' S 12 ø 23' E sea coast 

139 -- Silo Salvador do Congo * ... 6" 16' S i4 ø 15' E 560 rn 

140-- Saurimo (=H. de Carvalho) 9 ø 09' S 20" 24' E 1,080 rn 

141 -- Serpa Pinto ..................... 14" 40' S 17 ø 42' E 1,350 m 

142---Serra da Neve .................. 13ø40'S 13 ø10'E 1,250/1,450m 

143 -- Tando Zinze ..................... 5 ø 25' S 12 ø 25' E 150 m 

144 -- Tchivinguiro ..................... 15 ø 10'S 13 ø 18'E 1,690m 

145 -- Tombinga ........................ 9 ø 21' S 14 ø 47' E 320 rn 

146 -- Ueca ............................... 170 50' S 20 ø 10' E 1,050 m 

147--Uige ............................... 7 ø35'S 15ø00'E 830m 

148--Vale do Loge .................. 7 ø 17'S 14 ø 26'E 530m 

149 -- Vilar Flor ........................ 12 ø 40' S 15 ø 33' E 1,750 m 

150--Vila Nova do Seles ............ 11 ø 24'S 14 ø 18'E 1,000m 

151--Virei .............................. 15ø44'S 12ø57'E 450m 

152 -- Virei (25 km E) ............... 15 ø 45' S 13 ø 08' E 420 rn 

Estud., Ens. e 1)oc.- 134 25

4--ON THE ZOOGEOGRAPHY OF ANGOLA 

On Map 2, modified from Moreau (15), are shown the main biotic 

zones in Africa South of Sahara as well as the general position of Angola 

within the Ethiopian Region. 

Map 3 shows the tentative zoogeographical division of Angola as 

it was already proposed by the writers in a previous paper (16) on the 

Angolan mosquitoes of genus Anopheles M. 


5--LIST OF THE ANGOLAN SPECIES, SUBSPECIES AND FORMS 

In the establishment of the list of the Angolan members of genus 

Culex L., both our unpublished data and all published contributions 

available to us were taken into account. 

The basic systematic arrangement and taxonomic treatment followed 

by the writers in this paper is that of the World Catalog by Stone, Knight 

and Starcke (17) [including the Supplements (18) (19) (20) (21) and Corrections 

by Stone (22)], with the modifications introduced in the new edition 

by Knight & Stone (23). In a few instances, however, we adopted a somewhat 

divergent taxonomic treatment, as it will be mentioned under the 

corresponding taxa (see under C. annulioris, C. invidiosus, C. nebulosus, 

C. simpsoni, C. univittatus and also C. pipiens). 

On the other hand, some more recent taxonomic contributions by 

other Authors will be also confronted with the available evidence from 

Angola (see under C. pruina and C. univittatus). 

Some corrections made on the previously reported list of Culex species 

from Angola (see under 2), namely by treating C. annulioris and C. 

univittatus as monotypic species, lead to shorten it to only 34 members. 

On the other hand, it is established that the nominal subspecies of the 

polytypic C. trifilatus and C. tritaeniorhynchus are those occurring in 

Angola. 

Lastly, the list of the Angolan members of genus Culex L., as it 

is shown in Table 1, is now thought to comprise 47 names, 13 of which 

are here recorded for the first time in the country. From the viewpoint 

of the geographical distribution, our knowledge on the genus in Angola 

is also considerably extended with, on the whole, 447 new locality records 

(Table 1). 

Estud., Ens. e Doc.---134 29

6--KEYS TO FEMALES AND LARVAE OF THE ANGOLAN MOS- 

QUITOES 

The construction of our keys was mainly based on the classical 

works of Edwards (8) and Hopkins (24) as well as on other further contributions, 

such as those of van Someren (25) (26), Mattingly (27) and De 

Meillon et al. (28). On the other hand, attempts were also made by the 

writers in order to improve the usual keys by precising some diagnoses 

and by taking into account the range of variation observed in Angolan 

material. 

It must be noted, however, that individual larvae of C. antennatus, 

C. decens, C. invidiosus, C. telesilla, C. trifoliatus and C. univittatus are 

often indistinguishable, even though the identification of samples of these 

species will be usually feasible using the larval key here proposed. 


RIBgIR0, H. & RAMOS, H. Cunha -- <strong>Research</strong> on the mosquitoes o! dtlgOla X 

TABLE 1 

LIST OF THE ANGOLAN MOSQUITOES OF THE GENUS iCULEX 

Subgenera Species, subspecies and forms 

New records 

Taxa Localities 

Lutzia ............ C. tigripes Grandpr6 and Charmoy, 1900 41 

Maillotia ......... C. salisburiensis salisburiensis Theobald, 

1901 ............................................... q- 2 

[ C. horridus Edwards, 1922 .................... 1 

C. inconspicuosus (Theobald, 1908) ......... 14 

C. insignis (Carter, 1911) ........................ 1 

C. kingianus Edwards, 1922 .................. q- 2 

Eumelanomyia .. , C. quintetti Brunhes, Adam & Bailly-Chou- 

mara, 1967 ....................................... -- 

C. rima Theobald, 1901 ........................ -- 

C. rubinotus Theobald, 1906 .................. 2 

C. wigglesworthi Edwards, 1941 ............... q- 1 

: 

C. cinerellus Edwards, 1922 .................. 

Culiciomyia ..... i C. nebulosus cinereus Theobald, 1901 .................. 

[ I i 15 3 

[ C. annulioris Theobald, 1901 .................. [ 2 o 

C. antennatus (Becker, 1903) .................. 21 

C. argenteopunctatus kingi (Theobald, 1913) 9 

C. aurantapex ]iniaensis Edwards, 1941 ... q- 2 

C. bitaeniorhynchus Giles, 1901 ................ 

C. chorleyi Edwards, 1941 ..................... q- 5 

C. decens Theobald, 1901 ..................... 24 

C. duttoni Theobald, 1901 ..................... 36 

C. ethiopicus Edwards, 1912 .................. q- 11 

C. grahami Theobald, 1910 ..................... 3 

C. guiarti Blanchard, 1905 ..................... 4 

C. ingrami Edwards, 1916 ..................... 6 

C. invidiosus Theobald, 1901 .................. . q- 5 

C. invidiosus form vexillatus Edwards, 1941 2 

C. ornatothoracis Theobald, 1909 ............ 1 

C. per[idiosus Edwards, 1914 ................. q- 3 

C. per[uscus Edwards, 1914 .................. q- 1 

Culex ............. { I 

C. piplens piplens Linnaeus, 1758 ............ I 15 

C. pipiens quinque[asciatus Say, 1823 ...... 36 

C. poicilipes (Theobald, 1903) ............... ! 17 

C. pruina pruina Theobald, 1901 ............ + 1 

C. pruina eschirasi Galliard, 1931 ......... + 1 

C. simpsoni Theobald, 1905 .................. 40 

C. telesilla De Mei!lon and Lavoipierre, 1945 7 

C. thalassius Theobald, 1903 .................. 13 

C. theileri Theobald, 1903 ..................... 9 

C. toroensis Edwards and Gibbins, 1939 ... q- 1 

C. trifilatus tri[ilatus Edwards, 1914 ...... q- 1 

C. tri/oliatus Edwards, 1914 .................. 2 

C. tritaeniorhynchus tritaeniorynchus Giles, 4 

C. univittatus 

watti Edwards, Theobald, 

1920 ........................ 1901 ................. I q- -- 45 

C. weschei ? gediensis Edwards, 1941 ...... q- 2 

C. zombaensis Theobald, 1901 .............. [ 3 

Totals ............ 47 15 447 

6

RIBEIRO, H. & RAMOS, H. Cunha -- <strong>Research</strong> on the mosquitoes of Angola -- X 

Key to Females (') 

With 3-10 (usually, 4 or more) lower mesepimeral 

bristles; front and middle femora and tibiae each 

with a row of about ten small pale spots anteriorly; 

proboscis without a complete pale ring in middle 

(subgenus Lutzia) ............................................... 

Without this combination of characters .................. 

1 Acrostical bristles absent, though dorsocentral bris- 

tles and those above wing root are unusually long, 

strong and numerous (sub& Eumelanomyia, in part) 

--Acrostical bristles present, at least towards the front 

margin of scutum ................................................ 

3--No lower mesepimeral bristle .............................. 

--With 1-4 lower mesepimeral bristles ........................ 

-- Proboscis and tarsi dark (subg. Eumelanomyia, in part) 

--Proboscis and tarsi distinctly ringed (subg. Culex, 

in part) ............................................................ 

--Femora and tibiae with rows of small pale spots 

anteriorly .......................................................... 

--No such rows of spots (though scattered pale scales 

may be present) ................................................ 

--Wings with numerous scattered pale scales; probos- 

cis with at least a few white scales at tip above 

(before labella) ................................................... 

Wings with few or no pale scales; proboscis dark at 

tip (though labella may be pale) ........................... 

tigripes 

horridus 

rubinotus 

2 

4 

11 

poicilipes 

7--Abdominal tergites n-vn with complete pale apical 

bands ................................................................ ethiopicus 

(x) Female of Culex (E.) quintetti not described. 



101 

--Abdominal tergite tt entirely dark above and tergites 

IIt-V with apicolateral pale spots but without com- 

plete bands ................................................ bitaeniorhynchus 

--Thorax black, scutal scales nearly all black; tergites 

v-w with orange apical lateral spots ...... aurantapex jinjaensis 

Thorax brownish or scuturn with many pale scales; 

apical pale markings on tergites, if present, not 

orange .............................................................. 9 

_ Scutum with anterior -2/3 mainly pale scaled; 

abdominal tergites varying from almost all pale to 

all dark or nearly so (usually with pale lateral api- 

cal spots in addition to pale basal band, these pale 

markings on a mainly dark ground); costa entirely 

dark .................................................................. 

Scutal scales dark, pale scales either forming an indefi- 

nite mottled pattern or present only near scutellum; 

abdominal tergites with basal pale bands and without 

apical pale markings; costa with a small spot of pale 

scales at base ...................................................... 

annulioris 

111 Abdominal tergites with only apical pale markings 

(bands or lateral spots) or completely dark scaled; 

decumbent scales on vertex ranging from all narrow 

to mostly broad ............................................................ 12 

--Abdominal tergites with basal pale markings; all decumbent 

scales of vertex, including the ocular row, 

12 1 

Pale ring of female proboscis extended proximally, at 

least beneath; thorax brown, pale scales almost 

completely confined to near scutellum ............ tritaeniorhynchus 

--Pale ring restricted to about the middle third of proboscis, 

sharply defined proximally; thorax blacker, 

with some pale scales on front half of scuturn ..... thalassius 

quite narrow (subg. Culex, in part) ........................ 21 

All decumbent scales on vertex narrow (in kingianus, 

the flat whitish scales of sides extending upwards 

36 EstutL, Ens. e Doc. 

10 

134


13-- 

141 

15-- 

161 

17 1 

18q 

19 w 

as a narrow border to eyes) (subgg. Maillotia and 

Eumelanomyia) ................................................... 

With broadish scales on vertex, at least in a row 

adjoining eyes ..................................................... 

Pleurae brownish or dark brown ........................... 

Pleurae partly dark and partly light ..................... 

Mesepimeron with a large patch of scales; propleura 

with flat white scales which extend also over upper 

third of prosternum (subg. Maillotia) .................. s. salisburiensis 

Mesepimeron with very few or no scales; no scales 

on propleura and prosternum .............................. kingianus 

Erect scales on head all or almost all dark; middle 

third of mesepimeron only indistinctly pale ........... 

--Some of the erect scales on head whitish; middle 

third of mesepimeron clear yellowish ..................... 

13 

17 

14 

15 

rima 

Head with only a few of the erect scales entirely 

whitish .............................................................. insignis 

Head with a large patch of erect white scales ...... wigglesworthi 

Broad scales on head white and almost completely 

confined to the ocular row; abdominal tergites with 

more or less distinct patches of lighter scales on 

lateral margins and/or posterior corners (subg. Culi- 

ciomyia) ............................................................ 

--Head almost entirely clothed with broad, flat, mostly 

dark scales: all scales on abdominal tergites black- 

ish (subg. Eumelanomyia, mochthogenes group) ...... 

Pleurae somewhat shining, without grey dusting; no 

scales on mesepimeron ....................................... 

--Pleurae grey-dust'ed; mesepimeron scaled, with at 

least a few scales among hairs near base ............... 

Mesepimeron without a distinct scale-patch, usually 

with only very few scales among the hairs near base; 

propleuron with few or no scales ........................ 

16 

18 

20 

cinerellus 

19 

nebulosus 



Mesepimeron with a large patch of flat white scales 

extending over more than half its length; propleuron 

with a large patch of scales which just extends on to 

the upper corner of prosternum ........................... 

cinereus 

20--Hind femora indefinitely pale beneath, legs otherwise 

dark .............................................................. inconspicuosus 

w Hind femora (in $, as female is unknown) as dark 

as middle and front femora ............................... quintetti 

21--Tarsi, especially of hind legs, with narrow pale rings 

--Tarsi all dark .................................................. 

221 Middle tibia with a pale stripe antero-externally; postspiracular 

scales usually present ........................... 

Middle tibia without such pale stripe; no postspiracular 

scales ................................................................ 

23-- All tibiae and middle and hind femora with median 

pale anterior stripes running the whole length; hind 

tibia without an evident pale apical spot above but the 

anterior stripe reaches the tip; costal and subcostal 

veins with more or less numerous pale scales over the 

basal -2/3 of the wing ....................................... 

Femora and tibiae either unstriped or, if stripes are 

present on any of them, the hind tibia has a distinct 

pale apical spot and the anterior stripe does not reach 

the tip; pale scales on wing, if present, confined to 

base of costa ...................................................... 

22 

23 

duttoni 

() watti 

in part 

theileri 

24--Mesonotum with four spots of silvery-white scales; 

pre-alar scales present ........................ argenteopunctatus kingi 

() Some specimens of C. p. quinque[asciatus with lighter scales over hind 

tarsal joints may run out here in the key. The examination of the terminalia 

of the associated male is perhaps the best way to ascertain the identity of such 

palo females. 

38 Estud., Ens. e Doc.- 134 

24

RIBEIRO, H. & RAMOS, H. Cunha -- <strong>Research</strong> on the rnosquitoes of Angola m X 

--Mesonotum without silvery spots; with or without 

pre-alar scales ................................................... 

25 -- Pre-alar scales present .......................................... 

--No pre-alar scales ................................................ 

261 Costa (at least in female) with a small spot of pale 

scales at base; hind tibia usually with an anterior pale 

stripe, sometimes faint (often the other tibiae and middle 

femur also more or less distinctly striped) ............ univittatus 

--Wing scales all dark; hind tibia always dark, except for 

the conspicuous white spot at tip ........................... 27 

27 -- Postspiracular scales present; scutum with many brassy-yellow 

scales besides dark-brown ones, the former 

being abundant both on the anterior and posterior 

halves of scuturn ................................................ 

--No postspiracular scales; scales on the anterior 3 of 

scutum mostly brown .......................................... 

28--Abdominal sternites with narrow dark apical bands 

(usually extending forwards along the medium line 

of the segment); proboscis all dark ........................ 

25 

26 

29 

simpsoni 

28 

chorleyi 

in part 

--Sternites each with a median longitudinal dark stripe 

but without transverse bands; proboscis pale in middle 

beneath ............................................................. zombaensis 

29--Abdominal tergites dark or nearly so (without distinct 

pale bands or spots) ........................................... 

--Tergites with at least some distinct pale markings 

(bands and/or spots) .......................................... 

30- Abdominal tergites with pale basal bands, even if nar- 

row .................................................................. 

--At least some of the tergites without complete bands 

(though basal lateral pale spots present) .................. 

chorleyi 

in part 

Estud., Ens. e Doc. 134 39 

30 

31 

38

RIBEIRO, H. & RAMOS, H. Cunha -- <strong>Research</strong> o the mosquitoes o[ ,qngola X 

31- Venter mainly pale, unbanded (though median longitudinal 

dark markings may be present); proboscis pale 

beneath ............................................................. 

--Sternites with dark apical bands or almost completely 

dark; proboscis (in female) dark beneath ............... 

32- With 2-4 lower mesepimeral bristles () ................... 

--With only one lower mesepimeral bristle ............. 

'33--Tibiae each with a narrow pale ring at base ........... watti 

in part 

-- Tibiae dark at base or only slightly paler ... pipiens quinquefasciatus 

in part 

34--Upper fork-cell (in female) from 3.9 to 5.0 (mean 

about 4.4) times as long as its stem ........................ pipiens pipiens 

in part 

--Upper fork-cell (in female) from 2.1 to 3.7 (mean 

about 3.0) times as long as its stem ......... pipiens quinquefasciatus 

in part 

35--Scutal scales almost uniformly reddish-brown; male 

palpi all dark .................................................... 

--Scutal scales either almost uniformly dark brown 

(without obvious reddish tint) or with yellowish and 

brown scales intermixed; penultimate segment of male 

palpi with a line of white scales beneath .............. 

32 

35 

33 

34 

decens 

in part 

361 Pale spot at tip of hind tibia smaller, shorter than 

broad; distal - of anterior surface of hind femur 

dark ......................................................... trifilatus trifilatus 

--This spot on hind tibia more conspicuous, at least 

about as long as broad; usually, distal 1/5-1/3 (rarely, 

2/5) of anterior surface of hind femur dark ............. 37 

(D Among many hundreds of quinquefasciatus specimens from Luanda, a 

few males and females were found with 1-2, 1-3, 2-2 and even 2-3 lower mese- 

pimeral bristles. 

40 Eslud., Ens. e Doc.- 134 

36

RIBIglRO, H. & RAMOS, H. Cunha -- <strong>Research</strong> on the mosquitoes of Angola X 

37- Larger, wing-length 4.5-5.5 mm ............................ toroensis 

--Smaller, wing-length 3.5-4.5 mm ......................... chorleyi 

in part 

38 1 Mesepimeron bare of scales or almost so (a few scales 

may be present among the upper mesepimeral hairs, 

but none in middle of mesepimeron); thorax very pale, 

shoulders extensively so; 2-4 lower mesepimeral bristles 

................................................................... 

Mesepimeron with a distinct scale-patch in middle; 

thorax darker; with 1 or 2 lower mesepimeral brispruina 

tles ................................................................... 

39 

391 Antenna of female normal, each flagelar segment 

with only 4-6 verticillate hairs ............................. 

Antenna of female subplumose, first few flagelar 

segments each with 10-20 verticillate hairs ........... 

40--Venter pale, unbanded; halteres all yellowish; pro- 

boscis usually paler beneath ................................ 

--Sternites with at least narrow dark apical bands; knob 

of halteres dark; proboscis all dark ....................... 

41-- With broad lateral stripes of cream-coloured scales on 

abdominal tergites 6 and 7; hind tibia without any 

obvious pale spot at tip, though a few pale scales may 

be present; scales on propleura most of the colour of 

he integument, not white .................................... antennatus 

Tergites 6 and 7 without such lateral stripes; pale 

spot at tip of hind tibia usually evident; propleural 

scales whitish ................................................ pipiens pipiens 

in part 

421 Hind tibia with a pale spot at tip (this spot may be 

reduced and, in males, sometimes practically absent) 

--Hind tibia entirely dark ....................................... 

43- Venter with narrow apical bands, mainly pale; scales 

on anterior of scuturn almost uniformly brown, 


40 

49 

41 

42 

43 

47


without obvious reddish tint; penultimate segment of 

male palpi with a line of white scales beneath ......... 

--Venter with broad dark apical bands; either scutal 

scales mainly reddish-brown or scuturn with inter- 

mixed pale and brown scales; male palpi all dark ..... 

chorleyi 

in part 

44-- Hind femur with the apical - of the anterior 

surface dark; scuturn with a pair of spots of pale 

scales about middle; erect scales in middle of head 

yellowish ...................................................... ornatothoracis 

--Hind femur with only about /5-1/ of anterior sur- 

face dark; scuturn without spots or stripes of pale 

scales; erect scales of head all or nearly all dark 45 

45-- Scutal scales reddish-brown, without admixture of 

pale scales ........................................................ decens 

in part 

Scutal scales dark brown, without obvious reddish 

tint, sometimes some pale scales intermixed ........................ 46 

46-- Hind femur with barely distal 1/, of anterior sur- 

face dark ........................................................... invidiosus 

47 m 

48-- 

49-- 

Hind femur with fully 1/, anterior surface dark ...... trifoliatus 

Scales round prescutellar space dark; hind femur with 

the dark area at tip on anterior surface shorter 

than broad ....................................................... perfidiosus 

--Scales round prescutellar space pale; hind femur with 

the dark area at tip about as long as wide or longe 48 

Upper fork-cell scarcely more than twice as long 

as its stem ........................................................ 

Upper fork-cell 3-4 times as long as its stem ......... 

Hind tibia with a pale spot at tip; with two lower 

mesepimeral bristles ........................................... 

--Hind tibia all dark; usually, only one lower mese- 

pimeral bristle .................................................... 

44 

per/uscus 

telesilla 

grahami 

42 Estud.. Ens. e Doc.- 134 

50

RIBgIRO, H. & RAMOS, H. Cunha -- <strong>Research</strong> on the mosquitoes of Angola m X 

50-- Venter pale, unbanded (a few dark scales may be 

present on distal margins of sternites); proboscis 

pale beneath ................................................ weschei gediensis 

--Venter with dark apical bands; proboscis all dark 51 

511 Propleura with numerous scales; sternopleural scales 

more numerous, the two patches almost joined ... guiarti sudanicus 

--Propleura with few scales; the two sternopleural 

scale-patches smaller and well separated .................. ingrarni 

Key to Fourth Instar Larvae () 

1- Mouth brushes modified (predacity) ....................... tigripes 

--Mouth brushes unmodified .................................. 2 

2--Siphon with prominent apressed spines near apex ... 

--No such spines present ..................................... 

--Siphon with a bend towards the dorsal side near 

apex; 2 pairs of single or bifid subventral tufts .... 

Siphon straight; 3-4 pairs of subventral tufts near 

always with 2 or more branches ......................... 

--Spines near apex in a ventral group only; siphon in- 

dex 8-14 (2) ....................................................... 

--Spines either in a dorsal group only or both dorsal 

and ventral; index 6-8 ........................................ 

--Siphon swollen and strongly bilaterally convex to- 

wards middle ..................................................... 

--Siphon not strongly swollen towards middle, at most 

unilaterally convex ............................................ 

toroensis 

ingrami 

chorleyi 

(D The larvae of Culex (C.) ornatothoracis, Culex (E.) quintetti, Culex (C.) 

toroensis macrophyllus and Culex (C.) watti were not described. 

(2) All the figures of siphon indices in the key are in accordance with our 

mounted material. 


RT}ImO, H. & R^M0s, H. Cunha -- <strong>Research</strong> o the mosquitoes of Angola X 

6--Subventral tufts of siphon represented by long, single 

setae; siphon not spiculate .......................................... dutttoni 

() watti 

--These tufts strongly branched; siphon obviously spi- 

culate, at least in neighbourhood of pecten ........... 7 

__ Tufts of siphon (excepting the more distal pair) 6-10 

branched, with a mean number of branches per tuft 

of about 8.5; head seta d with 4 or 5 branches; spi- 

culation of siphon confined to the ventral side, in 

neighbourhood of pecten ................................. pruina pruina 

These tufts 9-16 branched, with a mean number of 

branches per tuft of about 12.5; head seta d with 

about 7 (6-9) branches; spiculation extending to the 

lateral and dorsal aspects of siphon, usually covering 

most of its surface ....................................... pruina eschirasi 

8--Pecten extending to 3/, or beyond ........................ 

--Pecten not extending beyond , seldom beyond 

--Comb composed of spines; spine on dorsal valve of 

siphon exceptionally long (longer than diameter 

of siphon apex), conspicuously denticulate towards 

base ...................................................... weschei ? gediensis 

Comb composed of scales; spine on dorsal valve of 

siphon small and simple, as usual ............... trifilatus trifilatus 

10q Mentum an equilateral straight-edged triangle with 

very numerous small teeth which are not visible 

under low magnifications ................................... 

Mentum not as above, obviously toothed .............. 

ill Seta } (abd. seg. w) with about 5 (4-7) branches, 

seta with 3-5 branches and seta with 4 or 5 

branches each side, making a total of about 25 (22-28) 

branches on both sides together ............... aurantapex jinjaensis 

Seta } usually with 2-3 (rarely, 4 or 5) branches, 

(x) According to Hopkins (24), apparently indistinguishable from duttoni. 

44 Estud., Ens. ßDoc.- 134 

9 

10 

11 

13

RIBgIRO, H. & RAMOS, H. Cunha -- <strong>Research</strong> o the mosquitoes of /tngola X 

seta with 2 (1-3) branches and seta with about 

3 (2-4) branches each side, making a total of about 16 

(10-22) branches on both sides together ........................... 12 

12 1 Head seta e nearly always with 2-3 branches, very 

rarely 4-branched ......................................... bitaeniorhynchus 

ethiopicus 

--Head seta e usually (in about 75 per cent of in- 

stances) with 4 or more slightly longer branches ...... annulioris 

13-- Head setae B and C both much shorter than head, 

with 2-3 and 2 branches, respectively; siphon with 

index about 5-6; subventral tufts sparsely plumose, 

longest tufts more than twice diameter of siphon ... inconspicuosus 

Not with this combination of characters ............. 14 

Comb composed wholly or partly of spines or spine- 

like scales ........................................................... 15 

Comb composed entirely of scales ....................... 22 

15-- Siphonal index about 4; siphon slightly bent upwards 

near apex; preclipeal spines short (shorter then half 

distance between bases), thick and blackish ........ 

Index about 4 or more; siphon not bent upwards near 

apex; preclipeals longer and thinner, usually also 

paler ................................................................ 

poicilipes 

161 Spine on dorsal valve of siphon exceptionally long 

(longer than diameter of siphon apex), conspicuously 

denticulate towards base ..................... argenteopunctatus kingi 

This spine short and inconspicuous, not denticulate 17 

17--Comb composed of spines, with no admission of 

scales ............................................................... 

--Comb composed partly of scales ......................... 

181 Siphon very long, index at least 10; subventral tufts 

single or double and minute, usually much shorter 

than diameter of siphon .................................... 

--Siphonal index not over 8-9; subventral tufts at 

16 

18 

20 

grahami 


RIBgIRO, H. & RAMOS, H. Cunha -- <strong>Research</strong> on the mosquitoes o[ Angola -- X 

least bifid (with 2-8 branches) and longer (though 

either longer or shorter than diameter of siphon) 

19-- Comb an irregular row of 5-9 spines; siphon index 

about 7 and over .......................................... guiarti sudanicus 

Comb a patch of about 30 spines; siphon index at 

most about 5 or 6 ............................................ theileri 

20-- Head setae B and C with 2-3 and 3-5 branches 

respectively ....................................................... zombaensis 

These setae with at most 2 branches ..................... 21 

21 I Subventral tufts of siphon usually bifid and not 

longer than siphon breadth; head seta [ with 4-10 

branches, usually 7 or 8-branched; comb of about 20 

(13-28) teeth ................................................ simpsoni form A 

Subventral tufts of siphon usually with about 4 

branches and longer than siphon breadth; head 

seta ! with 3-6 branches; comb of about 30 (26-36) 

teeth ......................................................... simpsoni form B 

22--Head seta B or C (or both) single () ................ 

--Each of these setae with at least 2 branches ........ 

23m Subventral tufts with 5-10 branches which are 

obviously longer than diameter of siphon; upper 

caudal seta with 4-5 branches .............................. salisburiensis 

Subventral tufts with 2-4 branches which are, at 

most, about as long as diameter of siphon; upper 

caudal seta single .............................................. horridus 

in part 

24--Siphon index usually about 4 or less, but if about 

5-6 then head setae B and C both with at least 4 

branches ........................................................... 

--Siphon index usually about 7 or more, but if about 

5-6 then head seta B (at least) only bifid or trifid .... 

0) Head setae A, B and C with 7/7, 1/2, and 1/1 branches, respectively, 

in the sole known Angolan larva of C. horridus. 

46 Estud., Ens. e Doc.- 134 

19 

23 

24 

25 

30

RIBEIR0, H. & RAMOS, H. Cunha -- <strong>Research</strong> on the mosquitoes o! Angola X 

25m Antenna less than half length of head; lateral seta 

of saddle distinctly frayed, more than twice length 

of saddle; anal papillae about 3-4 times length of 

saddle; siphonal index 3 or less; subventral tufts ex- 

tending to near base or siphon, at least to its basal 

fifth ................................................................ 

--Antenna at least half length of head; lateral seta 

not frayed and shorter, at most 1 times length 

of saddle; anal papillae usually shorter; siphon index 

usually more than 3; basal subventral tufts of siphon 

more distally placed ........................................... 

26 1 Antenna spiculate; comb of about 30 scales ........... 

--Antenna without spicules; comb of not more than 

18 scales ........................................................... 

27 1 Anal papillae short and rounded, dorsal pair at most 

about as long as saddle, usually less; upper caudal 

seta with at least 3 branches; preclipeals short and 

thick, with somewhat blunt tips .......................... 

Anal papillae narrowly triangular, dorsal pair usually 

longer than saddle; upper caudal seta usually with 

two branches, rarely single or 3-branched; preclipeals 

longer and thinner, as usual ............................... 

Head setae B and C with 2 or 3 branches, respec- 

tively; comb of about 30 scales; subventral tufts 

of siphon placed very close to mid-ventral line ...... 

--Both head setae B and C with at least 4 branches; 

comb with 40-60 scales; subventral tufts much more 

lateral in position .............................................. 

26 

27 

nebulosus 

cinereus 

thalassius 

28 

perfidiosus 

291 Proximal two pairs of subventral tufts of siphon 

with a mean number of branches per tuft of about 

2.5 (1.5-3.0); siphon often with a slight sigmoidal 

curvature ..................................................... pipiens pipiens 

--Proximal two pairs of subventral tufts of siphon 

with a mean number of branches per tuft of 

Estud., Ens. e Doc.- 134 47 

29

RIBgIRO, H. & R.s, MO8, H. Cunha -- <strong>Research</strong> on the mosquitoes of Angola X 

301 

about 8.5 (5.0-11.25); siphon without sigmoidal curvature 

................................................ pipiens quinquefasciatus 

Siphon very dark brown ..................................... horridus 

in part 

Siphon from pale brown to nearly colourless ........ 31 

31-- Pecten spines bifid at tip, with one large and some- 

times one small basal denticles .......................... 

321 

331 

34-- 

351 

Pecten spines not bifid at tip, often with more than 

two basal denticles .......................................... 

Usually, with at least some of the subventral tufts 

about 1 times the diameter of siphon at point 

of attachment, but if not so then preclipeal spines 

stout and dark brown ....................................... 

Subventral tufts smaller, at most about as long as 

diameter of siphon, usually shorter; preclipeals slen- 

der and paler .................................................... 

cinerellus 

Ventral brush without unpaired tufts outside the 

barred area; subventral tufts of siphon usually with 

3-4 (rarely, 2) branches; head seta C 3-branched; 

comb a patch of 30-45 scales ..................... tritaeniorhynchus 

Ventral brush with unpaired tufts outside the barred 

area; subventral tufts of siphon single or bifid, rarely 

3-branched; comb of about 50-60 scales ............. 34 

Siphonal index about 7-9; antenna infuscated for its 

full length; upper caudal seta 3 or 4-branched, with 

one long branch and usually other 3 much shorter 

ones of which 2 are dorsal and the other ventral 

to the main branch .......................................... wigglesworthi 

Siphonal index from 10.5 to 16; antenna infuscated 

at base and on about the apical 2/.; upper caudal 

seta with 2 branches, 'the short one dorsal to the 

main branch, as usual . ..................................... kingianus 

ß 

Head setae B and C only about half length of head, 

sparsely plumose; head seta d with 4 or 5 branches 

32 

33 

35 

rubinotus 


RIBEIRO, H. & RAMOS, H. Cunha -- <strong>Research</strong> on the mosquitoes of Angola X 

--Head setae B and C about as long as head, obviously 

plumose; seta d single or bifid ............................. 

361 Pecten spines with 4-6 fairly evenly spaced ventral 

denticles of about the same size which reach to near 

the tip; head seta d bifid beyond the middle in the 

sole described larva ............................................ 

--Denticles of pecten spines often less in number and 

always irregular in size and position and not extend- 

ing along the whole ventral side of the spine; seta 

d single ........................................................... 

37 1 None of the pecten spines with more than 2 basal 

denticles; antenna infuscated only beyond tuft ...... 

--Usually, some of the pecten spines with 3 or more 

basal dentic]es, but if none has more than 2 such 

denticles then antenna infuscated both at base and 

beyond tuft ........................................................ 

36 

insignis 

37 

perfuscus 

381 Lateral seta of saddle minute, with 4-6 branches; 

the most proximal subventral tuft of siphon proximal 

to the most distal pecten spine; comb with about 

25 scales .......................................................... () telesilla 

--Lateral seta at most 3-branched (occasionally with 4 

branches) about 1/,-t/2 length of saddle; usually all 

subventral tufts placed distally to the distal end of 

pecten; comb with about 40 or more scales ............ 39 

39--Siphon index about 10 or more ........................... 

--Siphon index about 9 or less ............................. 

Siphon index about 8 or 9; lateral seta of saddle 

single or double ................................................. 

38 

decens 

in part 

40 

decens 

in part 

invidiosus 

in part 

(x) On about 40 per cent of the Angolan larvae of telesilla none of the pectcn 

spines had more than 2 basal dentitles. 


RIBgIRO, H. & RAMOS, H. Cunha -- <strong>Research</strong> on the mosquitoes of Angola X 

--Siphon index about 5-7; lateral seta of saddle single 

or with up to 6 branches ................................... 

41--Lateral seta of saddle single ................................ 

--Lateral seta of saddle with 2-6 branches ................ 

41 

invidiosus 

in part 

42 

42--Head seta C with 4 branches .............................. univittatus 

in part 

--Head seta C with 2-3 branches ............................ 43 

43- Preclipeal spines finely drawn-out on the apical -; 

pecten spines usually with 1-3 basal denticles (occasionally 

4 or 5), often most of them with only 2 such 

denticles ............................................................ antennatus 

--Preclipeals somewhat stouter, not finely drawn-out 

appically: pecten spines usually with 2-6 basal denti- 

cles, but if none wit 4 or more then at least most of 

the pecten spines with 3 such denticles .................. 44 

44 1 Usually, none of pecten spines with less than 3 basal 

denticles; head setae A and C with 6-8 (usually 6 or 

7) and 2 branches, respectively; lateral seta of saddle 

2-branche .......................................................... 

Usually, some of pecten spines with 2 basal denticles; 

head setae A and C with 6-9 and 2 or 3 branches, respectively; 

lateral seta of saddle with 2-6 branches ... 

45q Head seta A with 6-7 branches; lateral seta of saddle 

usually 2-branched, sometimes trifid; anal papillae 

greatly variable in length but ventral pair only about 

ß /5-4/. of dorsal pair ......................................... 

--Head seta A with 7-9 branches; lateral seta of saddle 

usually with 3 branches; anal papillae about as long 

as saddle, dorsal pair only slightly longer than ventral 

pair .................................................................. 

trifoliatus 

45 

invidiosus 

in part 

univittatus 

in part 


7 -- SYSTEMATIC ACCOUNT 

Genus Culex L. is a cosmopolitan one, comprising near 600 described 

species and subspecies, about 150 of which occur in the Ethiopian 

Region. 

After the recent revisions by Sirivanakarn (29) (30), the Ethiopian 

species are thought to belong to seven subgenera: Lutzia Theo., Lasiosiphon 

Kirkpatrick, Barraudius Edws., Eumelanomyia Edws., Maillotia 

Theo, Culiciomyia Theo, and Culex L. Of these subgenera, Lasiosiphon 

and Barraudius are represented only by one and two species, respectively, 

confined to the northernmost part of the Ethiopian Region (Sudanese 

Arid biotic zone of Moreau). All the remainder five subgenera are widely 

distributed in Angola as well as in all the Africa South of Sahara. 

Basic descriptions of the Angolan Culex are to be found mainly in 

F. V. Theobald (6), F. W. Edwards (8), G. H. E. Hopkins (24), Galliard (31), 

De Meillon & Lavoipierre (23), De Meillon et al. (28), De Meillon (33) 

and Brunhes et al. (14), though other taxonomic contributions will be 

cited under the corresponding taxa. 

7.1--Subgenus I_,utzia Theobald, 190'3 

This subgenus with strong, predacious larvae is represented in Angola, 

as well as in all the Ethiopian Region, by a single generalized species. 

7.1.1- Culex (Lutzia) tigripes Grandpr6 & Charmoy, 1900 

The first published record of C. tigripes from Angola was that of M. 

Gamble (7), in 1914, from S. Salvador. F-rther records of this species 

Estud., Ens. e Doc. 134 51


were those by A. F. Gfindara (10) (11) from Caconda, Capelongo, Cuando, 

Dala Tando, Ganda, Luso and Huambo, by C. B. Worth & H. E. Paterson 

(12) from Cabinda, Caxito and Dundo and by one of us (34) from 

Lobito. 

MATERIAL EXAMINED. BAIA FARTA, 3 q plus larvae, 12.IX.1964; 

BIBALA, 16 larvae, 2.V.1969; BOCOIO, 1 2, 2 larvae, 8-9.X.1970; CABUTA, 

1 larva, 11.VI.1970; CALAI, 7 2, 9 /5, 21 larvae, 16-26.XI.1965, 23-25.V.1967 and 

9.VI.1967; CAPUNDA, 1 larva, 16.VI.1970; CARACULO, 1 2, 2 larvae, 14.V. 1969; 

CARIANGO, 1 larva, 16.X.1969; CARLAONGO, 1 2 plus 1 larva, 31.X.1969; 

CHAO DA CHELA, 2 larvae, 29.V.1969; CHIEDE, 3 larvae, 21.IV. 1971; CUAN- 

GAR, 1 $, 3 /5, 8 larvae, 10.IV. 1965 and 4.V.1966; CUANZA RIVER, I larva, 

12.VI.1970; CUBAL, 6 larvae, 12-14.IX.1970; DINDE, 3 2,2 /5, 2 larvae, 28.V.1971; 

EBANGA, 1 9, 4 larvae, 17-19.IX.1970; EQUIMINA, 1 $, 4 larvae, 21-22.V.1970; 

HUAMBO (=N. LISBOA), 5 /5, 30.XI.1966; IMPULO, 3 larvae, 23.111.1970; 

LANDANA, 1 , 2 /5, 7 larvae, 24-26.VIII. 1970; LUANDA, 1 , 17.111.1971; 

LUCALA RIVER, 1 larva, 25.VIII.1969; LUELE RIVER, 2 larvae, 13.IV. 1970; 

MALANJE, 2 larvae, 12.IV. 1971; MARCO DE CANAVEZES, 2 larvae, 12.IX.1970; 

MO(AMEDES, 8 $, 5 /5, 20 larvae and pupae, 18-26.X.1967 and 30.IV.1969; 

MONA QUIMBUNDO, 2 larvae, 10.IV.1970; M'PUPA, 1 /5, 2 larvae, 9.V. 1966; 

MUNHINO, 1 /5, 14 larvae, 7.V.1969; MUSSULO, 1 $, 1 /5, plus a few larvae, 

24.111.1971; ONCOCUA, 1 larva, 2.1111970; PANDA, 4 larvae, 13.IV. 1965; QUELA, 

1 larva, 29.1.1970; QUILENGUES, 1 larva, 27.V. 1971; SA DA BANDEIRA 

(20 km W), 1 larva, 6.VI.1969; SAURIMO, 6 larvae, 10-12.1V.1970; SERPA PINTO, 

2 larvae, 24.IV. 1971; SERRA DA NEVE, 1 larva, 29.V. 1970; TANDO ZINZE, 

1 $, 3 /5, 1 larva, 19.VIII. 1970; TCHIVINGUIRO, 4 larvae, 29.V.1969; UiGE, 

2 $, 2 larvae, 15.IV. 1971 and 25.IX.1969; VALE DO LOGE, 2 larvae, 26.IX.1969, 

all new locality records. Also DALA TANDO, 2 $, 1 /5, 2 larvae, 23-28.VIII. 1969; 

DUNDO, 13 larvae, 19.VIII. 1969, and HUAMBO, 12 $, 18 /5, 60 larvae, 

24.1.1964, 30.V-16.VI.1966, 3.XII. 1966, 28.I.1969 and 4-19.V.1971. 

TAXONOMIC NOTES. C. tigripes shows a marked variation through 

its range in Angola. Howewer, the comparison of our material, both of 

adults and larvae, with the series in the British Museum (Nat. Hist.) 

confirmed the infrasubspecific nature of this variation. 

DISTRIBUTION. As it can be seen on Map 4, C. tigripes is a fairly 

generalized species also in Angola. 

BIOEOOLOGICAL NOTES. Larval tigripes was recorded from 104 

different breeding places, 64 (61.5 per cent) of which were ground water, 

28 (27 per cent) were artificial containers and 12 (11,5 per cent) were 



rock-pools. Ground water biotopes included ground pools (31), river margins 

(14), irrigation drains (7), shallow earth-wells (6), pools in river beds 

(4), a pond and a crabhole. Artificial containers were represented by old 

rubber tyres (10), barrels (8), cement water-tanks (8) and drums (2). The 

nature of the breeding area of C. tigripes as shown by these data, points 

to an appreciable man-dependance of this species for breending. 

The breeding water was either limpid or turbid, sometimes even foul 

water with decaying vegetables (, sisal, fallen leaves, etc.). 

Living macroscopic aquatic vegetation was either present, sometimes 

even abundant, or completely absent. !n seYeral instances, the presence 

of filamentous algae was recorded. The pH of the breeding water measured 

in 50 different breending places, ranged from 5.0 to 8.0, though it 

became evident that C. tigripes shows a marked preference for acid 

waters, these representing 88 per cent of the total. The salt content 

(NaCl) of the water, deterre:reed in 16 biotopes, ranged from 0.1 to 

4.15 g/1 (mean 1.1 g/l), salinity being an important limiting factor for 

breeding of tigripes. 

Larvae of C. tigripes are voracious predators on other mosquito 

larvae and we observed a full grown tigripes larva devoring three third 

or fourth instar larvae of C. pipiens at intervals of about 15 minutes. 

It seems that, in fact, the presence of prey mosquito larvae is a decisive 

factor for breeding of tigripes. 

As to the associated mosquitoes, larvae of other species of Culex were 

present in 75 out of the 104 breeding places of tigripes, while Anopheles 

spp. occurred in 37 of these, Uranotaenia spp. in 10, Aedes spp. in 6, 

Ficalbia (F. mimomyial'ormis) in 3, and Aedomyia (Aed. [ur/urea) in 

only one. 52 species of associated mosquitoes were recorded from the 

Angolan larval biotopes of tigripes, belonging to genera Culex (22 species), 

Anopheles (19), Aedes (5), Uranotaenia (4), Ficalbia (1) and Aedemyia 

(1). The species most frequently associated with larval tigripes were, 

by decreasing order of frequency: C. duttoni and C. univittatus (each 

one recorded from 20 larval biotopes), An. coustani (18), C. p. piplens 

(11), C. p. quinque/asciatus and C. decens (9), An. gambiae (7), An. demeilloni, 

An. maculipalpis, C. annulioris, C. simpsoni and U. fusca (6), 

An. pharoensis, C antennatus and C. telesilla (5), An. cydippis, An. rhodesiensis, 

C. ethiopicus and C. poicilipes (4) and Ae. yangambiensis, An. 

r. rufipes, C. chorleyi, C. cinerellus, C. invidiosus, C. theileri and F. mimomyiaformis 

(3). 

Estud., Errs. e Doc.- 134 53

RIBEIRO, H. & RAMOS, H. Cunha -- <strong>Research</strong> on the mosquitoes o[ .4ngola -- X 

Figs. 1, 3, 6 and 12 show typical breeding places of C. tigripes. 

The association of larval tigripes with other predatory larvae of any 

of the Angolan species of Eretmapodites, Mucidus or Toxorynchites was 

never observed by the writers, pointing to competition as other important 

limiting factor for breeding of tigripes. It is also to be noted that in 10 

out of the 104 recorded larval biotopes of tigripes no larvae of any other 

mosquito were found, which seems to show that in an appreciable percentage 

of breeding places pupation of tigripes is attained at about the 

same time as the exhaustion of the associated prey larvae of other mosquitoes. 

Only 6 - 5 were caught as adults, the remainder of our adult 

material being laboratory-reared. All the males were caught at rest inside 

a hen-house, at about 7.30 a.m. Two of the females were taken outdoors 

in the afternoon, one of them resting on the inside walls of a drum, in 

peridomestic habitat, and the other at rest on plants in a public garden. 

The remaining four females were caught early morning (6.30-7.30) 

while resting indoors, on the walls of human habitation, two of them 

engorged. It is possible that these two females had fed on human blood, 

as it was also recorded by Haddow et al. (35) in Uganda. However, tigripes 

was never seen by the writers landing on man and there is a general 

agreement as to the marked zoophilic preferences of this species (36) (37). 

7.2 -- Subgenus Maillotia Theobald, 1907 

The World Catalog by Knight & Stone (23) lists only 9 species and 

2 subspecies under subgenus Maillotia endemic either of the Ethiopian 

Region or of the Mediterranean Subregion (of the Palaearctic). 

Of the 6 Ethiopian species, 2 are restricted to Madagascar and 3 

other species occurring in the mainland are endemic to the southernmost 

part of the East and South African Subregion. The only now known 

Angolan species of Mallotia is also the only widely distributed Ethiopian 

species of the subgenus. 

7.2.1--Culex (Maillotia) salisburiensis salisburiensis Theobald, 1901 

New record for Angola of the subgenus, species and subspecies. 



MATERIAL EXAMINED. BOCOIO, 1 9, 4 larvae, 7-10.IX.1970; EBANGA, 

1 9, 21 larvae, 17-19.IX.1970. 

TAXJONOMIC NOTES. Our material is in quite good agreement with 

the descriptions by Theobald (6), Edwards (8), Knight (38), and Hopkins 

(24). In particular, larvae show that Angolan salisburiensis is of the 

type form and not of ssp. naudeanus of Muspratt (39), as: (a) the comb 

consists only of scales; (b) the pecten teeth are straight and have 2-4 

basal denticles; (c) head seta C is much more delicate than B and is only 

about half the length of this one. 

DISTRIBUTION AND BI. OECOLOGICAL NOTES. Map 5 shows the only 

two known Angolan localities for C. salisburiensis. However, as salisburiensis 

seems to bc the most generalized Ethiopian member of the subgenus, 

it is expected to bc widely distributed also in Angola. 

Our females wcrc reared from larvae. Eight larval biotopcs of C. 

salisburiensis wcrc recorded by the writers. In six instances, larvae bred 

along the shallow margins of small rivers, while the other two breeding 

places wcrc rock-pools at river edges. The breeding water was always 

limpid and with aquatic vegetation, usually emergent. The pH of the 

water, measured in six instances, ranged from 5.4 to 6.4 (mcan, 5.8). 

C. annulioris, An. coustani, An. demeilloni and An. longipalpis were 

the most frequently associated mosquitoes, other Anophelines (ardensis, 

[unestus and ru[ipes) and Culex (decens, ethiopicus, p. pipiens, tigripes 

and univittatus) being also recorded from the breeding places of salisburiensis. 

C. salisburiensis seems to be a zoofilic mosquito as it was never 

caught by us biting man nor inside human habitations. 

7.3 -- Subgenus Eumelanomyia Theobald, 190'91 

Adopting the viewpoints of Sirivanakarn (29) (30), the new edition of 

the World Catalog by Knight & Stone (23) lists 64 species under subgenus 

Eumelanomyia Theo., most of them Ethiopian, but also Oriental and a 

few Australasian. Edwards' subgenus Mochthogenes is now considered 

as no more than a group of species within subgenus Eumelanomyia Theo., 

while Neoculex Dyar is now thought to be mainly Australasian and also 



Neotropical and Holarctic, but without any Ethiopian representatives. 

The Ethiopian species formaly belonging to Neoculex Dyar are now 

included in one of the subgenera E,melanomyia Theo. or Maillotia Theo., 

both resurrected from synonymy under Neoculex. 

Ethiopian species of Eumelanomyia are often relatively localized 

forms, though the now known 8 Angolan representatives are among the 

most widely distributed species of the subgenus in Africa South of Sahara. 

Yet, some of these may prove to be subspecifically distinct (see horridus, 

inconspicuosus and rubinotus). 

7.3.1- Culex (Eumelanomyia) horridus Edwards, 1922 

C. horridus was recorded for the first time from Angola (Sambo) by 

Colago (9), in 1952. The only other known Angolan records was that by 

Gfindara (10) (11), in 1956, from Cuchi. 

MATERIAL EXAMINED. SAURIMO (= H. de Carvalho), I larva, 

11.IV. 1971. 

TAXONOMIC NOTES. Appreciable differences concerning head chetotaxy 

were found between our larva and the description given by Hopkins 

(24). In the Angolan larva head chetotaxy is as follows: antennal 

tuft with 10 branches; seta A with 7 branches; seta B single or bibid; 

seta C single; d single; e 2 or 3-branched; f with 5 or 6 branches. 

It seems, thus, that our larva is nearer to Zambian larvae, as described 

by Robinson (in Hopkins, op. cit., footnote p. 265), horridus being 

probably a polpytypic species. Unfortunately, no Angolan adults are available 

for comparison with other adult local horridus populations (namely, 

from Accra, the type locality) nor the larva of horridus var. rageaui 

Hamon & Rickenbach (40) was yet described. 

DISTRIBUTION AND BIOECOLOGICAL NOTES. C. horridus seems to be 

a fairly generalized though relatively uncommon Ethiopian species. In 

Angola, the only known locality records of horridus belong to the Rbodesian 

Highland Zone (cf. Maps 3 and 5). 

Our horridus larva bred in a discarded tin of sardines filled with 

turbid rain-water, in association with larval Ae. aegypti, C. nebulosus 

.56 Estud., Ens. e Doc.- 134

RIBEIR0, H. & RAMOS, H. Cunha -- <strong>Research</strong> on the mosquitoes of Angola -- X 

and E. chrisogaster. This record is in accordance with the known pref- 

erence of larval horridus for small breeding places, including artificial 

containers. 

Very little is known about the bioecology of adult horridus (35) (36) (37) 

though it seems to be mainly a zoophilic mosquito also in Angola. 

7.3.2- Culex (Eumelanomyia) inconspicuosus (Theobald, 1908) 

The only Angolan published record of C. inconspicuosus is that by 

Gndara (10) (11), from Caconda, in 1956. 

MATERIAL EXAMINED. CANGANDALA (24 km SE), 2 larvae, 10.11.1970, 

CARIANGO, 2 larvae, 18.X.1969; DALA TANDO, 3 larvae, 22.VIII. 1969 and 

12.IV. 1970; DUNDO, 4 larvae, 19.VIII. 1969; EBANGA, 1 3, 9 larvae, 18.IX.1970; 

GUNGO, 1 larva, 21.XI.1969; HUAMBO, 1 larva, 12.IV.1970; INHUCA RIVER, 

2 larvae, 20.VIII. 1970; LUCALA, 2 larvae, 27.VIII. 1969; MUMBONDO, 7 larvae, 

8.V.1970; MUXIMA, 5 larvae, 30.IV-1.V.1970; SANZA POMBO, 1 larva, 24.IX. 1969; 

UIGE, 3 larvae, 25.IX.1969; VILA NOVA DO SELES, 4 $, 3 3, 4 larvae, 

20.XI. 1969. 

TAX;ONOMIC NOTES. Both the external morphology and terminalic 

characters of the Angolan adult inconspicuosus agree quite well with 

the descriptions by Theobald (6), Edwards (8) and Brunhes et al. (14). 

As to larvae, those from Cariango, Dundo, Ebanga, Gungo, Inhuca 

River, Mumbondo, Muxima, Uige and Vila Nova do Seles are also 

in good agreement with the descriptions given by De Meillon et al. (28) 

and Hopkins (24). It is to be noted, however, that the comb would 

be better described as consisting of scales, spinelike scales and spines 

altogether. 

Larvae from Cangandala, Dala Tando, Huambo, Lucala and 

Sanza Pombo, on the other hand, show some non-negligible disagreements 

with the above descriptions as well as with that given by 

Galliard (31) for C. nyangae. In any way, as these larvae are very near 

to inconspicuosus and considering the unfortunate absence of associated 

adults, they are only tentatively identified to C. inconspicuosus. 

DISTRIBUTION. Map 6 shows the known distribution in Angola 

of C. inconspicuosus, which seems to be mainly a species of the West 

African biomes (cf. with Map 3). 



BIOECOLOGICAL NOTES. All our adult specimens of C. inconspicuosus 

were laboratory-reared. 

16 larval biotopes were recorded for inconspicuosus, consisting of 

river margins (16), small ground pools (4), rock pools (3), small 

streams (2) and one large permanent pool. Aquatic vegetation was abundant 

in larger breeding places while fallen leaves were usually present 

in smaller ones. As a rule, appreciable additional shade was provided by 

surrounding vegetation. The breeding water was usually clean, with 

a pH, measured in five instances, of 5.4-6.9 (mean 5.9). 

The species of mosquito larvae associated with C. inconspicuosus 

in the 16 breeding places for this species were, by decreasing order 

of frequency: An. coustani (in 17 of them), An. demeilloni (6), F. mimomyaformis 

(4), An. funestus (3), An. obscurus, C. cinerellus, C. ingrami, 

C. perfidiosus, and U. mashonaensis (2, each) and, recorded 

only once, An. cinctus, An. gambiae, An. nili Congo form, An. 

rivulorum, An. r. rufipes, C. kingianus, C. simpsoni, C. tigripes, C. univittatus, 

C. weschei gediensis and U. balfouri. 

Fig. 2 shows a larval biotope of C. inconspicuosus. 

C. inconspicuosus is a widely distributed and relatively common 

species in Angola. It seems to be a zoophilic mosquito, as it was never 

seen by the writers either landing on man or resting inside human habitation. 

C. inconspicuosus was also reported by Hamon (37) as not bitting 

man in the Bobo Dioulasso area. 

7.3.3--Culex (Eumelanomyia) insignis (Carter, 1911) 

C. insignis was previously recorded from Angola (Maquela do 

Zombo) only by Worth & Paterson (12), in 1961. 

MATERIAL EXAMINED. SAURIMO (= Henrique de Carvalho), 1 9, 2 5, 

l 0.IV. 1970. 

TAXONOMIC NOTES. Both the external and terminalic characters 

of our adults agree with known descriptions by Carter (41), Edwards (8) 

and Hamon & Rickenbach (40). 

D1STRIBUTI,ON AND BIOECOLOGICAL NOTES. Map 5 shows the only 

two Angolan localities recorded for insignis. However, in spite of being 



a relatively rare mosquito, C. insignis is probably widely distributed 

also in Angola. 

The adults in collection were caught at rest inside the tents of 

our camp. C. insignis was never seen landing on man, which is in 

accordance with its known zoophily. 

7.3.4--Culex (Eumelanomyia) kingianus Edwards, 1922 

New record for Angola. 

MATERIAL EXAMINED. DALA TANDO, 2 larvae, 23.VIII. 1969; HUAMBO, 

3, 12 larvae 14.IV.1970 and 23.VII-1.VIII. 1966. 

TAXONOMIC NOTES. Our male agrees with the descriptions of 

kingianus given by Edwards (8) and by Mattingly & Lips (42). 

The larva tentatively attributed by Hopkins (24) to this species 

and described by him (pp. 255-6) seems not to be, in fact, that of 

kingianus. Anyhow, our larvae are in good accordance with the de- 

scription given by Mattingly (42). In particular, it is to be noted that 

all comb teeth on abdominal segment VIII are clearly scales and not 

spines. 

DISTRIBUTION AND BIOECOLOGICAL NOTES. Map 5 shows the only 

two Angolan localities recorded for C. kingJanus. 

Our male was reared from larva. All three larval biotopes recorded 

for kingianus consisted in ground water. Two of them were small 

pools at river margins with aquatic vegetation and moderately high 

organic matter content. In the other instance, larvae bred in the 

shallow margins of a small river with emergent and submerged 

aquatic vegetation. The pH of the breeding water, measured in two 

of the breeding places, was 5.6 and 6.4. 

As to the associated species in the larval biotopes of kingianus, 

C. duttoni and C. tigripes were recorded from two of these. An. demeilloni, 

C. cinerellus, C. inconspicuosus, C. p. pipiens, C. p. quinquefasciatus, 

C. toroensis, C. wigglesworthi and U. mashonaensis were also 

recorded as associated larvae in one instance. 

Fig. 3 illustrates a breeding place of C. kingianus. 

Estucl., Ens. e Doc.- 134 59


C. kingianus seems to be also a relatively uncommon zoophilic 

mosquito (35). 

7.3.5--Culex (Eumelanomyia) quirtterri Brunhes, Adam & Bailly- 

Choumara, 1967 

C. quintetti (male) was described by Brunhes e! al. (14) from Man, 

Ivory Coast, and recorded in the same paper also from Dundo, Northeastern 

Angola (Map 6), on the basis of specimens submitted by E. 

Abonnenc (op. cit., p. 49). 

Dr. A. de Barros Machado kindly informs us that Dundo specimens 

were almost certainly caught by means of one of his New Jersey lighttraps 

installed in the gallery forest of the Luachimo River, the specimens 

being sent to E. Abonrenc included in sandfly catcites. 

No specimens of C. quirtterri were caught by the present writers. 

7.3.6--Culex (Eumelanomyia) rima Theobald, 1901 

The only known Angolan record of C. rima is that of Gamble (7), 

in 1914, from S. Salvador (Map 5). 

No rima specimens were caught by the writers. 

7.3.7--Culex (Eumelanomyia) rubinotus Theobald, 1906 

C. rubinotus was previously recorded only from Cuchi by Grin- 

dara (10) (!1), in 1956. 

MATERIAL EXAMINED. CABINDA, 1 9, 23.VIII. 1970; LUANDO RIVER, 

29 9, 16.VI.1970. 

TAX!ONOMIC NOTES. All the 29 females caught at Luando River 

agree satisfactorily with the known descriptions by Theobald (6) and 

by Edwards (8). The Cabinda female, however, is here only tentatively 

attributed to rubinotus as it shows some non-negligible differences 

which cannot be properly evaluated in the absence of males from the 

same area. In any case, it will be readily keyed out with rubinotus 

if the above proposed key is used (see under 6). 


RIBEIRO, H. & R.lM08, H. Cunha -- <strong>Research</strong> o the mosquitoes o[ A,gola -- X 

DISTRIBUTION AND BIOECOLOGICAL NOTES. C. rubinotus is one 

of the most widely distributed 12u,,nelanomyia in the Ethiopian Region. 

This is probably also the case in Angola, though at present it is known 

there from only theree localities (Map 5). 

The Cabinda specimen was a gravid female caught at rest on bush, 

near the airport of the town. The 29 females from Luando River 

were caught biting man in the afternoon (17.15-17.45) during larval 

collecting (larvae of rubinotus not found) in the shallow overflown 

grassy margins of the river, near Quimbango, in the wooded savanna 

biome ((>). In this area, man was insistently bitten by C. rubinotus, 

Ae. argenteopunctatus, An. wellcomei ugandae, An. cydippis 

and M. uniformis, the first two being true pests. 

As far as we know, this marked anthropophily (and exophagy) of 

C. rubinotus had only been recorded by Hamon (37), in the Bobo 

Dioulasso area. 

7.3.8 Culex (Eumelanomyia) wigglesworthi Edwards, 1941 


MATERIAL EXAMINED. DALA TANDO, 8 larvae, 23.VIII. 1969. 

TAXONOMIC NOTES. Angolan larvae agree with the descriptions 

given by De Meillon et al. (28) and by Hopkins (24). The siphonal 

index in our eight mounted whole larvae ranged from 7.1 to 8.3 

(mean 7.5). 

DISTRIBUTION AND BIOECOLOGICAL NOTES. C. wigglesworthi seems 

to be mainly a West African mosquito, to which Subregion belongs 

the only known Angolan locality for this species (Maps 3 and 5). 

Larvae of C. wigglesvorthi bred in a small ground pool at the 

edge of a river in the Dala Tando humid forest of the Scarpment 

Zone. The breeding water was turbid, dark, foul water with many 

decaying leaves and vegetable debris. 

Larval An. demeilloni, C. cinerellus, C. kingianus, C. tigripes and 

U. mashonaensis were associated with wigglesworthi in the only An- 

golan breeding place recorded for this species (Fig. 3). 

Estud., E,s. e Doc.- 134 61

RIBgIRO, H. & R^M08, H. Cunha -- <strong>Research</strong> o, the mosquitoes of Angola -- X 

As it is the rule in subgenus Eumelanomyia, C. wigglesworthi 

seems to be also a relatively uncommon zoophilic mosquito. 

7.4-- Subgenus Culiciom¾ia Theobald, 1907 

Subgenus Culiciomyia cornprizes near 50 described species endemic 

to the Ethiopian, Oriental and Australian Zoogeographical Regions. 

Like for the preceding two subgenera, most of the 17 known 

Ethiopian members of subgenus Culiciomyia seem to have a relatively 

narrow range of geographical distribution. In Angola, are known to 

occur three of the most generalized Ethiopian species. 

7.4.1- Culex (Culiciomyia) cinerellus Edwards, 1922 

The only previous ecord of C. cinerellus is that by Grindata (10) (11), 

from Mulondo. 

MATIilRIAL IXAMINID. BOCOIO, 9 larvae, 8.IX.1970; CABINDA, 3 c, 

6 larvae, 19-20.VIII. 1970; DALA TANDO, 3 9, 3 c, 18 larvae, 23-26.VIII. 1969: 

DANGE-IA-MENHA, 2 57, 1 c, 2 larvae, 28.VIII.1969; DUNDO, 10 larvae, 

18-19.VIII.1969; UIGE, 1 c, 2 larvae, 25.IX.1969. 

TAXONOMIO NOTES. Both Angolan adults and larvae agree quite 

well with the descriptions given by Edwards (8) and by Hopkins (24). 

In larvae, however, dorsal pair of anal papillae are only about twice 

length of saddle while the ventral pair is appreciably shorter, about 4/, 

of dorsal pair. 

DISTRIBUTION. Map 7 shows the present known distribution of 

C. cinerellus in Angola, concerning mainly the West African Subregion 

(see also Maps 2 and 3). 

BIOIOOIaOIglCAL NOTES. All our adult cinerellus specimens were 

reared in the field laboratory. 

11 breeding places were recorded by the writers for cinerellus, 

usually in forested areas (climatic and gallery forests): 5 rock-pools, 

3 ground pools and niches at river edges (both kinds usually with abun- 

62 Estud., Etas. e Doc.- 134


dant fallen leaves), a pool in sandy river bed, a crabhole and a fallen 

banana leaf. 

The breeding water was usually turbid, sometimes even foul water 

with decaying leaves, or cut bamboos. The pH, measured 

in three instances, ranged from 5.4 to 8.0. 

Larval U. mashonaensis was the most frequently associated species 

(in 4 breeding places), followed by C. tigripes (3) and E. chrisogaster (2). 

Nine other species of Culex (annulioris, decens, inconspicuosus, ingrami, 

kingianus, p. pipiens, simpsoni, univittatus and wigglesworthi), three of 

Aedes (aegypti, nigricephalus and yangambiensis) and Anopheles (coustani, 

demelloni and rufipes), and also U. [usca were recorded only once. 

Ae. nigricephalus was the associated species in the crabhole breeding 

place. Figs. 1, 3 and 8 ilustrate some cinerellus larval biotopes. 

C. cinerellus is known to be a zoophilic species, which seems to 

be also the case for the Angolan populations of this mosquito. 

7.4.2--Culex (Culiciomyia) cinereus Theobald, 1901 

The only known previous Angolan record of C. cinereus is that 

by Grindata (lO) (11), from Huambo (= Nova Lisboa). 

MATERIAL EXAMINED. MALANJE, 9 larvae, 12.IV. 1971; SAURIMO, 

4 larvae, 10.IV. 1971; UIGE, 3 larvae, 15.IV.1971. 

TAXONOMIC NOTES. Angolan larvae of C. cinereus agree quite well 

with the description given by Hopkins (24), being easily identifiable. 

The siphonal index in our mounted material is only 1.5-2. 

DISTRIBUTION AND BIOECOLOGICAL IOTES. The known distribu- 

tion. of C. cinereus in Angola (Map 7) seems to indicate that it is 

mainly an West African mosquito. 

All our larvae were breeding in artificial containers in the peri- 

domestic habitat: gourds (4), barrels (2), discarded tins (2) and a drum. 

The only species we found in association with larval cinereus were 

A e. aegypti and C. p. quinquefasciatus. 

C. cinereus is known to be mainly a zoophilic mosquito (35) (37), 

though it was reported as biting man in Uganda, by Corbet et al. (43) 


RIBgIRO, H. & RAM08, H. Cunha -- <strong>Research</strong> on the mosquitoes of Angola X 

7.4.3--Culex (Culiciomyia) nebulosus Theobald, 1901 

The only known previous record of C. nebutosus in Angola is 

that by Colago (9), from a treehole at Sambo. 

MATERIAL EXAMINED. CABINDA, 3 larvae, 14.IV.1971; CAFIMA, 4 lar- 

vae, 20.IV. 1972; CUANGAR, 1 9, 2 , 16 larvae, 16.IV.1965; CUI. 5 larvae, 

13.III.1970; DALA TANDO, 25 larvae, 25-26.Vlli.1969; HUAMBO, 27 larvae, 

6-19.V.1971; JANGADA DO CUANGAR (50 km W), 4 larvae, 20.1V.1965: 

LUANDA, 2 larvae, 19.III. 1971; LUBIRI, 4 larvae, 30.III. 1972; MALANJE, 1 larva, 

12.IV.1971; N'GIVA, 2 larvae, 18.II.1970; PANDA, 35 larvae, 14-15.IV.1965: SAU- 

RIMO, 4 larvae, 10-11.IV.1971; SERPA PINTO, 10 larwte, 24.V.1971: TOMBINGA, 

4 larvae, 28.VIII. 1969. 

TAXONOMIC NOTES. Both our adult and larval specimens agree ouite 

well with the descriptions of C. nebulosus given by Theobald (6), 

Edwards (8) and Hopkins (24). 

Considering its known distribution in Angola (see below), both 

presumed subspecies of C. :ebutosus were expected to be represented 

in the country. According to the available evidence, however, we are 

not satisfied of the characters given by Edwards (op. cit.) and by 

Peters (44), on which is based the recognition of the nominal and pseudo.. 

cinereus subspecies. On the other hand, if we take into account the 

occurrence of both adult forms in the same area (e.g. Elisabethville), 

as it is cited by Mattingly (42) and by Peters (op. cit.), there seems to 

be strong indication to treat C. nebutosus as a polymorphic species and 

not as a polytypic one. In fact, polymorphism is to be expected in those 

monotypic species which are able to occupy novel ecological niches 

[Mayr (45), pp. 150 et seq.; p. 245], such as that offered by the immediate 

human environment. 

DISTRIBUTION. Map 7 shows the wide known distribution of C. 

nebulosus in Angola, extending to both West and East-South African 

Subregions (cf. Maps 2 and 3). 

BIOECOLOGICAL NOTES. Our adult specimens of C. nebutosus were 

all reared from larvae. 

37 nebulosus breeding places were recorded by the writers, 21 of 

which in the peridomestic habitat and 16 in the sylvatic habitat. As 

154 Estud., Etas. e Doc. 134

RIBgIRO, H. & RalVI08, H. Cunha -- <strong>Research</strong> o the mosquitoes of Angola X 

belonging to the first group, including all man-dependant water collections 

influenced by rains, were found, by decreasing order of frequency: 

old rubber tyres (10), barrels and drums (4), flower vases (2), cement 

water tanks (2), abandoned domestic utensils (2) and a discarded tin 

of sardines. As sylvatic breeding places, consisting in naturally occurring 

exclusively climate-dependant for water, were recorded: 

holes and axils in living trees such as Ficus sp. and Ricinodendron sp. (7), 

bamboo holes and stumps (5), holes in rotting tree (banana, papaya and 

baobab) trunks (3), and a rock-pool in gallery forest covered with decaying 

fallen leaves. Fig. 4 shows a typical larval biotope of C. nebulosus. 

The breeding water was always markedly shadded and ranged from 

clear limpid water to foul water with high organic matter content. 

The figure found for the pH of the breeding water in a Ficus hole 

was 7.0. 

In 12 out of the 37 larval biotopes recorded for C. nebulosus no 

other associated mosquito larvae were found, 8 of which in peridomestic 

and 4 in sylvatic habitats. In peridomestic habitat, the species found 

in association with neb.:losts were: C. p. quinque/sciatus (in 6 biotopes), 

A e. aegypti (5), C. tri/ilatus (2) and C. duttoni, C. horridus, C. univittatus 

and E. chrysogaster once each. In sylvatic habitat, the associated species 

were: (a) in holes and axils of living trees, Aedes spp.: calceatus (3), 

/ulgens (3), metallicus (2) and aegypti and marshalii once each; (b) in 

bamboos, Ae. angustus and E. chrysogaster, also only once, and (c) in the 

sole rock-pool recorded, E. chr?ogaster and U. mashonaensis. The apparent 

exclusion of C. tigripes as an associated species of C. nebulosus 

is perhaps a real one and it seems particularly intriguing as both species 

haYe partly common potential breeding areas. 

Though being the most common of the Ethiopian Culiciomyia, C. 

nebulosus is known to be also a zoophilic mosquito (35) (37), most probably 

an ornithophilic species. 

7.5--Subgenus Culex Linnaeus, 1758 

Nominal subgenus, with near 200 described species, is the largest 

of all Culex subgenera. Though subgenus Culex is represented in all 

Zoogeographical Regions, the group is mainly Ethiopian and Neotropical. 

From the 66 species of this subgenus occurri_g in the Ethiopian 


RIBgIRO, H. & RAMOS, H. Cunha -- <strong>Research</strong> on the mosquitoes of Angola q X 

Region, 3 of them are shared with other Regions (Palaearctic, Oriental 

and Australian) and other 6 species occur only outside the African Continent 

(Madagascar, Seychelles and Silo Tom). 

31 out of the 57 species of Culex s. str. endemic in Africa South of 

Sahara are here recognized as belonging to the mosquito fauna of Angola 

(Table 1). 

Group A (or sitiens) of Edwards (46), chiefly Oriental, is well represented 

in Angola by 13 of the about 20 Ethiopian species of the group. 

The larger Edwards' group B (or pipiens) is represented in the Ethiopian 

Region by some 35 species belonging to series pipiens and decens. Pipiensseries 

is mainly an East African subgroup and is poorly represented in 

Angola by only 7 species, some of which may prove to be subspecifically 

distinct (e.g. chorleyi, toroensis). Decens-series, chiefly West African, is 

in turn very well represented in Angola by 13 taxa, some of which may 

also prove to be new (e.g. our ingrami-like form). 

In the systematic account of the Angolan members of the subgenus 

that follows species were not grouped, each taxon being treated in alphabetical 

order. Distributional maps (Maps 8-17), however, were made by 

grouping the species according to the main groups of Edwards. 

7.5.1--.Culex (Culex) annulioris Theobald, 1901 

C. annulioris was firstly recorded in Angola by Gamble (7) from 

Sio Salvador, and later by Edwards (8), from Bi (Wellman coll.). In 

1956, this species was recorded again by Gfindara (10) (11), from Dilolo, 

Cuchi and Huambo. 

MATERIAL EXAMINED. ALTO CATUMBELA, 1 larva, 17.IX.1970; BE- 

LIZE, 1 /, 8 larvae, 20.VIII. 1970; BOCOIO, 7 ?, 5 3, 25 larvae, 7-10.IX.1970; 

BUCO ZAU, 2 larvae, 20.VIII. 1970; CAALA, 1 8, 13.V.1970; CABINDA, l 

4 8, 10 larvae, 18-26.VIII. 1970; CALULO. 1 larva, 9.VI.1970; CAPUNDA, 

4 larvae, 16-17.VI.1970; CARIANGO, 2 8, 1 larva, 18.X.1969; CEILUNGA, 9 lar- 

vae, 10-11.IX.1969; CHIQUITE, 1 8, 2 larvae, 29.V.1970; CHITEMBO, 1 

18.VI.1970; CUANDO, 8 9, 11 /, 4 larvae, 26.V.1970 and 12.V. 1971; CUBAL, 

2 $, 5 8, 9 larvae, 12-15.IX.1970; DALA TANDO, 1 9, 3 8, 3 larvae, 25.VIII. 1969; 

DUNDO, 1 larva, 19.VIII.1969: EBANGA, 10 9. 12 8, 29 larvae, 17-19.IX.1970; 

I-IUAMBO, 66 $, 78 /, 105 larvae, several dates from 25.I.1964 to 14.V. 1971; 

LONIAVA, 1 , 4 8, 7 larvae, 17-18.IX.1970; LUCE RIVER, 1 larva, 17.VI.1970; 

LUINGA, 1 /, 10 larvae, 1.X.1969; MALAIXTTE, 8 , 3 8, 10 larvae, 21-22.I.1970; 

66 Zstud., Ens. e Doc.- 134

RIBEIRO, I-[. & RAMOS, I-[. Cunha -- <strong>Research</strong> on the mosquitoes of Angola X 

MONA QUIMBUNDO, 7 2, 3 , 5 larvae, 10. IV.1970; MUSSENDE, 3 larvae, 

21.X.1969; QUELA, 5 larvae, 19.I.1970; QUIMBANGO (21 km N), 1 2, 1 larva, 

15.VI.1970; QUITEXE, 3 larvae, 27.IX.1969; SA DA BANDEIRA (20 km W), 1 2, 

4 larvae, 6.VI.1969; SERPA PINTO, 1 2, IV. 1971; TCHIVINGUIRO, 7 

28 larvae, 29.V.1969; UiGE, 4 larvae, 10-25.IX.1969. 

TAXONOMIC NOTES. As much as five forms of C. annulioris were 

described in addition to the : gambiensis, congolensis, 

major, consimilis (this one as tigripes var.) and mayumbae (as bitaeniorhynchus 

vat.). These were reduced to major and consimilis by 

Edwards (8) who, lesides the differences on abdominal tergites, also 

pointed out to some slight differences in phallosome. Later, Hamon & 

Ovazza (47), emphasizing the examination of phallosomes, came to the 

conclusion that only two subspecies were valid: C. annulioris annulioris 

(the typical form) and C. annulioris consimilis. 

It seems to be now quite evident, however, that the variation on 

the abdeminal tergites of C. annulioris is of a continuous kind. All the 

three kinds of tergal abdominal pale markings in this species (basal, 

latero-apical and ground pale markings) are subjected to a marked, 

largely (positively) correlated variation. They all may be either enterely 

absent (congolensis form) or more or less developed. In the other extreme 

of this variation, the abdomen becomes extensively pale (paler than in a. 

annulioris types), with the usual triangular basal and latero-apical mark- 

2g:. coalescing as the result ef the presence of many scattered pale 

scales on the ground (which is then perhaps better described as being 

pale with some scattered dark scales). 

On the other hand, in that concerns phallosome, variation in spiculation 

seems to be also continuous in kind. Moreover, sometimes this is 

apparently also the case in males reared from the same breeding place, 

some of these having the inner divisions of the , 

others presenting the strong spicules of the consimilis form (or even stronger 

than figured by Edwards), while others are intermediate. It is also 

to be noted that, sometimes, even the degree of spiculation of the inner 

division differs markedly from the left to the right side of the same phallosome. 

Lastly, this terminalic character is often in disagreement with the 

tergal abdominal pattern not only of the corresponding males but also 

of the associated females reared from the same breeding place. This was 

certainly the case for the form mayumbae of Galliard (31) which phallo- 



some was considered by Edwards (8) and by Hamon & Ovazza (47) as 

of the consimilis form, while the abdomen was surely distinctly pale 

marked, since the series was identified by Galliard as a variety of bitaeniorhynchus. 

Other instances of similiar discrepancies may be also found 

in Hamon & Ovazza, op. cit., and Gfindara (10) (11) (Dilolo records, 

p. 405). 

In that concerns larvae, we could also confirm that the different 

series of C. annulioris cannot be distinguished in the larval phase. 

According to the above remarks, it is not surprising, of course, 

that it becomes not possible to delimitate the geographical distribution 

of each of the two forms of C. annulioris in a way coherent with their 

presumed subspecific status. Such a delimitation is in fact not satisfactory, 

either if we base our diagnosis on the abdominal scaling of females, or 

on the terminalic characters of the males, or even less on both. And 

that this is true also outside Angola, even if we consider only the external 

morphology, is shown by the instances cited in the literature, as by 

Edwards (8) (Nairobi and Bole, p. 295; the records by Schwetz, p. 296), 

by Mattingly (48) (Kumasi and Takoradi) and by Hamon & Ovazza, op. 

cit. (Akaki, Addis-Abeba and Kolda), in which both forms occur at the 

same locality. 

These are the reasons why the writers propose that C. annulioris be 

treated as a monotypic polymorphic species with darker strains predominating 

in forest biomes, in a way similar to that one is now generally 

adopted for Ae. aegypti. This is also, naturally, the taxonomic treatment 

adopted in the construction of our keys. Accordingly, previous records 

of annulioris subspp. by Theobald (6), Edwards (8), and Gfindara (10) (11) 

are here not included in the list of the Angolan mosquitoes. 

The branching of head seta e will separate most of the larval specimens 

of annulioris from those of bitaeniorhynchus/ethiopicus (see larval 

key under 6). 

DISTRIBUTION. As it is shown on Map 8, C. annulioris is widely 

distributed in Angola, extending to both the West African and East-South 

African Subregions, with the exception of semiarid biomes (cf. Maps 2 

and 3). 

BIOEOOLOGICAL NOTES. The characteristics of 93 breeding places 

of C. annulioris were recorded, almost all of them (89) consisting in 

t58 Estud., Etas. e Doc.- 134

RIBEIRO, H. & R.,MOS, H, Cunha -- <strong>Research</strong> on the mosquitoes of Angola 

ground water. Naturally occurring breeding places such as river margins 

(26), ground pools at river edges (24) and permanent pools (7) 

constitute the main breeding area of C. annulioris. Man-made irrigation 

drains (19) and shallow earth wells (8) merely extended that main natural 

breeding area. Lastly, the occurrence of annulioris in rock-pools (5) is 

to be related to its (incipient?) ability to occupy the novel niches offered 

by artificial containers in peridomestic habitat (4): two flower vases, a 

drum and a discarded tin. Fig. 5 shows a small earth drain of turbid water 

where C. annulioris bred. 

In relation to this presumed trend to man-dependancy of annulioris 

for breeding, it is also to be noted that the breeding water is often turbid, 

dark and even foul water, apparently with a high organic matter content, 

not infrequently clue to human activity (decaying , contamination 

by human or cattle excreta, etc.). 

In 10 out of the 89 ground water larval biotopes, abundant filament 

tous green algae were recorded which, however, are not eaten by the 

larvae. In 61 out of the 93 breeding places the pH of the breeding water 

was measured, figures ranging from 4.8 to 7.0, with a mean of 5.8. 

Associated mosquito larvae were only recorded in 69 out of the 93 

breeding places of C. annulioris. Genera Anopheles (22 species) and 

Culex (19) were the only mosquito genera significantly associated with 

annulioris in its larval biotopes. In terms of species, the most frequent 

associates were: An. coustani (24 times), C. tigripes (10), An. cydippis 

and C. univittatus (9 times, each), An. rufipes rufipes (6), An. demeilloni, 

An. funestus, An. rnaculipalpis, C. duttoni and C. salisburiensis (5, each) 

and An. gambiae, An. marshalii, An. squarnosus, C. antennatus and 

C. argenteopunctatus (4, each). All the other species were associated 

occasionally, being found in less then 5 per cent of the above mentioned 

69 breeding places. 

In spite of being widespread and fairly common in Tropical Africa, 

C. annulioris is known to be mainly a zoophilic mosquito, though it may 

bite man rarely (35) (49) (43) (37). In Angola (Huambo), only two females 

were caught as such by the writers, both at rest, at 7.30 a.m., one of 

them in a hen-house and the other in a cow-house. All the other adult 

specimens in collection were laboratory-reared. 

Estud., Etas. c Dot'.- 134 69 

X


7.5.2--Culex (Culex) antennatus (Becker, 1903) 

C. antennatus was firstly recorded from Angola (Huambo) by Grin- 

data (lo) (11) and later by Worth & Paterson (12), from Caxito, and by 

one of us (34) (50), from Lobito and Luanda. 

MATERIAL EXAMINED. BOCOIO, 2 9, 8.IX.1970; CALAI, 26 9, 7 3, 

plus 48 larvae, several dates during 1965-67; CUANGAR, 47 larvae, 10. IV.1965 

and 5.V. 1966; CUBAL, 15 9, 14 3, 21 larvae, 14.IX.1970; DALA TANDO, 

1 larva, 28.VIII. 1969; DIRICO, 7 9. 8.V.1965; EBANGA, 1 larva, 17.IX.1970; 

HUAMBO, 3 9, 1 3, 1 larva, several dates 1964-1971; FOZ DO QUEVE, 7 larvac, 

29.X.1969; LUINGA, 1 3, 2 larvae, 1.IX.1969; MALANJE, I larva, 22.I.1970; 

MAZOZO, 5 larvae, 10.IV.1970; MO(AMEDES, 59 , 17 plus 45 larvae, 

18-23.X.1967 and 28.IV-3.V.1969; MOXICO, 3 larvae, 18.IV.1970; MUMBONDO, 

1 9, 1 3, 2 larvae, 7.V.1970; MUXIMA, I 3, 29.IV.1970; NANGURA, 19 ?, 

7.VI.1965; PORTO ALEXANDRE, 2 9, 16.V.1969; QUISSOL, 2 9, I 3, 4 larvae, 

23.I.1970; SACO DO GIRAUL, 1 , 4 larvae, 7.V.1969; S3.O NICOLAU, 3 9, 

4 plus 38 larvae, 28.IV.1969, all new locality records. 

TAX,OOMIC NOTES. Angelan antennatus agrees quite well with 

known descriptions, namely those by Edwards (8), Hopkins (24) and Larivire 

& Abonnenc (51). Males are easily identifiable through the examination 

of terrainalia. Females need to be carefully separated from those 

of univittatus and of C. pipiens pipiens, which may be better accomplished 

with the aid of our key. The difficulties found in the separation of 

antennatus larvae from those of decens, invidiosus, telesilla, trifoliatus 

and univittatus are well known. The larval key here proposed, however, 

will permit a correct diagnosis at least of the great majority of the 

specimens. As always, the association of specimens of different phases 

and sexes will be of great help, and it must be kept in mind that univittatus 

is an usual mosquito in the larval biotopes of antennatus, at least 

in Angola (see below). 

DISTRIBUTION. The general distribution of C. antennatus is a quite 

interesting one, the species being widely distributed throughout the Tropical 

Africa with an extension into the south-eastern Mediterranean 

Subregion of the Palaearctic (Egypt, Palestine, and Iran). In Angola, it 


RIBgIRO, H. & RAM08, H. Cunha -- <strong>Research</strong> o the mosquitoes of ,4ngola -- X 

is also a widespread and common mosquito in the savannas and highland 

biomes (Maps 15 and 3). 

BIOEGOLOGICAL NOTES. 42 breeding places were recorded from 

C. antennatus, 39 of them consisting of ground water: pools at river 

edges and backwaters (12), ground pools (10), irrigation drains (5), river 

margins, usualy overflown (4), pools in river beds and shallow earth 

wells (2, each) and a wheel track. Though being mainly a ground water 

breeder, larval antennatus was also recorded from cement water-tanks (2) 

and from a river boat. 

As a rule, the breeding places of antennatus were either scarcely 

shaded or even unshaded. The characteristics of the breeding water 

ranged widely from clean, limpid water, sometimes with Pistia or Lemna, 

to turbid water with an appreciable organic matter content. The pH of 

the breeding water, measured in 20 biotopes, varied between 5.3 and 8.0 

(mean 6.7). The salt content in 9 biotopes examined ranged from 0.2 to 

5.7 (mean 1.5) g/1 (NaCl). 

Geaera Culex (14 species) and Anopheles (7 species) were the only 

mosquito genera significantly associated with larval antennatus. The most 

frequently associated species were C. univittatus (in 23 biotopes), An. 

gambiae (13), An. squamosus, C. poicilipes and C. tigripes (8, each), An. 

coustani and C. p. pipiens (7, each) and An. argenteolobatus and An. 

pharoensis (4, each), while An. listeri, C. simpsoni form A and C. telesilla 

(3 times, each) occurred as associated species in the South-west Arid Zone 

(Map 3). Other species were recorded as occasional associates in less 

than 5 per cent of the breeding places. 

Figs. 6 and 14 illustrate two antennatus breeding sites. 

24 antennatus females were caught as such, not laboratory-reared. 

19 out of these were caught at rest in cow-houses (from 18.00 to 19.00 

hours), most of them gorged. Other 3 females were found at rest inside 

human habitations and only 2 antennatus females were seen biting man 

at dusk (18.00-18.30), in the open. Occasional man-biting was also 

observed by Haddow et al. (35), in Uganda, by Someren et al. (49), in 

Kennya, and by Hamon (37), in the Upper Volta. C. antennatus seems to 

be, thus, mainly a zoophilic mosquito which in areas of more intensive 

breeding may, in fact, attack man. 

Estud., Etas. e Doc.- 134 71

RIBEIRO, H. & RAMOS, [-I. Cunha -- <strong>Research</strong> on the mosquitoes of Angola X 

7.5.3- Cutex (Culex) argenteopunctatus kingi (Theobald, 1913) 

C. argenteopunctatus kingi was firstly recorded from Angola (Capule, 

near Sambo) by Colago (9) and later, by Gttndara (10) (11), from Cuando 

and Cuchi. 

MATERIAL EXAMINED. CARIANGO, 4 9, 1 3 plus 5 larvae, 14.X.1969; 

CEILUNGA, 1 3, 9 larvae, 10-11.IX.1969; EBANGA, 2 9, 2 3, 18-19.IX.1970; 

HUAMBO, 3 9, 3 3, 7 larvae, I.VI.1966, 21.IV-13.V.1970 and 12.V.1971; LUCE 

RIVER, 3 larvae, 18.VI.1970; MOXICO, 1 larva, 16.IV.1970; NANGURA, 1 larva, 

9.XII. 1965; RIO GANGO, 1 larva, 21.X.1969, all new locality records. Also 

CUANDO, 5 larvae, 12.V. 1971. 

TAX'ONOMIC NOTES. Our adult specimens of kingi agree quite well 

with known descriptions. Larvae are also highly characteristic. In Angolan 

material, the comb of seg. VIII has more numerous teeth (about 

30, from 20 to 35) than in Uganda larvae as described by Hopkins (24). 

Other particulars of our larvae to be mentioned are: preclipeals long and 

thin, delicate; head setae d minute and single, almost in line with bases 

of setae B; upper caudal seta occasionally single or trifid, though usually 

bifid; anal papillae long and narrow, dorsal pair about four times length 

of saddle, ventral pair slightly less. 

DISTRIBUTION. Map 11 shows the known distribution in Angola of 

C. argenteopunctatus kingi, mainly a mosquito of the Montane and 

Rhodesian Highland Zones (Map 3). 

BIOECOLOGICAL NOTES. All adult specimens examined were 

reared from larvae. 

C. argenteopunctatus kingi seems to be exclusively a ground water 

breeder, as no artificial containers or plant supports were recorded for 

breeding of this mosquito. 11 larval biotopes of kingi were examined: 

margins of rivers and streams (4), pools at river edges and irrigation 

drains (3, each) and a rain water ground pool, all of them only scarcely 

shaded by grasses. Abundant filamentous green algae were present in 

four of the breeding places though, as usual, the digestive tract of larvae 

was full of Diatomeae. 

Genera Anopheles and Culex, with 8 species each, were the only 

significantly associated mosquito genera in kingi larval biotopes. The 


RIBEIRO, H. & RAMOS, H. Cunha -- <strong>Research</strong> on the mosquitoes o[ Angola -- X 

species most often recorded as associates were An. cydippis (in 5 breeding 

places), C. annulioris (4), C. univittatus (3) and An. argenteolobatus, 

An. squamosus, C. duttoni and C. tigripes (2, each). 

C. argenteopunctatus kingi is known to be mainly a zoophilic mosquito, 

though it may bite man in the outside (35). 

7.5.4- Culex (Culex) aurantapex jinjaensis Edwards, 1941 

New record for Angola of both the species and subspecies. 

MATERIAL EXAMINED. LOMBE RIVER (W of Malanje), 1 $, 3 $ 

plus 6 larvae, II-1970; CALAI, 1 larva, 30.XI.1%5. 

TAXONOMIC NOTES. Angolan adult and larval specimens agree quite 

well with the descriptions of var. finjaensis of Edwards (8). 

According to Hopkins (24), larvae of the type form are indistin- 

guishable from those of var. jinjaensis. Larvae from the Lombe River, 

bowever, are surely attributable to the Uganda-Zambia form, to which 

quite probably also belong the larva from Calai. 

According to the available evidence in Angola, larvae of jinjaensis 

can be separated from those of C. annulioris, C. bitaeniorhynchus and 

C. ethiopicus if one takes into account the chactotaxy of abdominal 

seg. VIII (see larval key). 

DISTRIBUTION. Map 8 shows the only two Angolan localities known 

for finjaensis. 

BIOEOOLOGICAL NOTES. Two breeding places were recorded for 

jinjaensis: a pool at the edge of the Cubango River and the margins of 

Lombe River. In both instances the water was clean and limpid, only 

shaded by emergent grasses. In the first of the larval biotopes, the pH 

of the breeding water was 6.5. 

C. poicilipes was an associated species in both instances, while An. 

coustani, C. simpsoni (form A) and Ficalbia mal/eyti were recorded 

only once. 



Our female was caught when biting man in the Lombe River margins, 

at dusk (7.00 p.m.). This mosquito had already been recorded as 

biting man by Mattingly & Lips (42), in the Lubumbashi area. We may 

say that in Angola, C. aurantapex ]in]aensis is a relatively rare mosquito 

more liable to occur in the Highland Zone (Map 3) and readily biting 

man in the outside. 

7.5.5- Culex (Culex) bitaeniorhynchus Giles, 1901 

The only known Angolan record of C. bitaeniorhynchus is that from 

Lobito by one of us (13). Following Sirivanakarn (52), the Angolan bitaeniorhynchus 

population seems to be closer to form domesticus, as judging 

from the published photomicrograph of the male terminalia (Ribeiro, 

op. cit., Fig. 3). C. bitaeniorhynchus was seen to bite man in Lobito, such 

as it was found by Hamon (37) in the Upper Volta. 

No bitaeniorhynchus specimens ware caught by the present writers. 

7.5.6- Culex (Culex) chorleyi Edwards, 1941 


MATERIAL EXAMINED. CAALA, 1 9, 3 8 plus 6 larvae, 13.V.1970; 

CARIANGO, 5 9, 7 8, 7 larvae, 14.X.1969; CUANDO, 1 9, 1 8, 3 larvae, 

18.V.1970; EBANGA, 3 larvae, 17.IX.1970; HUAMBO (= Nova Lisboa), 7 larvae, 

14.IV, 13.V and 14.X.1970. 

TAXONOMIC NOTES. Our adults of chorleyi are much as described 

by Edwards (8), though in some specimens a few (1-5) pale pre-alar scales 

may be present on one or both sides of thorax and the pale bands of 

tergites are sometimes reduced or even practically absent. It is also to be 

noted that the length of the yellowish spot at tip of hind tibia is markedly 

variable, sometimes being even shorter than long. 

In that concerns male terminalia, most specimens are in quite good 

agreement with the description by Edwards (8). In the males from Cariango, 

however, there are a few small teeth in the outer division of lp 

and some slight differences in the shape of the lobe of coxite and of 

its appendages. 

74 Eslud., Etts. e Doc.- 134

RIBgIR0, H. & RAM08, H. Cunha <strong>Research</strong> on thc mosquitoes of Angola 

As to the larvae, they are also in good general agreement with the 

description by Hopkins (24), always bearing short siphonal tufts. In the 

specimens from Cariango, however, the larvae lack the ventral part of 

the siphonal of spines, as it was the case of seen by Hopkins (ibidem). 

In view of the above cited peculiarities of the terminalic and larval 

characters in the Cariango chorleyi population, this would be perhaps 

better treated as a subspecies. We feel, however, that further knowledge 

on the morphology and distribution of vansomereni, chorleyi and astri- 

dianus is needed before new taxonomic decisions be taken concerning 

this group. 

DISTRIBUTION. Map 14 shows the known distribution of C. chorleyi 

in Angola, all the five localities being at 1,500 meters and over in altitude. 

larvae. 

BIOEOOLOGICAL NOTES. All our adults of chorleyi were reared from 

3 out of the seven breeding places recorded for C. chorleyi were 

shallow vegetated river margins, other 3 larval biotopes were irrigation 

canals with emergent vegetation (and some decaying in one 

of them), while in the other instance the larvae were found in a grassy 

ditch. The pH of the breeding water, measured in six breeding places, 

was 5.8 in one of them and 5.4 in all the other. Larvae of An. cydippis, 

marshalii, squamosus, C. annulioris, duttoni, tigripes, toroensis and univittatus 

were found as associated species. Fig. 7 shows a breeding place 

of chorleyi. 

C. chorleyi seems to be a zoophilic mosquito of the Humid Montane 

Zone (Map 3). 

7.5.7--Culex (Culex) decens Theobald, 1901 

C. decens was recorded for the first time in Angola by Gamble (7) 

from So Salvador, and later by Gndara (10) (11), from Capelongo. 

MATERIAL EXAMINED. BIBALA, 2 larvae, 2.V.1969; BOCOIO, 8 $, 4 /5, 

8.IX.1970; CABINDA, 1 larva, 14.III. 1971; CAINDE, 5 larvae, 12.V.1969; CAR- 

LAONGO, 2 /5, 9 larvae, 31.X and 5.XI.1969: CUANGAR, 5 $. 7 , 4.V.1966; 


RIBEIR0, H. & RAM08. H. Cunha -- <strong>Research</strong> on the mosquitoes of Atgola -- X 

CUANZA RIVER (N. of Calulo), 1 9 2 larvae, 12.VI.1970; CUBAL, 1 9, 1 larva, 

12.IX.1970; DALA TANDO, 4 larvae, 25-28.VIII. 1969; DINDE, 5 larvae, 28.V. 1971; 

DUNDO, 48 larvae, 18-19.VIII. 1969; DUQUE DE BRAGAN(A, 2 larvae, 15.11.1969; 

EBANGA, I 9, 2 $, 3 larvae, 19.IX.1970; GUNGO, 2 larvae, 23.XI.1969; 

HUAMBO, 3 9, 2 $ plus 21 larvae, 4-12.V. 1970 and 5-12.V.1971; IMPULO, 

5 larvae, 23.II1.1970; JAMBA, 1 larva, 22.I1.1971; LUANDA, I larva, XI.1967; 

LUCALA, I larva, 27.VIII.1969; LUCALA RIVER, 4 larvae, 28.VII1.1969; LUINGA, 

1 9, 3 larvae, 30.IX and 1.X.1969; SA DA BANDEIRA (20 km W), 1 9,4 larvae, 

6.VI.1969; UiGE, 3 larvae, 25.IX.1969; VALE DO LOGE, I 9, 2 plus 2 larvae, 

26.IX.1969, all new locality records. 

TAXONOMIC NOTES. Larvae of decens, invidiosus and antennatus 

are not always surely distinguishable, though this distinction will be often 

feasible following our larval key (see under 6), at least in that concerns 

samples of Angolan material. The examination of as much as possible 

associated adults, however, is always highly desirable. 

DISTRIBUTION. Map 15 shows the known distribution of C. decena 

in Angola, a mosquito mainly associated there with the West African 

biomes (see also Maps 2 and 3). 

BIOEC/OIaOGICAL NOTES. 28 out of the 32 breeding places of decens 

consisted in several kinds of ground water: rock-pools, usually with 

fallen leaves and decaying plants (10), ground pools (9), river margins, 

often among masses of filamentous green algae (6) and pools in river 

beds (3). Artificial containers were recorded in 4 other instances: 

rubber tyros (2), a drum and a cement tank. 

Species of genus Culex were most common associates (14) followed 

by genera Anopheles (11), Aedes (3), Uranotaenia (2) and Ficalbia (1). 

The most common individual species in decens larval biotopes were, 

by decreasing order of frequency: Culex tigripes (10), Anopheles coustani 

(7), Uranotaenia fusca (5), C. p. piplens and C. univittatus (4) and 

C. simpsoni form B and C. telesilla (3). Two breeding places of C. 

decens are illustrated by Figs. 8 and 12. 

Females were never caught by the writers on humans exposed near 

prolific breeding places nor adults were found indoor human habita- 

tion, which seems to show that Angolan C. decens is also a zoophilic 

mosquito. 


RIl:Em0. H. & RAMOS, H. Cunha -- <strong>Research</strong> otz the mosquitoes o[ ,4ngola X 

7.5.8--Culex (Culex) duttoni Theobald, 1901 

C. duttoni was recorded for the first time in Angola by Well- 

man (4), from Catema (Bid) and Bailundo. The other published locality 

records for duttoni in Angola are those by Gamble (7), from S. Sal- 

,7ador, by Gtndara (10) (11), from Cuando, Dilolo, Mavinga and Moxico, 

by Colaco (9), from Luanda and that by Worth & Paterson (12), from 

Dundo. 

MATERIAL EXAMINID. BOCOIO, 8 q), 8 8,2 larvae, 8-9.IX.1970; BRUCO, 

2 larvae, 29.V. 1966; CAALA, 2 larvae, no date; CABINDA, 1 8, 10 larvae, 

26.II1.1970 and 14.II1.1971; CABUTA, 1 larva, 11.VI.1970; CARIANGO, 10 larvae, 

16.X.1969; CE1LUNGA, 1 2, 1 8, 8 larvae, 10-11.IX.1969; CHIEDE, 1 larva, 

21.IV. 1972; CHINGUAR, 14 larvae, 5 and 28.1V.1971; CUANGAR, I 9, 12 larvae, 

12.Xli. 1965; CUANGO, 5 larvae, 4.XII. 1970; CUBAL, 8 9, 14 8, 6 larvae, 

12. IX.1970; DALA TANDO, 7 9, 1 larvae, 25-26.VIl1.1969; DAMBA, 2 

9 larvae, 6.IV. 1971; DINDE, 2 9, 2 8, 2 larvae, 28.V. 1971; EBANGA, 1 

1 larva, 18-19.IX.1970; EBO, 4 9, 10 8, 4 larvae, 11.X1.1969; GUNGO, 2 larvae, 

20.XI.1969; HANHA, 3 9, 5 8, 11-13.IX.1964; iMPULO, 2 9, 3 8, 3 larvae, 

18.III.1970 and 29.V. 1971; JAMBA, 8 larvae, 22.IV. 1971; LUBIRI, 5 larvae, 

29.II1.1972; LUINGA, 1 larva, 30.IX.1969; MALANJE, 6 2, 4 8, 4 larvae 

25.I.1970; MOOAMEDES, 4 q), 5 8, 7 larvae, 26.X.1967 and 3.V.1969; MONA 

QUIMBUNDO, I 9, l larva, 10.IV./970; MUSSENDE 4 9, 3 8, I larva, 

20. X. 1969; MUTANGO, 1 8, 2.V.1966; NOVO REDONDO, 1 8, 7.V. 1971; 

QUIMBANGO, 4 larvae, 19.VI. 1970; QUISSOL, i 8, 23.I.1970; QUISSONGO, 

5 larvae, 10.VI. 1970; SAUR1MO, I 8, 19 larvae, 12.1V.1970 and 10-11.V. 1971; 

SERPA PINTO, 2 2, 3 8, 22 larvae, 24.V.1971; UiGE, 2 larvae, 25.IX. 1969; 

VALE DO LOGE, I 8, 1 larva, 26.IX.1969; VILA NOVA DO SELES, 10 

10 8, 3 larvae, 20.XI.1969, all new locality records. Also CUANDO, 1 

2 8, 3 larvae, 18.V. 1970; DUNDO, 1 q), 4 8, 25 larvae, 18-19.VIII. 1969: 

HUAMBO (= Nova Lisboa), 58 q), 80 8, 99 larvae, several dates from 25.V. 1966 

to 13.V.1971; LUANDA, 4 larvae, 7.IV. 1971, and MOXICO, 53 2, 5 larvae, 

16.IV.1970. 

TAXON'OMIC NOTES. Larval C. duttoni is quite characteristic and 

easily identified (see key under 6). Larvae of C. watti, however, though 

remaining undescribed, are said to be indistinguishable from those of 

C. duttoni (24). Distinction between adults of both forms, on the other 

hand, seems to have been based by Edwards (8) on differences not of 

the specific level. 

Estud., Etts. e Doc.- 134 77

RIBEIRO, H. & RAMOS, H. Cunha -- <strong>Research</strong> on the mosquitoes of Atgola -- X 

DISTRIBUTION. Map 10 shows the known distribution of C. duttoni 

in Angola, one of the most generalized and common mosquitoes in the 

country as well as in all the Tropical Africa. 

BIOEGOLOGICAL NOTES. Only 17 out of our 342 adult specimens 

of C. duttoni were in fact caught as adults, 10 of which (4 9, 6 ) 

were taken inside human habitation: one fernmale while biting man 

and the other specimens resting on walls. The remaining adults (5 9, 

2 3) were caught in the outside, resting near their breeding places: 

on the inner walls of barrels, on vegetation and in crabholes. C. duttoni 

seems to be, thus, a mainly zoophilic mosquito, though it easily 

enters houses and occasionally bites man. 

One hundred of breeding places were recorded by the writers for 

duttoni in Angola. 29 out of these consisted of naturally occurring 

water collections: ground water in 19 instances, rock-pools in 8, a 

banana axil and a hole in a fallen baobab. 21 out of the 71 man-made 

la:val biotopes of duttoni consisted of two main kinds of ground water; 

borrow-pits and shallow earth wells and holes, often with decaying 

, or other vegetables (16) and small ditches and 

earth drains (5). The remainder 50 man-made breeding places were 

represented by artificial containers in peridomestic habitat: 30 storage 

containers with water for domestic uses (barrels and drums), 16 discarded 

objects (11 rubber tyres and 5 vases, tins, etc.) and 4 cement 

water tanks. 

The breeding water was sometimes clean and limpid, but more 

often it was turbid and foul from decaying plants. The pH was measured 

in 25 breeding places, showing figures from 5.1 to 8.0. Most 

biotopes (21), however, had an acid water (5.1-6.7), while only in 2 

of them the water was neutral, and figures over 7.0 were only found 

in other 2 instances (7.2 and 8.0). The salt content of the breeding 

water from a crabhole at Moqttmedes (coast) was measured, giving a 

figure of 1.29 g/1 (NaCl). 

Many species were found as associates in the breeding biotopes 

of C. duttoni, the most common of which were, by decreasing order 

of frequency: C. p. quinquefasciatus (24 instances), C. tigripes (20), 

C. p. pipiens (6), An. coustani and C. univittatus (5), Ae. aegypti (4) 

and C. annulioris, C. decens and C. ethiopicus (3). C. p. quinquefasciatus 

was the main associate of duttoni in water storage containers followed 



by C. tigripes, Ae. aegypti, C. p. pipiens, C. decens, C. trifilatus and 

C. pruina. 10 out of the 20 breeding sites with the associate tigripes 

were ground water, while other 6 consisted of artificial containers 

and 4 were rock-pools. 

From the above data concerning breeding of C. duttoni, it seems 

probable that this mosquito was once mainly a plant breeder and that, 

later, as it is usual in such cases, it was able to use rock-pools. From 

these, also as usual, artificial containers in peridomestic habitat could 

then be invaded, which seem to be now the most favoured kind of 

all the larval biotopes of duttoni, at least in Angola. At the same time, 

however, it seems that C. duttoni was also able to develope another 

trend for breeding in ground water which has also become an important 

breeding area for this mosquito. As the relative importance of 

these two kinds of breeding areas is reversed in the case of its redoubtable 

predator and main associate C. tigripes (see under 7.1.1), it would 

be interesting to ascertain the impcrtance of the selection pressure 

exerted by this predator on the development of the ability of C. duttoni 

for exploring peridomestic habitat. 

Fig. 9 illustrates a typical breeding site of C. duttoni. 

7.5.9--Culex (CuIex) ethiopicus Edwards, 1912 


MATERIAL EXAMINED. All the following locality records for C. ethiopicus 

are based on the examination of laboratory reared adults: BERO RIVER (N. of 

Virei), 7 9, 12 /5 plus 7 larvae, 12.V.1969; CAINDE, 4 9, 2 /5, 16 larvae, 

same date; CALAI, 1 9, 2 /5, 4 larvae, 27.IV. 1966 and 24.V.1967; CARACULO. 

8 2, 6 /5, 14.V. 1969; CHITADO, 1 /5, 21.II.1970; CUBAL, 4 9, 1 /5, 15.IX.1970; 

LAGOA DOS PARALELOS, I 9, 1 /5, 4 larvae, 7.V.1969; LUINGA, 1 /5, 

1.X.1969; MACULAMA, 1 ,o, 5 larvae, 21.V.1969; MOnAMEDES, 4 9, 9 /5, 

13 larvae, 20.X.1967 and 29.IV. 1969; MUNHINO, 4 9, I /5, 6 larvae, 7.V.1969. 

TAXONIOMIC NOTES. In our adult specimens of C. ethiopicus the 

pale apical bands on tergites are sometimes widened laterally, while 

in some of the specimens tergites vI-VIII are mostly pale. The colour 

of these pale markings on tergites varies in the Angolan material from 

pure white to yellow. The distinction between adult ethiopicus and 

Estud., Ens. e Doc. -- 134 79


the form of bilaeniorhynchus occurring in Africa (? domesticus) is 

given in the adult key. 

As stated by Hopkins (24), larvae of C. ethiopicus are indistin- 

guishable from those of C. bitaeniorhynchus, though both can usually 

be separated from larval annulioris. On the other side, all our ethio- 

picus and annulioris larvae are distinguishable from those of auran- 

tapex }injaensis by means of the chaetotaxy of abdominal segment viii 

(see larval key). 

DISTRIBUTION. Map 9 shows the known distribution of C. ethio- 

picus in Angola, mainly confined to the driest areas of the country 

(see also Maps 2 and 3). It is probable, however, that ethiopicus be 

a qdite generalized species and that several of our larval records 

of the group throughout Angola (Cariango, Dala Tando, Dinde, Dirico, 

Ebanga, Huambo, Hunguria, Lonjava and Quilengues) belong, in fact, 

to this species, namely those from the aridest of these areas. 

BIOEClOLOGICAL NOTES. All our adult specimens of C. ethiopicus 

were reared in the laboratory from larva or pupa. 

C. ethiopicus is a ground water breederø Twelve breedings places 

were recorded by the writers for this mosquito: 8 pools in river beds, 

3 open ground pools and a small pond. The larval biotopes were always 

unshaded or scarcely shaded, usually with clean water and often without 

any vegetation. No filamentous green algae were recorded. The breed- 

ing 'wate,r wais ialways acid, With a pH, measured in 10 biotopes, 

varying from 5.3 to 6.7 (mean 6.2). The salt content (NaCl) of the 

water, also examined in 10 biotopes, was always low, ranging from 0.1 

to 2.1 (mean 0.8) g/1. 

Seven Anophelines and four other Culicines were recorded in the 

ethiopicus breeding places. The species more often found as associates 

were: An. gambiae (7 biotopes), An. listeri and An. pretoriensis (6 each), 

C. simpsoni form A (4) and C. univittatus (3). Fig. 10 shows a typical 

ethiopicus breeding place. 

As no adults were ever found inside human habitations nor collectors 

were ever bitten when exposed near its prolific breeding places, 

it seems that C. ethiopicus is a zoophilic mosquito also in Angola. 

80 Es,ud.. Etts. c Doc.- 134

RIBEIRO, H. & RAMOS, H. Cunha -- <strong>Research</strong> on the mosquitoes o[ Angola X 

7.5.10--Culex (Culex) grahami Theobald, 1910 

The only published Angolan record of C. grahami is that by Grin- 

dara (10) (11), from Lobito. 

MATERIAL EXAMINED. CANGANDALA, 2 larvae, 15.11.1970; CANGAN- 

DALA, 24 km SE (road to Bembo), 2 larvae, 10.I1.1970; MALANJE, 7 larvae, 

1 .II. 1970. 

TAXON'OMIC N.OTES. Our larvae of C. grahami exhibit some non- 

negligible variation. The number of subventral tufts of siphon (always 

with 2-4 branches) varies from 8 to 14 (both sides together), some 

of the specimens approaching in this respect, var. farakoensis of 

Hamon (53). The length of these tufts is also noticeably variable, as 

they may reach in some specimens to about 2//3 diameter of siphon. 

Tlne most proximal of the subventral tufts are often inserted before 

the last 1-3 pec[en teeth, as figured by Hopkins (op. cit., Fig. 207) but 

not mentioned in the text. Also tb, e number of comb spines may be 

as high as 36, though it is usually about 20. Finally, it is to be noted 

that the last abdominal segments, including the distal of saddle, 

of the Angolan larvae are obviously spiculate. 

DISTRIBUTION. Map 17 shows the known distribution of C. gra- 

hami in Angola. 

BIOEOOLOGICAL N.OTES. C. grahami seems to be a ground water 

breeder. 2 of the larval biotopes recorded by the writers were grassy 

rain-water ground pools and the other one consisted of a pond with 

abundant aquatic vegetation, including water-lilies. In one of the ground 

pools, near a village, the water was dark and turbid, with high organic 

content, but it was clean in the other instances. 

Larvae of ,4n. coustani, An. squamosus and C. weschei were found 

as associates in two of the biotopes, while Aedomyia africana was only 

recorded from the pond. 

According to our data, C. grahami seems to be a relatively uncom- 

mon zoophilic mosquito in Angola. In Kenya, Someren et al. (49) 

reported grahami as biting man very rarely, in bush. 

Estud., Ens. e Dot'.- 134 81

RIBgIRO, H. & RAMOS, H. Cunha -- <strong>Research</strong> otz the mosquitoes of Angola -- X 

7.5.11 -- Culex (CuIex) guiarti Blanchard, 1905 

C. guiarti was firstly recorded from Biff and Bailundo by Well- 

man (4), in 1905, and, much later, by Worth & Paterson (12), from 

Caxito, in 1961. 

MATle,,RIAL EXAMINED. AMBRIZETE, I larva, 4.VIII. 1790; CABINDA, 

2 2, 12 larvae, 18.VIII.1970; CEILUNGA, 1 larva, 11.IX.1969; UIGE, 2 larvae, 

25.IX.1969, all new locality records. 

TAXONOMIC NOES. The green colour of thorax is obvious in one 

of our females, though in the other one the thorax is brownish. Both 

females were reared from larva. 

Larvae agree quite well with Hopkins' description. We must note, 

however, that in two of them there are only 5 comb spines on each 

side and that either the subventral tufts of siphon, the upper caudal 

seta, or the lateral saddle hair may have 4 branches. 

DISTRIBUTION. C. guiarti is widely distributed in Tropical Africa, 

including Madagascar. In Angola (Map 17), all the known locality 

records for guiarti are in the northern half of the country. 

BIOEOOLOGICAL NOTES. C. guiarti breeds in ground water. Four 

larval biotopes were recorded by the writers: 2 small ponds with abundant 

aquatic vegetation, including filamentous green algae, 1 large 

borrow-pit with greenish water (microscopic green algae) and 1 small 

ground pool with brackish water. In this last breeding place larvae 

of Ae. irritans, C. thaIassius and U. biIineata were found in association 

with guiarti. In the other biotopes the associated larvae were those of 

An. constahl, C. annuIioris, C. univittatus, F. (114.)mimomyiaformis 

and F. (M.) uniformis. 

Fig. 11 shows a guiarti breeding site. 

C. guiarti is thought to be a relatively rare zoophilic mosquito in 

Angola, though it may bite man rarely (49) (54). 

7.5.12--CuIex (Culex) ingrami Edwards, 1916 

The only known Angolan record of C. ingrami is that by Worth 

& Paterson (12), from Caxito, based on four adults (almost certainly 


RIBgIR0, H. & RAMOS, H. Cunha -- <strong>Research</strong> on the mosquitoes of Angola m X 

females). All the Angolan material examined by the present writers, 

here dealt with, is only provisionally identified as C. ingrami Edws. (see 

below). 

MATERIAL EXAMINED. CABINDA, 2 larvae, 2 pupae, 19.VIII. 1970; CA- 

BUTA, 1 larva, 12.VI.1970; CARLAONGO, 1 , I larva, 5.XI.1969; LUCALA 

RIVER, 6 larvae, 28.VIII. 1969; MUMBONDO, 2 9, 2 , 15 larvae, 7-8.V.1970; 

MUXIMA, 17 larvae, 30.1V.1970. 

TAXON, OMIC NOTES. Since the original description of C. ingrami 

(Ghana males) by Edwards (55) and that of the larvae and pupae by 

Macfie & Ingram (56) (57), many larval and some adult (terminalic) 

varieties have been described, though not named, by Galliard (58), 

Edwards (8), Hopkins (24), Rageau & Adam (59), Peters (60), Service (61), 

Pajot (62) and others. It seems quite evident that much of this variation, 

particularly that concerning larvae, must be interpreted as polymorphic 

in nature. Angolan larvae, however, besides exhibiting a marked polymorphism 

of :he kind described by these different authors, constantly 

show the presence of a tergal plate on abdominal segment viii, as described 

byd Pajot (op. cit.) on Cameroon larvae, which separates them immediately 

from ingrami s. str. Moreover, these larvae are associated with 

males whose terminalia also show non-negligible differences from those 

of both C. ingrami s. str. and of Pajot's form. These are the reasons 

why this Angolan form here provisionally determined as C. ingrami is 

thought to be, in fact, a distinct though closely allied taxon, to be dealt 

with in another paper. 

DISTRIBUTION. Map 17 shows the known distribution of the Angolan 

form of C. ingrami s. l. It is assumed that the previous Caxito record 

by B. Worth & Paterson also concerns this Angolan form. 

BIOECDIaOGICAL NOTES. These notes refer exclusively to the Angolan 

form examined by the writers, closely allied to ingrami. This is a ground 

water breeder, as it is the case for all the taxa of the group. Both stagnant 

and running water are used by the present form, as it can be seen from 

the 8 larval biotopes recorded by the writers: niches and pools at river 

edges (3), rain-water ground pools (2), overflown river margins (2) 

and one stream. The breeding water was either turbid or clean, being at 

Estud., E.s. e Doc,- 134 83

RIBgIRO, H. & RAMOS, H. Cunha -- <strong>Research</strong> on the mosquitoes o[ Angola -- X 

times greenish from microscopic algae. Some kind of macroscopic vegetation 

was usualy present in the breedifig water, while the larval biotope 

was often shaded by surrounding vegetation. 

Larvae of 10 Culicines and 4 Anophelines were found as associates, 

those of An. rivuIorum, C. decens and C. inconspicuosus being the 

most common. Fig. 1 shows a larval biotope of C. ingrami. 

Like C. guiarti, both C. ingrami s. str. and the closely allied Angolan 

form seem to be relatively rare mainly zoophilic mosquitoes. 

7.5.13 -- Culex (Culex) invidiosus Theobald, 1901 

C. invidiosus s. str. is a new record for Angola. 

MATERIAL EXAMINED. DUNDO, 1 3, 15 larvae, 19.VIII. 1969; HUAMBO, 

1 3, 1 larva, 24.I.1964; MONA QUIMBUNDO, 4 9, 6 3, 5 larvae, 10.IV. 1970; 

MUXIMA, 1 9, 1 3, 29.IV. 1970; SAURIMO, 1 9, 1 3, 12.IV. 1970. 

TAXONOMIC NOTES. For a general taxonomic discussion of C. invidiosus, 

see var. vexillatus (7.5.14). 

As it is well known, larvae of antennatus, decens, invidiosus s. l., 

trifoliatus and univittatus (larval ornatothoracis remains undescribed) 

ressemble closely one another and it is not uncommon that individual 

specimens cannot be separated. Neverthless, if representative samples 

of the different local populations are available, a specific determination 

becomes often possible, even in the case of the closest allied larvae of 

decens and invidiosus. Within C. invidiosus (s. l.), the larva of ssp. vicinalis 

was not described and that of C. invidiosus s. str. cannot be se- 

parated from that of var. vexillatus. 

In all our invidiosus s. str. male terminalia d, d', e and ! are present. 

Setae d and d' are minutely barbed aplically, e is a narrow delicately 

striate foliole and f is always a larger obviously striate, nearly symmetrically 

expanded foliolar process. 

DISTRIBUTION. C. invidiosus s. l. is a widespread mosquito in Tropical 

Africa, though mainly associated with West African biomes. The 

known Angolan locality records of C. invidiosus s. str. (male records), 

all of them at latitudes less than 13 ø S, are shown on Map 16. 

84 Estud., Ens. e Doc. -- 134


BIOEOOLOGICAL NOTES. Seven larval biotopes were recorded for 

invidiosus s.s.: margins of slow flowing rivers (3), rock-pools (2), a small 

ditch and a cement tank. The breeding water was usually clean, sometimes 

with abundant aquatic vegetation. Larvae of C. tigripes were found 

as associates in two biotopes and C. annulioris, C. alecerts, C. duttoni, 

C. perfidiosus and C. telesilla were recorded once. Fig. 12 ilustrates a 

breeding place of C. invidiosus (typical form). 

In that concerns adults, invidiosus s.s. was never seen biting and only 

one male was caught resting indoors, which seems to point to the 

zoophily (and exophily) of this mosquito. According to Someren et al. (49), 

C. invidiosus (s. l.) may bite man rarely. 

7.5.14--Culex (Culex) invidiosus var. vexillatus Edwards, 1941 

The only published Angolan records of var. vexillatus are those from 

Lobito, by Worth & Paterson 02) and by Ribelto & Mexia (34). 

MATERIAL EXAMINED. CABINDA, 1 /t, 7 larvae, 26.VIII. 1970; L.iN- 

DANA, 2 $, 3 /t plus 3 larvae, 24.VIII. 1970. 

TAXONOMIC NOTES. The male terminalia from Cabinda and Lfindana, 

here attributed to the form vexillatus, differ from those of the type 

form as follows: f more expanded and assymmetrically so, somewhat 

like a flag, and d and d' slightly flattened and delicately striated. Ap- 

pendage e is a relatively narrow, obviously striated foliole, much like that 

of the type form. 

The taxonomy of C. invidiosus s. l. and its allied forms is far from 

being in a satisfactory state. The examination of male terrainalia of 

C. invidiosus Theo. enabled Edwards (8) to describe var. vexillatus from 

Kampala (Uganda) and to establish for this form a distribution (op. cit., 

p. 339) strictly within the West African Subregion of Chapin (63), includ- 

ing Stanleyville (now Kisangani) and Ibadan (Southern Nigeria). At the 

same time, however, Edwards (op. cit., p. 338) recorded males of the 

type form from a larger area within Tropical Africa which included also 

Kampala and Southern Nigeria (Lagos). The sympatry of both the type 

form and var. vexillatus being not compatible with a subspecific status, 

the question remains if var. vexillatus must be trated as a synonym of 

Estud., Ens. e Dot'.- 134 85


invidiosus or as a distinct, closely allied species. And if one considers 

both the kind and degree of the differences between both forms and the 

existence of intermediate stages of development of seta f, we would be 

inclined to the first hypothesis. 

This seemed to be, in fact, the decision implicitly favoured by De 

Meillon & Lavoipierre (32) when these Authors described, in 1944, the 

form vicinalis from Yangambi (near Stanleyville - Kisangani) as a subspecies 

of invidiosus. However, as the type form was recorded by Edwards 

also from this area (Stanleyville), the taxonomic difficulties arise now 

with the subspecific treatment of vicinalis. On the other hand, vicinalis 

being a member of series decens, we cannot be but reluctant as 

to accept such a subspecies differing so drastically from the rioruminate 

one in the number and shape of the coxite lobe appendages. Finally, 

the situation becomes even more delicate if we consider the previous 

description by Edwards (op. cit., p. 340), also from Stanleyville, of a male 

termina2ia quite similar to that of vicinalis. Thus, the areas of Yangambi- 

Stanleyville (Zaire), Kampala (Uganda), Ibadan-Lagos (Southern Nigeria) 

and, now, that of Banana-Cabinda (Congo River mouth) are the 

key-areas for a taxonomic analysis of C. invidiosus and its closest allied 

forms. 

According to the distributional and morphological data available and 

in the absence of mating experiments, the suggestion of the present 

writers is that var. vexillatus be treated as a junior synonim of invidiosus 

and that a specific rank be given both to vicinalis and to the form 

described by Edwards (op. cit., p. 340 and Fig. 118 g) as which in his own opinion (p. 340). 

In the writers' views, the affinities of the different forms (species) 

within series decens of Edwards would be better established by grouping 

them according to the number of appendages in the coxite lobe: group 

decens/invidiosus/ornatothoracis/trifoliatus, with d, d', e and f; group 

perfidiosus/perfuscus, with d, e and f; group of Edwards/vicinalis, with only e and f. The range of 

variation in terminalic and external morphology characters within each 

of these smaller groups is yet in need of extended research efforts. 

DISTRIBUTION AND BIOEOOLOGICAL NOTES. Map 16 shows the known 

distribution of var. vexillatus in Angola. 


RIBEIRO, H. & RAMOS, H. Cunha -- <strong>Research</strong> otz the mosquitoes of Angola X 

No bioecological differences could be put into evidence between var. 

vexillatus and the type form. 

Our adults of var. vexillatus were all reared in the field laboratory. 

Three vexillatus breeding places were recorded by the writers: two 

enlarged crabholes in the mangrove swamp of the Chiloango River mouth 

and a slow flowing cement channel draining muddy water to the sea. 

Aedes argenteopunctatus kingi, C. annulioris and C. tigripes were recorded 

as vexillatus associates in its larval biotopes. 

According to the findings of Hamon (37) in Bobo Dioulasso area, 

concerning var. vexillatus, this would be mainly a zoophilic mosquito. 

7.5.15- CJdex (Culex) ornatothoracis Theobald, 1909 

The only known Angolan record of C. ornatothoracis is that by 

Worth & Paterson (12), from Cabinda. 

MATERIAL EXAMINED. LIFUNE, 1 ex pupa, 21.IV. 1970. 

TAXONOMIC NOTES. Scutum with many yellowish scales round margins 

and on posterior half, as described. Hind femur, however, practically 

without pale spot at tip. Phallosome much like those figured by 

Edwards (8) (Fig. 118- d, e) though the small external tooth present on 

the outer division on type phallosome from Accra (but not figured in the 

Toro specimen) is much more developed in our male. On the coxire lobe, 

/ is a narrow pointed blade, the other appendages being as in type. 

DISTRIBUTION AND BIOEOOLOGICAL NOTES. Map 16 shows the only 

two Angolan localities known for ornatothoracis. 

The larval biotope recorded by the writers for this species was a 

large rain-water ground pool with floating vegetation and shaded by 

palms. The breeding water was turbid, with a high organic matter content, 

and greenish owing to microscopic algae. 

C. ornatothoracis is thought to be a rare zoophilic mosquito in 

Angola, as it is in Kenya (49). Haddow et al. (35), however, recorded this 

mosquito as biting man by day, in the open. 

Extud., Ens. e Doc.- 134 87

RIBgIRO, H. & RAMOS, H. Cunha -- <strong>Research</strong> on the mosquitoes of At,gola -- X 

7.5.16 -- Culex (Culex) perfidiosus Edwards, 1914 


MATERIAL EXAMINED. DUNDO, 10 larvae, 19.VI11.1969; GUNGO, 

3 larvae, 23.XI.1969. MUMBONDO, 2 larvae, 8.V.1970. 

TAXONOMIC NOTES. All our larvae agree closely with the description 

given by Hopkins (24). Head setae B and C are always trifid and the 

mentum has 8-10 teeth, these characters distinguishing larval per/idiosus 

from the larva of schwetzi described by Peters (44). For the separation 

of the larvae of C. per/idiosus from those of C. pipiens forms, see the 

larval key under 6. 

DISTRIBUTION AND BIOECOLOGICAL NOTES. Map 16 shows the three 

Angolan localities known for C. per/idiosus, all of them in Northern 


Six larval biotopcs of per[idiosus wcrc recorded by the writers: 4 

rock-pools with decaying leaves and 2 rain-water ground pools, all of 

them in forest. The pH of the breeding water, measured in a ground 

pool under Laurisilva forest in the $carpmcnt, was 6.2. 

Larvae of 13 other species were found in association with per[idiosus. 

From these, C. decens, C. inconspicuosus, C. invidiosus, C. telesilla 

and C. tigripes were recorded twice, while An. coustani, An. [unestus, 

An. implexus, An. obscurus, An. rivulorum, C. trifoliatus and U. [usca 

were found only once. Figs. 2 and 12 show typical larval biotopes of 

C. perfidiosus. 

As far as we know, perfidiosus has not been reported as biting man. 

7.5.17- Culex (Culex) perfuscus Edwards, 1914 


MATERIAL EXAlVIINED. SAURIMO, 1 , 10.IV. 1970. 

TAXONOMIC NOTES. Our per[uscus female agree quite well with 

known descriptions. It can be separated from female ornatothoracis 

mainly by its entirely dark hind tibia and from female telesilla by its 

shorter upper fork cell, which is only about two times as long as the stem. 


RIBEIRO, H. & RAMOS, I-[. Cunha -- <strong>Research</strong> o. the nosquitoes of Angola -- X 

DISTRIBUTION AND BIOEOOLOGICAL NOTES. Map 16 shows the only 

Angolan locality known for C. periuscus. The unfed female was caught 

resting inside a tent of our camp near the hospital of the town. In Kenya, 

C. per/uscus is also a relatively rare mosquito which, however, may bite 

man rarely (49). In Uganda, Haddow et al. (35) have reported perfuscus 

as not biting man. 

7.5.18- Culex (Culex) pipiens pipiens Linnaeus, 1758 

Though our data on the Angolan members of the piplens complex 

are not in fayour of a subspecific treatment (see also under quinquefasciatus), 

we maintain the nomenclature of Stone, Knight & Stareke (17) 

and Knight & Stone (23) considering the need for a world-wide revision 

of the systematics of the complex, not carried out in the present paper. 

C. p. piplens was firsfly recorded from Angola (Bi6) by Theobald (6) 

in 1907, as Heptaphlebomyia simplex males (Wellman coll.). Other published 

locality records are those by Gfindara (10) (11), from Huambo 

(8, 9) and Capelongo (9, L), and by Ribeiro & Mexia (34), from 

Lobito (8, L). 

MATERIAL EXAMINED. ALTO CATUMBELA, 1 larva, 17.IX.1970; BO- 

COIO, 1 larva, 9.IX.1970; CALAI, 2 9, 6 8, 29 larvae, 15-16.V. 1965; CHAO DA 

CHELA, 1 larva, 29.V.1969; CUANGAR, 8 larvae, 4.V. 1966; CUBAL, 10 9, 

8 8, 5 larvae, 12-14.X.1970; DALA TANDO, 1 9, 1 8, 10 larvae, 27.VIII. 1969; 

EBANGA, 2 8, 4 larvae, 18-19.X. 1970; FOZ DO CUNENE, 1 9, 30 larvae, 

22.V.1969; GUNGO, 1 8, 5 larvae, 22-23.XI. 1969; LUINGA, 1 larva, 1.X.1969; 

MO(AMEDES, 31 9, 36 8,218 larvae, several dates 1967-1969; M'PUPA, 2 larvae, 

9.V. 1966; TCHIVINGUIRO, 1 larva, 29.V. 1969; UIGE, 2 larvae, 25.IX. 1969, all 

new locality records. Also HUAMBO, 15 9, 49 8, 64 larvae, several dates 1964- 

1971. 

TAXONOMIC NOTES. A general taxonomic discussion on the piplens 

complex in Angola will be carried out under C. p. quinque[asciatus. 

The phallosome of the Angolan pipiens sensu stricto is quite typical. 

The inner divisions are stout, strongly divergent, with truncate, striated 

tips. The inner parts of the outer divisions are narrow, subparallel for 

most of their lengths, distally bent at almost right angles and short, their 

points do not reaching to the outer margins of the inner divisions. The 



DV/D ratio (64), SO, seems to be usually negative, this being the case 

for all the apparently undistorted mounts measured by the writers fol- 

lowing Barr's method (65). 

In all pipiens females associated with males, the ratio of the length 

of the upper fork cell to that of its stem ranged from 3.9 to 5.0 (mean 

4.4). This ratio was shown to be a satisfactory criterium for separating 

the females of the two forms () of the piplens complex thought to occur 

in Angola. The scaling of the venter, giving less constant results, was 

abandoned. 

The larvae associated with pipiens males were also examined in 

search for reliable morphological diagnostic characters, according to the 

suggestions given by Knight (67). The subdorsal abdominal hair of 

segments hi-IV was always double in our piplens larvae. However, as 

this hair was also often branched in the larvae associated with quinquefasciatus 

males, this character was abandoned as it is not diagnostic. In 

that concerns the siphon index, it is clear that in the Angolan material 

the highest values correspond to pipiens s. str. The usual marked variation 

of this index, however, makes it a non-reliable diagnostic character, 

much overlap being observed. The same is true, and even in a greater 

extent, in that concerns the length of the anal papillae or the number 

of the pecten teeth, though this one is usually smaller in pipiens. As 

to the shape of the larval siphon, the sigmoidal curvature is often 

marked in pipiens, but it is usually too an elusive character to be relied 

upon. 

The only valuable diagnostic character in Angolan larvae was found 

in the branching of the siphonal tufts. The investigation of this character 

was also suggested by Knight, though being presented as one > (op. cit.). After the examination of many 

thousands of pipiens and quinquefasciatus larvae, it became obvious to 

the writers that the last form has a greater branching of the siphonal 

tufts than has pipiens. However, this character is of diagnostic value 

only if one ignores the distal tufts (the branching of which does not differ 

significantly in both forms) and takes into account only the two most 

(D The term is used in the sense of Knight & Malek (66) and others, 

that is, to designate each of the components of the (pipiens) complex regardless 

of its questionable taxonomic status. 

90 E.vtud., Ens. e l)oc.- 134


proximal pairs of subventral tufts. For these tufts, in the larvae asso- 

ciated with piplens males the mean number of branches per (proximal) 

tuft was 2.4 (range 1.5-3.0), individual tufts being either single or 

with 2-4 branches. This character consistently separated our pipiens 

larvae from those of quinque/asciatus, with only a quite small fraction 

of overlap (see also under quinque/asciatus). 

DISTRIBUTION. C. p. pipiens is widely distributed all over Central 

and Southern Angola (Map 13), while it seems to be at least quite uncommon 

in the West African biomes of Northern Angola (Maps 2 and 3). 

It is to be noted that pipiens s.s. is also absent from the coast N of 

Lobito, the northernmost localities recorded (Dala Tando, Luinga and 

Uige) being at altitudes from about 700 to 1,200 meters, in the Scarpment. 

Being a rare mosquito in the Lobito area (34), pipiens s.s. becomes 

common at Moqfimedes, southwards in the coast, and is apparently 

the only member of the complex at Foz do Curterie, the southernmost 

coastal locality in Angola (mouth of Cunene River). At these higher 

altitudes, pipiens was also relatively common in South-eastern Angola, 

being the only member of the complex recorded from Cuangar, Calai 

and M'Pupa (altitudes about 1,000 meters), along the Cubango River. 

Besides these areas, pipiens s.s. seems to be particularly associated with 

the Scarpment south of the parallel 12 (' S (Gungo, Bocoio, Cubal, Cho 

da Chela, Alto Catumbela), at altitudes from 900 to 1,250 meters, or 

with the Montane biome (Bi, Ebanga, Huambo, Tchivinguiro) from 

about 1,700 to 2,000 meters. And it is perhaps significant that at Ebanga 

(2,000 meters) piplens s.s. was found to be again the sole member of the 

complex present. Summarizing, we can say that pipiens s.s. in Angola 

shows a distribution characteristic of a species belonging to the East-South 

African Subregion, being commoner in the higher latitudes and/or altitudes, 

which is in general accordance with the views expressed by other 

Authors, such as Edwards (8) and Mattingly (68). Finally, we must note 

that the available distributional data in Angola are clearly in fayour 

of the assumption that piplens s.s. is either a native form or, at least, 

an early and quite successful introduced one. 

BI.OECOLOGICAL NOTES. 52 pipiens s.s. breeding places are thought 

to have been recorded by the writers, though 32 of them were only presumptive 

as they were based on the examination of larvae and/or of 

EstutL, Ens. e Doc.- 134 91

RIBgIR0, H. & RAMOS, ]-[. Cunha -- <strong>Research</strong> o. the tnosquiloes of A.gola -- X 

reared females. No breeding dependant upon an inadequate sewage 

disposal nor upon stored water for domestic uses was ever observed by 

the writers in the Angolan populations of piplens s.s. 

The following account, however, refers only to those 20 breeding 

places which could be surely confirmed as being of pipiens s.s. 

through the examination of genitalia of males reared from larvae. 

These 20 breending sites were of three main kinds: ground water (14), 

artificial containers (4) and vegetable containers (2). Ground water 

breeding places consisted of: river margins (6), open shallow earth 

wells (4), vegetated rain-water ground-pools (2), borrow-pits (1) and 

crabholes (1). Artificial containers were used by pipiens for breeding as 

follows: boats in river margins (2), pottery vase with rain-water (1) 

and choked cement drain (1). Breeding in vegetable containers was 

recorded in two instances: in a fallen banana leaf with rain-water and 

in an old abandoned gourd with rain-water and decaying fallen leaves. 

Thus, C. pipiens pipiens seems to be mainly a ground-water breeder 

(65 0er cent of the breeding places) though also exhibiting a marked 

ability for breeding in small, non-earthy containers, usually of vegetable 

origin and not envolved in the stGrage of water for domestic 

uses. In other words, from the evidence available. it seems that in 

Angola the breeding area of pipiens s.s. shows little or none dependence 

upon human activities. 

The breeding water was often clean and limpid, usually with emergent 

and floating vegetation, though several instances were recorded 

of turbid, greenish water full of microscopic green algae and/or with 

fallen leaves and vegetable debris. The presumed high organic matter 

content of such breeding waters, however, seems to be always of vegetable, 

non animal, origin. The pH of ten breeding places ranged from 

5.4-7.0, with a mean of 6.4. The chloride content of the water, measured 

in three larval biotopes, was 0.82, 2.28 and 9.83 g/1 (NaCl), which 

shows some tolerance of pipiens s.s. to brackish water. 

In 5 out of the 20 pipiens s.s. breeding places, no other mosquito 

larvae were found. 17 Culicines and 2 Anophelines were recorded from 

the remaining larval biotopes of pipiens s.s. as associated species. 

C. p. quinquefasciatus (confirmed by male terminalia) was the main 

associate (7 breeding places, 35 per cent of all confirmed pipiens s.s. 

larval biotopes), followed by C. univittatus (3) and C. antennatus, C. decens, 

C. duttoni and C. tigripes (2, each). An. coustani, An. listeri, 

92 Extud., E.s. e Doc.--134

RIBEIltO, H. & RAMOS, H. Cunha -- <strong>Research</strong> ott the mosquitoes o[ Angola -- X 

C. annulioris, C. cinerellus, C. kingianus, C. salisburiensis, C. simpsoni, 

C. thalassius, C. theileri, C. toroensis, C. tri[oliatus, C. tritaeniorhyn- 

chus and E. chrysogaster were found only once in association with 

piplens s.s. 

Only one male and five females (ratio of anterior fork to its stem 

3.9-5.0) were caught as pipiens s.s. adults. From these, one unfed 

female from Cherugema and the male, from Mogfimedes, were caught 

at rest in crabholes (no larvae found). Another unfed female (wing 

ratio 5.0) was caught when biting man outdoors, at Mogfimedes. The 

remainder three females were caught when resting on walls inside 

human habitation, at about 9 h a.m., two of them gorged (one from 

Cubal and the other from Huambo) and the other unfed. 

Unfortunately, the nature of the blood in the stomach of the two 

gorged females was not investigated nor tests for anautogeny and 

eurigamy were carried out. In spite of this, however, the above remarks 

on the larval and adult bioecology of the Angolan pipiens s.s., 

besides showing a marked contrast with quinque[asciatus (see below) 

strongly suggest the absence of the form molestus from Angola. In 

fact, unlike quinque[asciatus, the only other Angolan form of the 

pipiens complex seems to be clearly a non man-associated, zoophilic 

mosquito unable to breed in the sewage system or in surface waters 

contamined by human excreta as ,well as in containers related to domestic 

activities, rarely entering houses and biting man only exceptionally. 

7.5.19 -- Culex (Culex) piplens quinquefasciatus Say, 1823 

As it was said under C. p. pipiens, we maintain here the nomenclature 

of Stone, Knight & Starcke (17), and of Knight & Stone (23), 

though the evidence available to us is clearly in desagreement with 

a subspecific treatment of the two forms of the pipiens complex known 

to occur in Angola. 

C. p. quinque/asciatus was firstly recorded from Angola (Luanda) 

by Colago (9), in 1952, as C. /atigans Wied. . Other published 

locality records are those by Gfndara (10) (11) from Lobito (9), Capelongo, 

Cuando and Moxico (L), and by Worth & Paterson (12) from 

Cabinda and Mogmedes (adults, sexes not mentioned). Other papers 

dealing with quinque/asciatus in Angola are those by Ribeiro & 


RIBEIRO, H. & RAMOS, H. Cunha -- <strong>Research</strong> on the mosquitoes o[ ,qngola -- X 

Mexia (34), from Lobito (9-, 8, L. many hundreds) and by Ribeiro (50) 

(69), from Luanda (9, 8, L, several thousands). 

MATERIAL EXAMINED. AMBRIZETE, 9 ?, 2.VIII.1970; BAIA DOS TI- 

GRES, 8 9, 15 8 plus 8 larvae, 3.VI.1969; BAiA FARTA, 2 ?, 7 8, 12.IX.1964; 

BARRA DO DANDE, 2 9, 2 8, 24.IV.1970; BENGUELA, 4 ?, 12 8, 5 larvae, 

13-14.IV.1971; BOCOIO, 1 9, 1 larva, 8-9.IX.1970; CACUACO, 20 9. 1 8, 8 larvae, 

6.XI.1967, 25.IV.1970 and 23.III.1971; CARACULO, 60 9, 32 8, 14.V.1969; 

CASSOALALA, 3 9, 7.XI.1967; CATETE, I 8, 5 larvae, III-IV.1971; CAXITO, 

2 , XI.1967; CHERUGEMA, 1 9, 4.X.1972; CHINGUAR, 10 larvae, IV.1972; 

CUBAL, 6 9, 2 8, 1 larva, 12-14.IX.1970; DAMBA, 3 9,6 8,6 larvae, 6.IV.1971; 

GUNGO, 5 larvae, 23.XI.1969; HANHA, 6 9, 10 8, 12.IX.1964; HUAMBO, 60 

76 8, 549 larvae, several dates 1964-1971; JAMBA, 3 9, 2 8, 5 larvae, 21.1V.1971; 

LUBIRI, 6 larvae, 30.III. 1972; LUCIRA, 10 larvae, 30.IV.1969; MALANJE, 3 

4 8, 51 larvae, 12-13.IV.1971; MUSSULO, 4 9, 32 larvae, 24.III. 1971; MUXIMA, 

2 9, 29.IV.1970; N'G1VA, 3 ?, 17.II.1970; NOVO REDONDO, 16?, 13 

38 larvae, 26.XI.1969 and 7-8.V.1971; PORTO ALEXANDRE, 5 

14-16.V.1969 and 10.IV.1971; PORTO AMBOIM, 4 9, 1 8, 12. IX.1946; QUILEN- 

GUES, ½ 2, 2 8, 27-29.IV.1791; SA DA BANDEIRA, 9 larvae, 12.IV.1971; SACO 

DO GIRAUL, 1 9, 11.IV.1971; SAURIMO, 13 ?, 15 8, 10 larvae, 11.IV.1970; 

and 10-11.IV.1971; SERPA PINTO, 2 2, 13 larvae, 24.IV.1971; UiGE, 3 

4 larvae, 15.IV. 1971; VILA FLOR, 2 9, I 8, 27.X.1968; VIREI, l ?, 12.V.1969, 

all new locality records. Also CABINDA, 17 ?, 2 8, 11 larvae, 18-21.VIII. 1970 

and 14.IV.1971; LOBITO, 3 9, I 8, 12 larvae, 8.V.1971; LUANDA, 6 

13 larvae, 26.V. 1967 and 8.VI.1967; MO(AMEDES, 70 9, 22 8, 105 larvae, 

X-XI.1967, IV-V. 1969 and 11.IV.1971, and MOXICO, 4 9, 19.IV.1970. 

TAXONOMIC NOTES AND DISTRIBUTION. The phallosome of the 

Angolan quinquefasciatus is quite typical, as judging from the de- 

scriptions and figures of Edwards (70), Belkin (71), Dobrotworsky (72) 

and others. The inner divisions appear shorter and obviously slender 

than in pipiens s. str., with blunt tips, never sharply truncate, and 

usually noticeably convergent or, at most, subparallel. The sickle-shaped 

parts of the outer divisions are large, moderately sclerotized, concave 

plates diverging from each other for most of their lengths. Their 

points largely surpass the inner divisions, so that the DV/D ratio is 

always largely positive, ranging from about 1.0 to 1.5 in the appar- 

ently undistorted mounts measured by the writers (Barr's method). 

In all our females associated with quinque/asciatus males, the ratio 

of the length of the anterior fork-cell to that of its stem ranged from 

2.10 to 3.60 (mean 2.96). As it was stated under pipiens s. str., this 


RIBgIRO, H. & RAMOS, H. Cunha -- <strong>Research</strong> on the mosquitoes of Angola m X 

ratio was thereafter used by the writers as a satisfactory criterium for 

separating females of both Angolan forms of the pipiens complex. 

As to the larvae associated with quinquefasciatus males, only the 

branching of the proximal two pairs of siphonal tufts were considered 

of diagnostic significance (see under pipiens s.s.). For these tufts in 

such associated larvae, the mean number of branches per tuft was 8.5 

(range 5.0-11.25), individual tufts exhibiting 4-13 branches. After the 

examination of our samples of the Angolan larvae of the pipiens 

complex, however, a small fraction of specimens was found, in samples 

without associated adults, in which the mean for these tufts was about 

4. Even if this may indicate a slight overlap in the variation of this 

character in both taxa, such individual specimens usually belonged 

to samples that, as such, had means within the range surely known 

for quinquel'asciatus in Angola (5.0-11.25). 

The preceding notes, showing the marked degree of morphological 

distinctiveness between pipiens s..tr. and quinquefasciatus in Angola, 

are related to the delicate and, as it seems to us, yet unsolved question 

of the tay.nomic status of both forms. In fact, on purely morphologica' 

g:omds, more and greater differences (concerning both adults 

and larvae) are found between these two forms not only than those 

separating mosquito subspecies but also between many of the species 

of the smaller groups within the pipiens group of Edwards. In other 

words: the morphological differences between pipiens s. str. and quinquefasciatus, 

as such, are of the specific level. 

Certainly, taxonomy is much more than morphology and the main 

question arises as to if the spatial distribution of both forms is or is 

not in accordance with the apparent level of the morphological differences. 

It must be said, however, that the purpose of the present writers 

is not to carry out a general discussion on the world-wide distributed 

C. pipiens complex but only to contribute to such discussion 

through the analysis of the Angolan situation concerning members of 

the complex. As it is shown on Map 13, quinquefasciatus is widely 

distributed in Angola, from Cabinda to N'giva and from the coast to, 

at least, Saurimo and Moxico. In the northern half of the country this 

seems to be, by far, the predominant form of the complex, becoming 

its sole representative along the costal belt north of the parallel 12 ø S. 

From this latitude southwards, quinquefasciatus is largely sympatric 

with pipiens s. sir. and, with the exception of the extreme Southern 

Estud., Ets. e Doc.- 134 9,5

RIBIglRO, H. & RAMOS, H. Cunha -- <strong>Research</strong> on the mosquitoes o[ Angola X 

Angola, the records of both forms are intermingled. Moreover, from 

a few Angolan localities both forms (male terminalia examined) were 

recorded at the same time. This simultaneous occurrence of both forms 

(male records) had already been reported in Lobito (34), each form 

occupying there its own breeding place. Though this is largely the rule 

in Angola, in a few instances --such as at Cubal, Huambo and Moffimedes-- 

both pipiens s. str. and quinquefasciatus were seen breeding 

in association in the same breeding place, as it was established by the 

examination of the terrainalia of reared males. 

Obviously, the preceding notes on the distribution of both forms 

of the pipiens complex in Angola are in favour of the specific level of 

distinctiveness and in accordance with the presumed specific rank of the 

morphological differences. 

The question now arises, however, as to the existence of naturally 

occurring h-/bridization between the sympatric local populations of both 

forms and its extent. It must be clearly stated here that no sure 

indication that such hybridization occurs in nature could be found by 

the writers in Angola. What we mean by can 

perhaps be clarified as follows: (a) the terminalia of males of both 

forms occurring simultaneously at the same localities or even reared 

ex larva from the same breeding places could be easily attributed either 

to pitiens s.s. or to quinquefasciatus; this decision was based on the 

well known differences in the morphology of the phallosome (see above) 

and, for the undistorded mounts, also on the determination of the 

DV/D ratio (see above for ranges); (b) no (let us say, 

with DV/D ratios between 0.2 and 0.8) were found, though a marked 

variation was observed in the terminalia of quinquefasciatus from Mofamedes, 

namely in that concerns the shape of the inner division, 

the width of the sickle-shaped division and the appendages of the 

coxite lobe. In fact, if hybridization occurred freely in the southern 

half of Angola (where both forms are largely sympatric) and if hybrids 

had the same rate of survival, it was expected that vast swarms 

of hybrids would be found, rare being the places, if they existed, where 

more or less pipiens and quinquefasciatus terminalia were not 

linked by a continuous gradiant of . By the contrary, 

from the available evidence it seems quite clear that, if hybridization 

occurs in Angola, either it is a rare event or hybrids cannot establish 

themselves in nature, or both. In other words, according to our data, 

96 Estud., Efts. e Doc.- 134


the Angolan populations of pipiens s. str. and quinquel'asciatus seem to 

be reproductively isolated and would be treated as two distinct species. 

Interestingly enough, this was also the same view already expressed by 

an eminent entomologist working in South Africa, who wrote: [J. Muspratt (73).] In fact, the situation in 

Southern Africa (Southern Angola included) is obviously quite different 

from that found in the United States or even in Australia, and it seems 

that the understanding of these differences may be of decisive importance 

to the understanding of the systematics of the pipiens complex 

itself. 

As in all generalized species, the adult external morphology of quinquefasciatus 

is subjected to marked variation which was accounted 

for in the construction of the adult key given above under 6. 

BIOECOLOGICAL NOTES. Data on the bioecology of the quinque- 

/asciatus populations of Luanda and Lobito were already published (34) 

(50) (69). The following account concerns only the unpublished obser- 

vations on quinque/asciatus since then carried out throughout the 

country. 

From the 36 breeding places here recorded (confirmed by the 

terminalia of reared males), 19 were artificial containers, 16 ground 

water biotopes and 1 was a hole in a rotting fallen baobab trunk. The 

artificial containers (16 in peridomestic habitat and 2 others inside 

human habitation) were as follows: barrels and drums with stored 

water for domestic uses (10), old rubber tyres (4), cement tanks and 

drains (3) and ornamental vases (2). As ground water breeding sites 

of quinque/asciatus, the following were recorded: oppen shallow earth 

wells and borrow-pits, usually with turbid, greenish water and decaying 

plants and debris (7), open sewage efluents (5), grassy margins of slow 

flowing streams (3) and a dich with fallen leaves and vegetable debris 

(1). The breeding sites were either completely exposed to sunlight, 

sometimes with a surface temperature of 30øC, or markedly shaded 

or even indoor biotopes. Besides the occurrence of quinquefasciatus in 

association with the sewage disposal, foul water was also common in 

the remaining of its breeding area. On the other hand, there is also 

some ability of this mosquito to breed in brackish water. The salt 


RIBIgIRO, H. & RAMOS, H. Cunha -- <strong>Research</strong> on the mosquitoes o[ Angola -- X 

content of the breeding water was determined in 10 biotopes, ranging 

from 0.4 to 4.15 (mean 2.3) g/1 (NaCl). The FH of the breeding water, 

measured in 10 biotopes, ranged from 5.4 to 8.0 (mean 6.6). 

In 18 out of the 36 quinque/asciatus breeding places here concerned 

no larvae of other mosquitoes were found, while in the remaining 

half of the biotopes larvae of 13 Culex spp., 2 Anopheles spp. and 2 

A edes spp. were recorded as associates. These were as follows, by 

decreasing order of frequency: C. p. pipiens (in 7 instances, confirmed 

by the terminalia of reared males), C. dttttoni and C. tigripes (4, each), 

An. coustani, Ae. aegypti and C. simpsoni form A (2, each) and An. 

listeri, Ae. natronitts, C. cinereus, C. kingiants, C. nebulosus, C. thalassius, 

C. theileri, C. toroensis, C. tri/ilatus, C. tritaeniorhynchus and 

C. univittatus (once). Differently from Luanda where, in 1971, the 

invader A e. aegypti was, by far, the main associate of quinque/asciatus 

(69), it seems that in stabilized situations this role is played by 

pipiens s. sir. in the areas where both forms are sympatric, and by 

duttoni and tigripes in other areas. Anyhow, the characteristic low percentage 

of quinque/asciatus breeding places with other associated mosquito 

larvae seems to be due to the ability of this form to breed in 

waters with a high organic matter content of animal origin, not 

suitable for other species. Under favorable conditions, larval associations 

become relatively more frequent as other types of waters are 

used by quinque/asciatus and, inversely, under stabilized situations of 

heavy competition (e.g., as those following the invasion of a town by 

A e. aegypti), its larval associations decrease with the confinement of 

quinque/asciatus to its more characteristic breeding area of foul 

waters. 

Beyond the data already published on adult quinquefasciatus from 

Lobito and Luanda (op. cit.), the following observations are based on 

358 adults of this form caught throughout the country. From these, 

115 were males while 243 were females, these ones being identified 

as quinque/asciatus on the basis of the ratio of the length of the upper 

fork-cell to its stem (see above). Only 2 males and 6 unfed females 

were caught in the outside: the males and 2 of the females resting near 

their breeding places and 4 females when biting man in the open, at 

dusk. All the remaining 350 adults were caught in human habitation, 

though many catches of other mosquito species were carried out in 

cow-houses and pigsties. Te ratio of the number of males to that of 



females caught inside human habitation (113/237--0.40) is near to 

that already found in Lobito (about 0.30) and Luanda (0.37), suggest- 

ing that this may be an expression of some real bioecological cha- 

racteristic of this form. 

160 out of the 350 indoor caught females were either freshly fed 

or gravid, 58 other females were unfed and the remaining 19 were 

biting man (collectors or residents) at the time of the catch. All gravid 

females, so, were found inside human habitation. Both males and 

females could be found indoors at any time, these ones either resting 

or biting, though there seems to be an evening and a morning peak (34) 

for the indoor density of quinque/asciatus. The main indoor resting 

sites of this form were the bed mosquito-nets (when they existed) and 

*.he walls (of any material), though the ceilings (also of any kind), 

clothes, the underside of beds, furniture, etc., were also often used. 

The present remarks on C. p. quinque/asciatus, confirming the 

close association to man of this mosquito, a true house-pest wherever 

it occurs in Angola, also show the high degree of distinctiveness 

between this form and piplens s. str. in that concerns behaviour and 

ecological niches. Here again, as on the morphological and distributional 

grounds, the available evidence is clearly in accordance with 

a specific treatment of each of the two forms of the C. pipiens 

complex known to occur in Angola. 

7.5.20--Culex (Culex) poicilipes (Theobald, 1903) 

The first records of C. poicilipes in Angola are those by Giles (2) 

from Benguela (as taeniorhynchoides) and by Wellman (3) (4), from 

Biff and Bailundo. Later, it was found at Capelongo, Cuchi, Luanda, 

Mavinga, and Mulondo by Gfindara (10) (ll), at Caxito, Huambo and 

N'Giva by Worth & Paterson (12) and at Lobito by one of us (34). 

MATERIAL EXAMINED. CACUAGO ,2 larvae, 22.V. 1967; CALAI, 62 9, 

23 c plus about 250 larvae, several dates between 19.V. 1965 and 15.VI.1967; 

CARIANGO, 2 larvae, 18.X. 1969; CUANGAR, 6 9 plus a few larvae, 12.IV. 1965 

and 4.V. 1966; CUBAL, 7 larvae, 14-15.IX.1970; DANDO, 1 9, 1.11.1970; DIRICO, 

20 larvae, 8.V. 1965 and 5,V. 1966; FOZ DO CUNENE, 2 larvae, 22.V. 1969; LAGOA 

BANDA, 2 larvae, 8.VI. 1967; LOMBE RIVER, ! 9, 4 larvae, 31.1.1970; MO,- 

Estud., Etas. e Doc.- 134 99


MEDES, 1 larva, 20.X.1967; MUMBONDO, 1 9, 2 larvae, 7.V.1970; MUTANGO, 

13 larvae, 5-12.VI.1965, 2-4.V.1966 and 12.VI.1967; MUXIMA, 5 larvae, 29.IV.1970; 

NANGURA, 2 9, 12 larvae, 9.XII.1965, 26.IV. 1966 and 4.V.1966; PANDA, 

2 9, 21 larvae, 24.IV. 1965, and UECA, 3 larvae, 6.V.1966, all new locality records. 

TAXONOMIC NOTES. C. poicilipes is a quite distinctive species, 

easily identifiable using the adult and larval keys proposed in this paper 

(see under 6). The siphon index in the Angolan larvae (30 specimens 

from several localities examined) ranged from 3.0 to 4.7 (mean 3.8). 

DISTRIBUTION. C. poicilipes is a largely distributed mosquito within 

the Ethiopian Region. According to Edwards (8), it also penetrates 

into the Mediterranean Subregion (of the Palaearctic) through the 

Nile Valley. In Angola, as it is shown on Map 9, poicilipes is mainly 

associated with East African biomes (see also Map 3), a pattern of 

distribution already pointed out by Edwards (op. tit., p. 290). 

BIOECOLOGICAL NOTES. C. poiciIipes is a breeder in stagnant and 

semi-stagnant ground waters. 82 larval biotopes of C. poiciIipes were 

recorded, as follows: shallow margins of slow flowing rivers (31 biotopes), 

backwaters and overflown margins of receding rivers (20), small 

ponds (15), pools in river margins (6) rain-water ground pools (4), and 

borrow-pits, ditches and pools in swamps (two each). The breding water 

is usually clean, with aquatic vegetation and exposed to sunlight. The 

pH, measured in 62 biotopes, ranged from 5.0 to 7.0 (mean 5.8). 

In 10 out of the 82 poicilipes breeding places, no larvae of other 

mosquito species were found. In the remainder 72 larval biotopes, representatives 

of genera Anopheles (in 60 biotopes), Culex (39), Aedeomyia 

(11), Ficalbia (7), Uranotaenia (2) and Aedes (once) were found. 

The larval mosquito species most commonly found as associates of C. 

poiciIipes were, by decreasing order of frequency: Anopheles squamosus 

(in 48 breeding places), An. pharoensis (30), An. ziemanni (24), Culex 

univittatus (21), Aedeomyia furfurea (1l), C. theileri (10), An. coustani 

s. str. and C. antennatus (9 each), An. distinctus (8) and An. argenteoIobatus 

(7). Two poicilipes breeding sites are illustrated in Figs. 6 and 13. 

As to the adults, 52 poicilipes females (but no males) were caught 

by the writers throughout Angola, mainly along the margins of the 

100 Estud., Ens. e Doc. 

134


rivers Lombe, Cuanza and Cubango. 50 of the females were caught 

when biting man in the open, at dusk (from 6.00 to 7.00 p.m.). Another 

female was caught indoors, also landing on man, at 19.00. Finally, 

one unfed female was caught at rest, in the outside, at about 18.30. 

Thus, according to our experience, C. poicilipes must be regarded 

as an important man-biting mosquito in the Angolan savannas, which 

is in accordance with the findings of other Authors in other parts of 

Africa (35) (37). 

7.5.21--Culex (Culex) pruina pruina Theobald, 1901 


MATERIAL EXAMINED. DALA TANDO, 3 9, 9 plus 2 larvae, 

25-26.VIII. 1969. 

TAXOIOMIC NOTES. In the Angolan larvae of the nominal subspecies 

of C. pruina head seta d has 4 or 5 branches and, with the exception 

of the more distal pair, subventral tufts of siphon have 6-10 branches 

per tuft (mean about 8.5). The spiculation of the thorax and abdomen 

seems to be strongly variable. Head seta e is bifid, and not single as 

described by Hopkins (24), f has 3-6 branches and the lateral seta of 

saddle is 3- or 4-branched. Other larval characters, as described by 

Hopkins (op. cit.). 

The adults, including the male terminalia, agree quite well with 

known descriptions (74) (8). 

For further taxonomic analysis, see also under subspecies eschirasi. 

DISTRIBUTION. Map 17 shows the only Angolan locality known for 

C. p. pruina, within the West African zoogeographical Subregion. 

BIOEOOLOGICAL NOTES. Our adults of C. p. pruina were laboratory 

reared. 2 larval biotopes of pruina s.s. were recorded: a sunlit rock-pool 

with greenish water in the margins of the Capecha river, and a shaded 

bamboo stump in a bamboo gallery forest remnant. In both instances 

the breeding water was foul and turbid, with decaying vegetable debris 


RIBgIRO, H. & RAMOS, H. Cunha -- <strong>Research</strong> on the nosquitoes of At,gola -- X 

(

RIBEIRO, H. & RAMOS, H. Cunba -- <strong>Research</strong> oJt the mosquitoes of Angola X 

of the river Zuunga, in a degraded mangrove area about four kilometers 

north of Cabinda. The breeding water was foul, dark and turbid, with 

a pH -- 5.4. Culex annulioris and C. duttoni were recorded as associates 

of eschirasi in this larval biotope. According to these data, it seems that 

both pruina forms would compete for breeding places if they were 

sympatric. 

Our two females of C. p. eschirasi were reared from larvae. However, 

according to Haddow et al. (3'5), subspecies eschirasi may bite man by 

day, in the outside. 

7.5.23- Culex (Culex) simpsoni Theobald, 1905 

Culex simpsoni was firstly recorded by Gfindara (10) (11), from 

Luanda. The only other published record for this species in Angola is 

that of Lobito, by one of us (34). 

MATERIAL EXAMINED. BERO RIVER (N of Virei), 2 9, 1 8 plus 

9 larvae, 9.V.1969; BOCOIO, 6 larvae, 7-8.IX.1970; CAINDE, 5 9, 6 8, 14 larvae, 

12.V. 1969; CALAI, 3 9, 2 8, 1 larva, 19.XI.1965 and 24.V.1967; CARACULO, 

1! 9, 7 8, 44 larvae, 14.V. 1969; CHERUGEMA, 3 9, 7 cS, 4-6.X.1972 (Dr. R. A. 

Capela coll.); CHIQUITE, 2 8, 3 larvae, 27-29.V. 1970; CHITADO, 4 9, 5 8, 

9 larvae, 21.II.1970; CUANGO, 4 9, 2 8, 8 larvae, 3.II.1970; CUANZA RIVER 

(N of Calulo), 2 8, 3 larvae, 12.VI.1970; CUNENE RIVER (S of Espinheira), 

4 9, 4 8, 29 larvae, 20.V. 1969; DANGE-IA-MENHA, ! 8, 5 larvae, 28.VIII. 1969; 

DINDE, 1 larva, 28.V. 1971; DUQUE DE BRAGANnA, 5 larvae, 21.11.1969; 

EQUIMINA, 4 9, 3 8, 9 larvae, 20-22.V. 1970; GUNGO, 3 9, 3 8, 6 larvae, 

22.XI.1969; HUNGUIgRIA, 1 larva, 12.V. 1969; LAGOA DOS PARALELOS, 19 9, 

11 8, 31 larvae, 7.V.1969; LOMBE RIVER (W of Malanje), 1 larva, 11.1970; 

LUCALA, 1 larva, 27.VIII. 1969; LUCIRA, 1 9, 1 8, 17 larvae, 30.IV.1969; 

LUINGA, 2 larvae, 1.X.1969; MACULAMA (NE of Iona), 4 9, 4 8, 45 larvae, 

21.V. 1969; MO(AMEDES, 29 9, 29 8, 77 larvae, 18-26.X.1967, 24-29.IV.1969 

and 7.V. 1969; MUNHINO, 1 9, 2 8, 1 larva, 7.V.1969; MUSSULO, many larvae, 

1972, (Dr. R. A. Capela coll.); N'GIVA, 5 larvae, 23.IV.1972 (Dr. R. A. Capela, 

coll.); NOVO REDONDO, 4 9, 1 8, 3 larvae, 24.XI.1969; PEDIVA, 3 8, 6 larvae, 

6-8.X.1972 (Dr. R. A. Capela coll.); PICO DO AZEVEDO, 8 9, 8 8, 120 larvae, 

5.V.1969; PORTO ALEXANDRE, 1 9, 1 8, 19 larvae, 14.V.1969; QUICUCO, 

1 larva, 28.V. 1971; RIO GANGO, 2 8, 6 larvae, 21.X.1969; SACO DO GIRAUL, 

1 9, 1 8, 3 larvae, 9.V. 1969; SA DA BANDEIRA (20 km W. of), 1 9, 1 larva, 

6.VI.1969; SAO NICOLAU, 8 larvae, 28.IV. 1969; SERRA DA NEVE, 4 larvae, 

29.V. 1970; TOMBINGA, 3 larvae, 28.VIII. 1969; VIREI, 11 9, 8 8, 35 larvae, 

7.V. 1969, and VIREI (25 km E of), 21 larvae, 12.V. 1969, all new locality records. 

Estud., EJts. e Doc. - 134 1703

RIBEIR0, H. & RAMOS, 1-. Cunha -- <strong>Research</strong> o the mosquitoes of Agola -- X 

TAXONOMIC NOTES. Adults of C. simpsoni show little variation and 

agree quite well with Edwards description, including the terminalic characters 

of male. It is perhaps worth to note, however, that the scutal 

pattern described by Edwards is by no means obvious and the decumbent 

scales of head are mostly pale even toward sides. 

In that concerns larvae, on the contrary. much variation of 

apparently different kinds has been reported by Hopkins (24), 

Mattingly (in Hopkins, op. cit.), Mattingly & Lips (42), Mattingly 

& Brown (75), Hamon et al. (76), Mouchet et al. (77) 

and others. Most of the described variation in larval C. ,simpsoni 

was also found by the present writers in Angola. Besides, after 

careful examination of thirty full-grown larvae from eight different 

Angolan localities, it was possible to distinguish two main larval forms 

(A and B), almost always separable on the basis of the characters used 

in the key (see under 6). Besides these ones, the following characters 

may also be useful for distinguishing between the two larval forms: head 

seta C is bifid in about 80 per cent (and single in about 20 p.c.) of 

form A larvae, while these percentages are inverted in larval form B; 

the mean number of pecten teeth in form A larvae is 10.1 (range 8-12), 

while in larval form B we found 11.8 and 10-15 teeth, respectively; lastly, 

the saddle hair has 2-5 branches (mean 3.5) in form A larvae and is 

single, bifid or, at most, trifid (mean 2.4 branches) in larval form B. 

Even if no differences have been found in external and terminalic 

characters between adults associated with each one of the larval forms A 

and B, it is quite apparent that we are dealing with an incipient process 

of speciation within C. simpsoni (s. l.), an hypothesis that seems to be 

confirmed by distributional data in Angola (see below). 

DISTRIBUTION. C. simpsoni is an Ethiopian species known to occur 

in the African mainland, Madagascar, Mauricius, Seychelles (75), and 

S. W. Arabia (38). As it was pointed out by Mattingly & Brown (op. cit.), 

this species seems to be more common and more widely distributed 

throughout East and South African biomes than in West African ones, 

at least according to the available data. In Angola, as it is shown on 

Map 11, C. simpsoni is a quite common and widespread mosquito in the 

South-west Arid Zone and also occurs in the West African Savanna Zone 

(Map 3). A most interesting finding is the apparent association of each 

of the two larval forms described above with each one of these biotic 

104 Estud., Ets. e Doc.- 134

RIBgIRO, H. & RAMOS, H. Cunha -- <strong>Research</strong> ott the mosquitoes of Angola X 

divisions: while form A is associated with the SW Arid biome (East 

and South African Subregion of Chapin), form B is to be found in the 

Southern Congo Savanna Zone (West African Subregion). This allopatry 

of forms A and B of C. ximpxoni emphasize the taxonomic significance 

of the morphological differences found in the larvae and strongly suggests 

a subspecific treatment of both aimpsoni populations. 

BIOECOI.,OGICAL NOTES. 85 breeding places of (7. simpsoni were recorded: 

19 pools in river beds, 18 rock-pools, 13 shallow earth wells, 10 

pools at river edges, 9 borrow-pits, 6 cement tanks and their efluents, 5 

river margins, 2 ponds, an earth drain, a salt-marsh and a hot saline spring. 

These larval biotopes were usually unshaded and often without any macroscopic 

vegetation. The breeding water was usually clean fresh water, 

with a pH (measured in 51 biotopes) ranging from 5.4 to 10.0 (mean 6.4). 

The salt content of the breeding water, determined in 61 out of the 85 

simpsoni larval biotopes, ranged from 0.1 to 23.4 (mean 2.6) g/l (NaCl), 

which points to the remarkable tolerance of larval simpsoni to chlorides, 

already shown in Lobito (34). Larvae of C. ximpsoni seem to be also 

adapted to relatively high temperatures. Breeding water temperatures of 

29øC and 31øC were recorded by the writers, as well as breeding in a 

hot spring at a certainly higher temperature (unfortunately not measured). 

In 11 out of the 85 simpsoni larval biotopes no other mosquito 

larvae were found. In the remainder 74 breeding sites larvae of 16 

species of Culex, 13 Anopheles, 5 Aedes, 2 Ficalbia and 2 Uranotaenia 

were found as associates of C. simpsoni. The following were associated 

species, by decreasing order of frequency: Anopheles listeri (28 times), 

Culex univittatus (17), An. gambiae (15), An. pretoriensix (13), C. ethiopicus 

(10), An. rufipes rufipes (8), An. coustani, An. rhodesiensis rhodesiensis 

and C. tigripes (6 each), An. azevedoi, An. ziemanni, C. thalassius, 

A edes natronius and Uranotaenia fusca (4 each), A e. vittatus, 

C. antennatus, C. decens, C. pipiens pipiens and C. theileri (3), Ae. 

durbanensis angolae, An. longipalpis, An. squamosus, C. annulioris, 

C. pipiens quinque/asciatus and C. poicilipes (twice) and only once, 

Ae. aegypti, Ae. irritans, An. demeilloni, An. nili Congo form, C. antennatus, 

C. aurantapex }iniaensis, C. argenteopunctatus kingi, C. cinerellus, 

C. ethiopicus, C. inconspicuosus, C. perfidiosus, C. tritaeniorhynchus, 

Ficalbia malfeyti, F. mimomyiaformis and U. bal/ouri. Figs. 5 

and 10 show two larval biotopes of C. simpsoni form A. 


RIBEIRO, H. & R^MOS, H. Cunha -- <strong>Research</strong> ou the mosquitocs of /lngola -- X 

Only five females and eleven males of C. simpsoni form A were 

caught in nature, all the remainder adult specimens being laboratoryreared. 

One of the females was caught when biting man in the open, 

at sunset. Two unfed females and eight males were caught early morning 

at rest in earth-holes and rock-holes near simpsoni breeding sites. 

The other two females and three males were caught resting inside 

human habitation, in the afternoon, one of them freshly fed. 

As to the bioecology of each one of the forms A and B of C. 

simpsoni, it is worth to note that all the records of brackish and salt 

water biotopes concern only form A. In relation to this is the occurrence 

of larval form A, and not of form B, in association with some 

halophilic mosquito species, namely Ae. natronius, An. azevedoi, An. 

listeri and C. thalassius. Inversely, some other species, such as C. decens 

and U. /usca, were found only in association with larvae of form B. 

Lastly, the breeding waters with the highest temperatures (and, quite 

probably. also the lowest ones)seem to concern only form A, this 

being related to the occurrence of this form of larval simpsoni in arid 

biomes. All our data on adult C. simpsoni are attributable to form A. 

7.5.24--Culex (Culex) telesilla De Meillon & Lavoipierre, 1945 

C. telesilla was known from Angola only through the larval records 

by Gindara (10) (11), in 1956, from Capelongo (= Folgares) and Luso 

(= Moxico). 

MATERIAL EXAMINED. CALAI, 1 plus 2 larvae, 7.IV.1966, 25.V.1967 

and 6.VI.1967; CUANGAR, 3 larvae, 10-12.IV. 1965; DUNDO, 10 larvae, 19.VIII. 

1969; LIFUNE, 5 9, 5 < plus 6 larvae, 21.IV. 1970; LUANDA, 5 larvae, 5.IV.1971; 

MUMBONDO, 1

RIBI:IItO, H. & RAM08, H. Cunha -- <strong>Research</strong> o, the mosquitoes o) .,gola X 

(group B) of Edwards, there are distinct and completely free d, d' and e, 

besides a much narrower f. Larvae of C. telesilla are quite similar to 

those of C. perfuscus and of other species of this decens series. C. telesilla 

may be separated from all of them, however, owing to the combination 

of the following characters: relatively low siphonal index, 

bifid head setae B and C, antennae infuscated both at base and beyond 

tuft, tufts of siphon and saddle hair with 4-6 branches and comb with 

no more than 30 narrow scales. These were the main characters used 

in the construction of the larval key under 6. The peculiar shape of 

the antennae, as it was described by De Meillon et al. (28), is by no 

means an obvious character in Angolan larvae. 

DISTRIBUTION. C. telesilla is an Ethiopian species described from 

Zaire and also known to occur, at least, in Liberia and Mozambique. 

Map 16 shows all the Angolan localities known for C. telesilla. 

As it can be seen, this is a relatively generalized mosquito in the 

country, though not a much common one. 

BIOEOOLOGICAL NOTES. Twelve larval biotopes were recorded for 

C. telesilla: 5 pools at river edges and overflown margins, 3 rock-pools, 

2 backwaters of receding rivers, an old rubber tyre and a drum. Decaying 

leaves and vegetable debris were often present in the telesilla 

breeding places. 

The following are the mosquito species found in association with 

larval C. telesilla: Culex tigripes (6 times), C. univittatus (3), Anopheles 

squamosus, C. antennatus, C. decens, C. perfidiosus, C. poicilipes and 

Uranotaenia [usca (twice) and Anopheles argenteolobatus, An. gambiae, 

An. pharoensis, Culex invidiosus and U. balfouri (once). Fig. 6 

shows a typical breeding area of C. telesilla in Southern Angola. 

No adults of C. telesilla were caught in the wild by the present 

writers, all our adult specimens being reared from larvae. No records 

are known to us of telesilla as biting man. 

7.5.25-- Culex (Culex) thalassius Theobald, 1903 

C. thalassius was recorded for the first time in Angola by Co- 

laqo (9), from Luanda. The other two Angolan localities for thalassius, 

Estud., Ens. e Doc.---134 ]07

RIBIglRO, H. & RAM08, H. Cunha -- <strong>Research</strong> o the mosquitoes o[ /ttgola -- X 

Lobito and Nova Lisboa, were added by G'findara (10) (11), a few 

years later. The Lobito and Luanda records could be confirmed also 

by Brooke Worth & Paterson (12) and by one of us (34) (50). 

MATERIAL EXAMINED. AMBRIZETE, 4 9, 1 8 plus 10 larvae, 4-5.VI1l. 

1970; BAIA FARTA, 2 9, 12.IX.1964; BARRA DO DANDE, I 9, l 8, plus 

3 larvae, 23.IV.1970; CABINDA, 1 9, 18.VIII. 1970; CUROCA RIVER. 1 larva, 

9.V.1969; EQUIMINA, 5 9, 2 8 plus 2 larvae, 22.V.1970; GUNGO, 1 8, 21.XI. 

1969; MO(AMEDES, 45 9, 16 8 plus 102 larvae, 18-27.X.1967; 23-29.IV.1969. 

2 and 26-27.V.1969 and 10-12.IV. 1971; NOVO REDONDO, 25 9. 14 8 plus 

12 larvae, 20-24.XI.1969; PORTO ALEXANDRE, 14 9, 12 8 plus 58 larvae, 

14-16.V.1969 and 10.IV.1971; SACO DO GIRAUL, 28 9, 19 8 plus 17 larvae, 

9.V.1969; SAO NICOLAU, 1 9, 8 larvae, 8.V.1969; VIREI; 2 8, 27 larvae, 

12.V.1969, all new records. Also LOBITO, 3 larvae, 8.V.1971. and LUANDA, 

I 9, 1 8 plus 4 larvae, X.1969 and 27.IV.1970, besides the data previously 

published concerning these two localities (34) (50). 

TAXONOMIC NOTES AND DISTRIBUTION. Both adults and larvae of 

C. thalassius are quite characteristic and will be easily identified following 

the keys given under 6. 

C. thalassius is widespread throughout the coastal areas of the 

Ethiopian Region, being also known from Syria. 

As it is shown on Map 10, C. thalassius occurs along almost all 

the Angola coast, from Cabinda to Porto Alexandre. Interestingly 

enough, however, this species was not found further south (e.g., at the 

Cunene river mouth) nor it is known from anywhere in the Atlantic 

coast of Africa south of Porto Alexandre. 

BIOECICIaOGICAL NOTES. 36 breeding places of C. thalassius are 

here recorded: 9 salt works, 6 shallow earth wells, 5 borrow-pits, 4 

ground pools, 4 drains and ditches, 3 crabholes, 2 Rhiaophora-Avicennia 

mangroves, and the backwaters of a river, a boat and an old 

rubber tyre. The larval biotopes are usually exposed to sunlight and 

never much shaded, though halophilic vegetation in often present. 

The breeding water may be either clean and limpid or turbid and even 

foul water, with a pH (measured in 17 biotopes) ranging from 6.0 to 

9.0 (mean 6.9). 25 out of the total 36 breeding places were recorded 

either as brackish or salt water larval biotopes. In 21 of them, the 

chloride content of the breeding water was examined, ranging from 

]08 Estud., Etas. e Doc.--134

RIBEIRO, H. & RAMOS, H. Cunha -- <strong>Research</strong> o, the mosquitoes of Angola -- X 

2.9 to 93.6 (mean 29.7) g/1 (NaCl), confirming that thalassius is one 

of the most halophilic Angolan mosquitoes. 

In 7 out of the 36 breeding sites of C. thalassius here recorded no 

larvae of other mosquitoes were found. In the remainder 29 biotopes 

the following species were associated with thalassius, by decreasing 

order of frequency: Anopheles listeri (17 times); An. azevedoi (13), 

Aedes natronius (8), Culex pipiens pipiens (5), Ae. durbanensis angolae 

and Ae. irritans (4 times each), C. simpsoni (3), C. tritaeniorhynchus 

(twice), and An. gambiae, An. pretoriensis, C. guiarti, C. pipiens quinquefasciatus, 

C. theileri, C. univittatus and Uranotaenia bilineata, only 

once. It is quite interesting to note that the species most often found 

as associates of C. thalassius are also halophilic mosquitoes or, at least, 

showing some degree of tolerance as to the chloride content of the 

breeding water. 

Most of our adult specimens were reared in the laboratory, though 

24 females were also caught in the wild. Of these, 6 females were 

caught when biting man in the open, at about 19.00 h., 17 other females 

(15 unfed and 2 freshly fed) were caught at rest inside human 

habitation (including walls of bedrooms) at several hours from 6.00 

a.m. to about 24.00 h. Finally, another female was caught at about 

10.00 a.m., when resting in a rock-hole near salt works. 

These observations seem to confirm the main characteristics of 

C. thalassius, as they were already put into evidence in Lobito (34): 

halophily, anthropophily and exophagy. 

7.5.26--Culex (Culex) theileri Theobald, 1903 

Culex theileri was recorded (as C. creticus) for the first time in 

Angola already by F. C. Wellman (78), from the Bid area. Much later, 

it was again recorded by G5ndara (10) (11), from Bundas, and by 

Brook Worth & Paterson (12), from N'Giva. 

MATERIAL EXAMINED. CALAI, 4 9, 3 plus 69 larvae, 3-11.VI.1965, 

23-24.V. 1967 and 15.VI. 1967; CUANGAR; l 9, V. 1965; DIRICO, 1 9, VI. 1965; 

FOZ DO CUNENE, 2 larvae, 22.V. 1969; JAMBA, 1 larva, 1971; MO(AMEDES, 

5 9,4 plus 9 larvae, 18-26.X.1967; MUTANGO, 1 larva, 12.VI. 1967; NANGURA, 

2 9, 21 larvae, 7-11.VI. 1965; PANDA, 1 9, VI. 1965. All these are new locality 

records. 

E, vtud., Ens. e Doc.- 134 709

RIBgIRO, H. & RAMOS, H. Cunha -- <strong>Research</strong> ot the mosquitoes oj Atgola -- X 

TAXONOMIG NOTES. Notwithstanding the occurrence of some va- 

riation, both adult and larval C. theileri will be easily identified using 

the keys given under 6. Male terminalia are also quite characteristic. 

The following are some particulars concerning Angolan larvae (15 full- 

grown larvae examined, 8 from Mog/tmedes and 7 from the Calai area): 

head setae A usually with 7 or 8 branches (range 5-11), B with 2 

(1-3), C with 3 (2-4), d bifid or trifid, e usually with 5 (3-6) branches 

and / with 7 (6-9) branches; comb of about 30 (18-40) spines; siphon 

index about 4.5 (3.4-5.5); pecten usually with 8 or 9 (6-10) curved 

teeth; subventral tufts of siphon each usually with 8-10 simple branches 

(range 6-12 branches); saddle hair with 2-4 branches, usually bifid; 

gills at times as long as twice the length of saddle, usually not longer 

than the anal segment; eighth abdominal segment obviously spiculate. 

DISTRIBUTION. C. theileri occurs throughout the East and South 

African Subregion (of the Ethiopian Region) and the Mediterranean 

Subregion (of the Palaearctic), extending into the Northern Oriental 

Region (8) (79) (20). In Angola, as it is shown on Map 12, C. theileri 

occurs, in fact, in East and South African biomes, being apparently 

most common in semiarid country. 

BIOECOI_,O(IC:AL NOTES. 24 breeding places of C. theileri were 

recorded: 11 ground pools in overflown river margins, 7 small ponds, 

4 borrow-pits, a shallow earth-well and an old rubber tyre. The larval 

biotopes were usually exposed to sunlight, with little vegetation. The 

breeding water had a pH (11 biotopes examined) ranging from 5.0 to 

8.0 (mean 5.8). In six instances, the chloride content of the water was 

examined, being as follows: 1.4, 2.3, 3.7, 4.2, 7.0 and 9.8. The temperature 

of the breeding water was at times high (30øC were recorded). 

The following mosquitoes occur as associates of C. theileri in its 

breeding places: Culex poicilipes (in 9 out of the 24 biotopes recorded), 

Anopheles pharoensis and An. squamosus (8 times, each), An. coustani 

group (6), Culex simpsoni (5), C. p. pipiens (4), C. tigripes (3) Aedes 

natronius and Culex univittatus (twice, each) and Aedeomyia /ur/urea, 

Anopheles listeri, An. gambiae, Culex decens, C. p. quinque/asciatus 

and C. thalassius, only once. In 3 out of the 24 larval biotopes no associated 

mosquito larvae were found. A typical larval site of C. theileri 

is illustrated in Fig. 13. 

IlO Estud., E,s. e Doc. -- 134


As theileri adults, only 13 unfed females were caught, all of them 

in the act of biting man in the outside, from 17.30 to 18.00 hours. The 

agressive theileri females were caught during May and June, in the 

Calai area, either along the margins of the Cubango river or in the 

villages bordering the flood plaine. Here, however, they were never 

found inside human habitation. 

7.5.27--Culex (Culex) toroensis Edwards & Gibbins, 1939 


MATERIAL IgXAM][NED. NOVA LISBOA, 6 9 plus 4 c, 31.III, 14.IV 

and 13.V. 1970. 

TAXONOMIC NOTES AND DISTRIBUTION. Our knowledge on the 

morphology, variation, distribution and taxonomy of the tri/ilatus sub- 

group (within Edwards' group B or piplens) is still a very unsatisfactory 

and scanty one. This seems to be particularly true in the case of the 

closed allied species C. toroensis and C. vansomereni and their forms, 

arisen during the process of speciation which took place along the 

discontinuous montane biomes of East and South Africa and which cer- 

tainly extended also to Angolan highlands and mountains. 

The form found in Nova Lisboa, at 1,700 m (Map 14), is here 

attributed to C. toroensis as none of the specimens has prealar scales, 

the anterior surface of hind femur may be dark for as much as the 

distal "-/3, and the foliole in the male terrainalia is large, longer than 

seta [. The identification of these Angolan specimens with C. toroensis 

macrophyllus, however, is quite doubtful, as the arfical lobe of coxite 

is deeply divided (as in C. vansomereni vansomereni) and the hind 

femur may be dark for only as little as the distal 1/3. It seems that, 

in fact, the Angolan form should be better treated as a new subspecies 

of C. toroensis. Curiously enough, Mouchet et al. (77) arrived 

at a similar conclusion when studying macrophyllus-like material from 

Cameroon. 

BIOEOOLOGICAL NOTES. Our adult specimens of C. toroensis were 

reared in the field laboratory. Unfortunately, however, all the immature 

Estud., Ens. e Doc. -- 134 11]


material was lost. 3 breeding places of C. toroensis were recorded: 

a sunlit cement water-tank, a shallow earth-well and a borrow-pit, both 

of these well shaded biotopes. The pH of the breeding water, meas- 

ured in two biotopes, was 5.6 and 6.4. Larvae of C. tigripes were found 

as associates in all the breeding places, while the following species 

were recorded only once: Anopheles coustani s. str., An. cydippis, 

Culex argenteopunctatus kingi, C. chorleyi, C. duttoni, C. kingianus, 

C. p. pipiens, C. p. quinquefasciatus, C. trifilatus, and C. univittatus. 

The Angolan form of C. toroensis seems to be a relatively un- 

common zoophilic mosquito of the Humid Montane biome. 

7.5.28--Culex (Culex) trifilatus trifilatus Edwards, 1914 

Only C. trifilatus s. I. was known to occur in Angola, on the basis 

of the finding of four adult specimens (certainly females) caught by 

Worth & Paterson (12) at Nova Lisboa (,: Huambo). 

MATERIAL EXAMINED. CUI, 3 Q plus 2 ex pupa, 15.1II.1970, new 

locality record. Also HUAMBO (= Nova Lisboa), 2 9, 1 3 plus 47 larvae in 

in different instars and a pupa, 15.I11.1970 and 6-13.V.1971. 

T,XONOMIC NOTES. Our C. trifilatus are here identified with 

the nominate subspecies in view of possessing the following characteristics: 

apical lobe of coxite (male terminalia) with setae d and e present, 

as well as a stout hooked seta f; distal third to distal half of the 

outer (anterior) surface of hind femur dark. Larvae are also slightly 

different from those of ssp. aenescens and will be described in another 

paper. 

DISTRIBUTION. C. trifilatus is widely distributed throughout the 

Ethiopian Region, though the nominate subspecies seems to be mainly 

associated with East and South African highland biomes. Map 14 

shows the only two Angolan localities known for C. t. trifilatus (at altitudes 

of 1,700 and 1,800 m). 

BIOEOOLOGICAL NOTES. 15 breeding places of C. t. tri/ilatus were 

recorded, all of them artificial containers in peridomestic habitat: old 


RIBEIRO, H. & RAMOS, H. Cunha -- <strong>Research</strong> on the mosquitoes o1' Angola -- X 

rubber tires (7 biotopes), barrels (5), 2 cement tanks and a discarded 

tin. Curiously enough, this marked man-association of the nominate 

subspecies of C. trifilatus is in contrast with the known breeding preferences 

of ssp. aenescens (24). The breeding water was either clean 

or turbid, dark or greenish water. The pH, measured in one instance, 

was 6.4. 

In 3 out of the 15 larval biotopes no larvae of other mosquitoes 

were found. The associated species in the remainder 12 biotopes were 

the following, by decreasing order of frequency: Culex p. quinque/asciatus 

(in 7 biotopes), Aedes aegypti (5), C. tigripes (3), and Aedes 

marshalii, Culex annulioris, C. duttoni, C. nebulosus, C. toroensis and 

Toxorhynchites brevipalpis, only once. As it would be expected, all of 

these associated species are also known to be able to breed in artificial 

cortainers. Lastly, it is to be noted the absence of tril'ilatus from 

the peridomestic biocoenosis of the coastal belt in Angola. 

C. t. tri/ilatus females seem to take only animal (avian?) blood. 

7.5.29 -- Culex (Culex) tri/oliatus Edwards, 1914 

The only previous record of C. tri/oliatus in Angola was that by 

Worth & Paterson, from Luso (--Moxico) (12). 

MATERIAL EXAMINED. CUNGO, 3 9, ! 3 plus 5 larvae, 23.XI. 1969; 

BOCOIO, 1 plus 3 larvae, 7.IX.1970. 

TAXONOMIC NOTES. Male specimens of C. trifoliatus can be easily 

identified on account of the characteristic leaf-like appendages e and ! 

in the apical lobe of coxite. Examination of the external morphology 

of adults is not always conclusive beyond a group diagnosis within 

series decens (but see key under 6). 

Described the same year by Van Someren (80) and by Ovazza 

et al. (81), the larvae of C. trifoliatus are most able to be confused 

with those of C. univittatus. However, these larvae will be almost 

always separated by the branching of head setae A and C and saddle 

hair, as well as by the denticulation of the pecten spines, these being 

the characters used in the larval key given under 6. In our trifoliatus 

Estud., Ens. e Doc.--134 ]]3


larvae the antenna is infuscated at base as well as from a little before 

tuft to the tip, seta A may have as much as 8 branches, seta d is 

usually single (bifid in about 15 per cent of the instances), the siphon 

index is from 5.5 to 7, and both the saddle hair and the upper caudal 

seta are bifid. In two full-grown larvae there is a dorsal bend of siphon 

near apex, much as it is found in larvae of C. toroensis. The other larval 

characters are as described by the above-cited Authors. 

DISTRIBUTION AND BIOECtOLOGICAL NOTES. C. trifoliatus is an 

Ethiopian species widely distributed throughout Africa South of the 

Sahara, though most records refer to East African biomes. Map 16 

shows the three now known Angolan records for trifoliatux, at altitudes 

of about 900, 1,000 and 1,300 meters. 

All the adult specimens available to the writers were reared in 

the field laboratory. 2 larval biotopes were recorded, both along the 

margins of mountain streams in the Scarpment Zone (Map 3). Larvae 

of Anopheles implexus, Culer annulioris, C. decens, C. perfidiosus, and 

C. p. piplens were recorded as associates of trifoliatus, all of them 

only once. 

C. trifoIiatus is not known to bite man. 

7.5.30-- Culex (Culex) tritaeniorhynchus tritaeniorhynchus Giles, 1901 

The only previously known Angolan record of C. tritaeniorhynchus 

was that from Lobito, by one of us (13). The subspecific name is re- 

corded for the first time. 

MATERIAL EXAMINED. BARRA DO CUANZA, 2 9, 2 3 plus 10 larvae, 

15.IV.1970; MOChAMEDES, I 9, 2 3 plus 19 larvac, 18.X.1967, 28.IV.1969 and 

3.V. 1969; NOVO REDONDO, 2 plus 7 larvac, 24.XI.1969; TCHIVINGUIRO, 

1 3, 29.V.1969. 

TAXONOMIC NOTES AND DISTRIBUTION. It seems that the form 

of C. tritaeniorhynchus occurring in Africa is the introduced type form 

(type locality: Travancore, SW India) brought by human agency from 

the western Indian coasts (8), possibly aldready in historical times. 

In fact, no taxonomically significant differences were ever reported 

114 Estud.. Ens. e Doc. -- 134


between African specimens and those from Western India (83) (24) (84) 

(82). In Angola, also 4 out of the 5 known localities for C. tritaeniorhyn- 

chus are coastal (Map 10). In that concerns the morphology of the 

Angolan specimens (both adults and larvae), no significant disa- 

greements could also be found between our material (including male 

terrainalia) and known descriptions concerning the type form, namely 

the more recent redescription by R. Reuben (84). The only exception 

to be noted (though certainly not significant, given the usual variability 

of this character) is the reduction of the anal papillae in Angolan lar- 

vae (particulary in those from Barra do Cuanza and Novo Redondo) 

which are only - length of saddle, much as described by Kirkpa- 

trick in Egypt (op. cit.). In the male terrainalia, the appendages of 

the apical lobe of coxite (not mentioned in Reuben's redescription) are 

as described by Ovazza et al. (81), from Ethiopian males. 

For distinguishing between C. tritaeniorhynchus and its closely 

allied C. thalassius, see characters used in the diagnostic keys under 6. 

In the female tritaeniorhynchus, the proboscis is usually mostly pale 

(either beneath and on the upper side) for about the basal two thirds. 

BIOEOOLOGICAL NOTES. The following 8 breeding places of the 

ground breeder C. tritaeniorhynchus were recorded, all of them well 

exposed to sunlight: 4 borrow-pits, 2 shallow earthwells and the margins 

of 2 marshy pools. The breeding water was often turbid with pH (six 

biotopes examined) of 7.0 (4 times), 6.7 and 10.0. The chloride content 

(NaCl) of the breeding water, measured in six biotopes, ranged from 

0.5 to 19.4 (mean 5.5) g/l, thus showing the preference of tritaeniorhynchus 

for salt water breeding. 

This ability of the Angolan population of C. tritaeniorhynchus 

for breeding in salt water seems to be related, on the one hand, to the 

above cited reduction of the anal papillae of larvae and, on the other 

hand, to be also reflected in the larval associations of this mosquito. 

The following are the associated species recorded: Culex thalassius and 

Anopheles listeri (4 times each), C. p. piplens (3), An. azevedoi, An. 

ziemanni, C. antennatus and C. univittatus (twice), and Aedes durbahensis 

angolae. Ae. natronius, An. gambiae, C. p. quinque/asciatus 

and C. simpsoni form A (once), five of which are well-known halophilic 

species. It is also perhaps worth to note the frequency of the 

association of the introduced C. tritaeniorhynchus with the autochtho- 


RIBgIR0, H, & RAMOS, H. Cunha -- <strong>Research</strong> on the mosquitoes of Angola -- X 

nous C. thalassius. This strongly suggests competition between both 

closely allied species for basically the same ecological niche, which 

could account for the relative rarety of the invader tritaeniorhynchus 

along the Angolan coasts. 

No adults of C. tritaeniorhynchus were caught as such by the pres- 

ent writers in Angolan, though females are known to bite man, at least 

occasionally (36) (49) (85) (79) (37). 

7.5.31- CuIex (Culex) univittatus Theobald, 1901 

C. univittatus (s. I.) was recorded for the first time in Angola by 

Wellman 8,: Fay (5), from Bailundo. Other early records are those by 

Gamble (7), from Sgto Salvador do Congo, and by Edwards (8), from 

Bie and Benguela. In his revision, Grindata (lo) (11) added Caconda, 

Capelongo, Cuchi, Duque de Braganqa and Nova Lisboa as new loca- 

lities for C. univittatus, this species being also recorded some years 

later from Caxito and Luanda by Worth 8,: Paterson (12) and Ri- 

beiro (50), respectively. Meanwhile, C. univittatus var. neavei had been 

recorded by one of us (13) from Lobito. 

MATERIAL EXAMINED. BARRA DO CUANZA, 2 larvae, 15.IV.1970; 

BIBALA, 28 larvae, 2.V.1969; BERO RIVER (N of Virei), 1 larva, 12.V.1969; 

BOCOIO, 5 2, 6 / plus 7 larvae, 7-10.IX.1970; CABUTA, 1 larva, 11.VI.1970; 

CAFIMA, 1 larva, 20.IV.1972 (Dr. V. M. R. Casaca coll.); CAINDE, I $, 3 ½ 

plus 28 larvae, 12.V.1969; CALAI, 108 9, 122 ½ plus 278 larvae, several dates 

from 27.IV.1965 to 15.VI.1967; CANGANDALA, 1 ½ plus 2 larvae, 13.11.1970; 

CAPUNDA, 2 larvae, 16.VI.1970; CARACULO, 1 $, I plus 59 larvae, 11-14.V. 

1969; CHO DA CHELA, 4 larvae, 29.V.1969; CHICALA, I , 1 , 17.VI.1970: 

CHIEDE, 1 larva, 21.IV.1972 (Dr. V. M. R. Casaca coll.); CHINGUAR. 9 larvae, 

28.IV.1972 (Dr. V. M. R. Casaca coll.); CHITADO, 2 , 5 plus 5 larvae, 21.II.1970; 

CUANGAR, 16 larvae, 10.IV.1965; CUBAL, 11 9, 5 ½ plus a few larvae, 

12-14.IX.1970; CUROCA RIVER (N. of Porto Alexandre), I , 10.V.1969; 

DIRICO, I larva, 5.V.1966: EBANGA, 7 , 8 plus 13 larvae, 17-19.IX.1970; 

EQUIMINA, 5 larvae, 21-22.V.1970; GIRAUL RIVER, 2 , 3 plus 13 larvae, 

7.V.1969; IMPULO, 4 larvae, 22-23.II1.1970; INHUCA RIVER, 3 larvae, 20.VllI. 

1970; IONA, 1 $, 1 plus 37 larvae, 19-21.V.1969; LAGOA DOS PARALELOS, 

I , 7.V.1970; LANDANA, 4 larvae, 26.VIII.1970; LUELE RIVER (Source of), 

3 larvae, 13.IV.1970; LUSO (= MOXICO), 1 larva, 16.IV. 1970; MOCAMEDES, 

18 , 17 ½ plus 26 larvae, 20.X.1967, 23.IV.1969, 2-7.V.1969 and 11.1V.1971; 

M'PUPA, I , 3 plus 52 larvae, 9.V.1966; MUMBONDO, 5 larvae, 7.V.1970; 



MUNHINO (20 km W), 1 9, 1 c, 7.V.1969; MUTANGO, 14 larvae, 9.V1.1965: 

4.XI1.1965 and 2-3.V.1966; NANGURA, 1 c plus 4 larvae, 14.VI. 1965, 9.XII. 1965 

and 2.V. 1966; N'GIVA, 6 larvae, 23.IV. 1972 (Dr. V. M. R. Casaca coll.); PANDA, 

6 larvae. 13.IV.1965 and 2.IV. 1966; PICO DO AZEVEDO (Pedras Salvadoras), 10 

larvae, 10.V.1969; QUELA, 3 larvae, 29.I.1970; QUIMBANGO, 6 larvae, 15-16.VI. 

1970; QUISSOL, 2 larvae, 21.1.1970; SO NICOLAU, 13 larvae, 28.IV.1969; 

TCHIVINGUIRO, 6 larvae, 29.V.1969; UiGE, 1 9, 2 plus 3 larvae, 25.IX.1969; 

VIREI, 3 9, 3 plus 13 larvae, 7-9.V. 1969, all new locality records. Also CEI- 

LUNGA (circa Bi6), 1 c plus 13 larvae, 11.IX.1969; LUANDA, I larva, 8.IV.1971, 

and HUAMBO (= NOVA LISBOA), 59 9, 47 plus 56 larvae, 24-25.I.1964, 

11o19.IX.1965, 9.XII. 1965, several dates in 1966 and 1970, and 10.V.1971. 

TAXONOMIC NOTES. Since the description of C. univittatus, C. perexiguus 

and naevei, as distinct full species, by Theobald (6) (86), the 

taxonomic status of these closely allied and largely sympatric forms 

has been subjected to much controversy by Edwards (87) (8), Mattingly 

(68), Senevet et al. (88), Mattingly & Knight (85), Jupp (89) (90), 

White (91), and others. In the World Catalog by Stone, Knight & 

Starcke (17) the views of Edwards in this matter were followed, C. perexiguus 

being treated as a junior synonym of C. univittatus, while C. 

neavei was sunk into > status. In the new edition of the World 

Catalog by Knight & Stone (23), however, C. neavei is given again 

a full specific rank, in agreement with the views expressed by Jupp. 

On the other hand, in the more recent paper by White (91), not only 

neavei but also perexiguus are tentatively reinstated to full species rank, 

a key being provided to the adults of the three African species of this 

univittatus group (op. cit., pp. 321-322). 

In Angola, as it was to be expected on distributional grounds, 

form perexiguus does not occur, which could be stated through the 

examination of the well-developed on the lateral aedeagal plate 

in a sample of 17 males from several localities. 

In that concerns univittatus s. str. and neavei, both Jupp and White 

agree that females may be separated by the presence, in the former, of 

a complete pale stripe along the anterior surface of mid femur. This same 

criterium is used by White also for separating the male specimens, while 

Jupp prefers the shape of the foliole on the apical lobe of coxite, 

expressed as his >. 

In 6 (3 , 3 ) out of the 48 (22 , 26 ) specimens examined from 

several Angolan localities, there was a practically complete anterior pale 



stripe on mid femur. However, reared from the same batch of larvae 

(same breeding place), there were also other females and males with 

practically all possible gradations, from the absence of even a faint 

indication of a stripe to a well-marked one interruped only a little before 

the knee spot of the mid leg. As to the foliole on the subapical lobe of 

coxite, the mean in 23 Angolan males examined was 48, 

with range 37-60. The indices found in 3 of these males exhibiting 

striped mid femora were, in fact, of the (45, 45 and 48). 

However, indices such as these and even lower were also recorded 

in other 10 out of the 23 males examined (37, 38, 38, 39, 41, 45, 46, 47, 

47 and 48), all of them whithout a complete stripe and most of them 

with no more than a faint indication of it. 

The shape of seta f on the subapical lobe of coxite and the extent 

of the dark scaling of hind femur expressed as a were 

also considered by Jupp as useful characters to separate C. univittatus 

from C. neavei. According to our experience, however, this seems not 

to be also the case in Angola. In that concerns seta f, it was found to 

vary markedly in our sample of 23 Angolan males, from practically non 

modified setae (or only very slightly so), in about half the specimens, 

to setae with conspicuously swallen tips, in four of the males. However, 

no useful correlation could be found between the shape of f and the 

leaflet index, the most markedly modified setae belonging to males with 

indices ranging from 45 to 54, while poorly modified setae (if at hall) 

were found with both the lowest (37) and the highest (60) indices. The 

mean fernoral index was 63 (range 48-83) in the 22 Angolan females 

examined, and 65 (range 40-94) in the 26 males. According to the 

figures found by Jupp for this index and to his taxonomic interpretation 

of them, 36 out of the 48 Angolan specimens examined should 

be neavei (17 females in 22 and 19 males in 26) arid only one male 

(index 94) should be identified as univittatus (s.s.), the remainder specimens 

falling into the area of overlap. Again, no reliable correlation was 

discernible between fernoral indices and the scaling of mid femora or 

the leaflet indices. Namely, none of the 6 specimens with striped mid 

femora had femoral indices surely of the (females: 77, 

71 and 55; males: 76, 60 and 54), while the only male with index over 

92 the upper limit of neavei range, according to Jupp) was pratically 

devoided of pale scales on the anterior surface of mid femora. In turn, 

males with leaflet indices 48 and less had fernoral indices 40 to 87. 


RIBEIRO, H. & RAMOS, H. Cunha -- <strong>Research</strong> ot the mosquitoes of Azgola -- X 

According to the preceding notes, it is the view of the present 

writers that C. univittatus Theo. sould be treated as a polymorphic 

species, forms neavei and perexiguus being of the infrasubspecific level. 

Ribeiro et al. (92), in a quite recent paper dealing white Portuguese mosquitoes, 

have also arrived to the conclusion that form neavei would be 

better treated as an infrasubspecific variation of C. univittatus Theo. 

Other important characters of the Angolan adult population of C. 

univittatus are as follows: postspiracular scales present (and numerous) 

in all the 48 specimens (22 , 26 ) examined; anterior surface of hind 

tibia with an obvious pale stripe in 42 out of the 48 specimens, absent 

or faint only in 1 female and 5 males; spot at apex of hind tibia conspicuous; 

a few, at least, pale scales present at base of costa; ornamentation 

of the abdominal tergites and sternites markedly variable, much 

as described by Edwards (8). 

In that concerns larval univittatus it is also known to be subjected 

to much variation, though no larval forms were ever described. The 

main characters of Angolan larvae (30 specimens examined, from the 

same breeding places as the above studied reared adults) are as follows: 

head setae A usually with 7 or 8 branches (range 6-9), B with 2-4 branches, 

C usually with 3 or 4 branches, occasionally bifid, and d always 

single; comb of abdominal segment viii with 32-65 scales (mean 46); 

siphon index 4.5-7.5 (mean 5.5), usually 5-6; pecten with 8-14 spines, 

mean 11.3; mean number of branches in siphonal tufts about 3, though 

individual tufts may be reduced to a single seta; saddle hair with 2-6 

branches, usually trifid. The variation found on larval univittatus from 

Angola was also of an obvious infrasubspecific kind. 

DISTRIBUTION. C. univittatus is widely distributed in the Ethiopian 

Region, with two other isolated refuge populations, a western Mediterranean 

one, known from Portugal (93), Spain and the Tell of Algeria (88), 

and an eastern Mediterranean population which extends through Middle 

East to the North-west of India. 

In Angola, as it is shown on Map 12, C. univittatus is also a widely 

distributed species, the most generalized and one of the most common 

Angolan mosquitoes. 

BIOECIOIaO(ICAL NOTES. Larval univittatus was recorded from 150 

different breeding places, 136 of which (90.7 per cent) were ground 

Estud., Er'. e Doc.--134 119

RIBIglRO, H. & RAMOS, H. Cunha -- <strong>Research</strong> o the mosquitoes o[ Agola -- X 

water biotopes, while 8 (5.3 per cent) were artificial containers, and 6 (4 

per cent) were rock-pools. Ground water biotopes consisted of margins of 

rivers and streams (38), ground pools from rains or floods (35), ponds 

(16), earth drains (15), shallow earth-wells and water holes (13), pools 

in river beds (13), borrow-pits (4), and coastal salt-water pools (2). As 

artificial containers, were recorded cement tanks (3), barrels with water 

for domestic uses (2), old rubber tyres (2) and the bottom of a boat. 

The breending conditions of C. univittatus in all these sites were quite 

varied, as it was to be expected of a mosquito occurring from the semidesert 

in the South-western Angola to the gallery forest in the North of 

the country. The breeding water was either limpid or turbid, high organic 

matter contents being presumed in 34 of the biotopes. Exposition to sun 

light and accompanying aquatic vegetation were also extremely variable. 

Temperatures of 31øC were recorded in semiarid country. The chloridecontent 

of the breeding water was usually quite low, though in a few 

instances sal-water breeding was recorded, including what was thought 

to be a sea-water ground pool (salinity not measured). In the 35 biotopes 

in which the salt content was determined a mean of 1.I g/1 (NaCl) 

was found, with a range 0.1-15.8. The pH of the water, examined in 

74 larval biotopes, ranged from 4.9 to 8.0 (mean 6.0). The marked preference 

of C. univitlatus for acid waters was put into evidence, as this was 

the case in 63 of the breeding places examined (85 per cent), while the 

water was neutral in 9 of them and had pHs of 7.5 and 8.0 in the 

other two. 

In 10 out of the 150 univittatus breeding places no associated mosquitoes 

were found. As to the remainder 140 biotopes, larvae of other 

species of Culex were present in 103 of them, while Anopheles spp. 

occurred in 84 and A edes spp. were present only in 6. On the whole, 56 

species of mosquitoes were found in association with C. univittatus in 

its breeding sites: 24 species of Culex, 23 Anopheles, 4 Ficalbia, 3 Aedes, 

1 A edeomyia and 1 Uranotaenia. The species most often recorded as 

associates of larval univittatus were, by decreasing order of frequency: 

Anopheles gambiae (29 times), Culex tigripes (28), An. squamosus (24), 

C. antennatus (23), C. poicilipes (22), An. coustani s.s. and C. simpsoni 

form A (15 times each), An. pharoensis and C. annulioris (13 times each), 

An. listeri (12), C. p. pipiens (11), An. pretoriensis and An. ziemanni (9 

each), An. rhodesiensis and An. rufipes (7 each), An. [unestus s.s. and 

C. duttoni (6 each), Aedeomyia furlurea and Ficalbia mimomyiaformis 

120 Estud., Etzs. e Doc.- 134

RIBEIRO, H. & RAMOS, H. Cunha --- <strong>Research</strong> ou the mosquitoes o[ Agola X 

(5 each), and An. argenteolobatus, An. cyddipis, C. argenteopunctatus, 

C. decens, C. telesilla and Uranotaenia bal[ouri (4 times each). The halophilic 

mosquitoes A edes durbanensis angolae, C. thalassius and C. tritaeniorhyinchus 

were also recorded as associates, each in two of the univittatus 

breeding sites. Figs. 5, 6 and 7 illustrate three typical breeding sites 

of C. univittatus. 

Only 13 9 and 5 were caught as adults, the remainder of our specimens 

being laboratory-reared. Of them, 8 9 and 4 were caught at rest 

on several kinds of outdoor shelters. Other catches were made by early 

morning (6.00-8.00) either in cattle-sheds (2 ?) or in human habitation 

(3 9 --one of them fed, in a window trap--and 1 ). Two of these 

three females were taken at Calai (inland plateau) while the other one 

was caught at Bocoio (ScarpmenU, both localities being at about 1,000 

meters in altitude. These observations, showing the predominant zoophily 

of C. univittatus in Angola, seem to be in general agreement both with 

the findings of Someren et al. (49), in Kenya, and with those of 

Hamon (37), in the Upper Volta. 

7.5.32 --- Culex (Culex) watti Edwards, 1920 

C. 4,atti was not found by the present writers. 

The only known records of C. watti in Angola are those by G'andara 

(lo) (l), from Dala Tando (14 reared males and females), Portugtlia 

(circa Dundo) and Caconda (1 female each). 

So far as we know, with the exception of those by Gandara, no 

other new distributional data on C. watti were reported anywhere since 

the original ones given by Edwards himself (8). As it was noted above 

under C. dttttoni, the differences used by Edwards to distinguish watti 

from duttoni seem to be more of the kind of an infrasubspecific variation. 

In favour of this view is also the assumption of Hopkins (24) that 

the larvae of watti (even though not yet described) are

RillgiRO, H. & RAMOS, H. Cunha -- <strong>Research</strong> ot, the mosquitoes of Angola -- X 

quite well with the description of watti. The associated males and/or 

larvae, when they exhisted, were, however, invariably quinquefasciatus. 

7.5.33--Culex (Culex) weschei ? gediensis Edwards, 1941 

C. weschei is a new record for Angola. 

MATERIAL EXAMINED. CANGANDALA (24 km SE), 7 larvae, 10.11.1970; 

MALANJE, 7 larvae, 1.11.1970. 

TAXONOMIC NOTES. Larval C. weschei is highly characteristic, no 

other larva being known with a long pecten and a comb of spines. 

However, our larvae disagree in some aspects with the description of the 

nominate subspecies (24), much as the Kenya larvae tentativelly attributed 

to ssp. gediensis by Someren et al. (49). Namely, Angolan larvae have 

the extraordinary long, stout and basally branched spines on the dorsal 

valves of siphon described by these Authors, similar to those of 

C. argenteopunctatus khgi. Other characteristics exhibited by our larvae 

are as follows: preclipeals dark, flattened, blunt-tipped, and somewhat 

spatulate; head seta A with about 10 (7-13) plumose branches, B bifid 

and C usually (76 per cent) single, sometimes also bifid; d single, e and f 

with 3 or 4 and 3-5 branches, respectively; mentum with 4 major teeth plus 

1 or 2 smaller ones each side of the central tooth; comb with about 6 

(4-8) spines; siphon darkened for the distal 1/2-'/,, with index 6-8; 

subventral tufts consisting of 3-6 pairs of single, delicate setae of different 

lengths, some of them very long; spine on dorsal valve of siphon 

longer than diameter of this at apex, with basal branches or denticles of 

increasing lengths; ventral valves with long hooks; upper caudal seta 

laterally branched somewhat beyond base, with 1-3 secondary branches; 

lower caudal seta single; lateral saddle hair usually bifid (80 per cent), 

sometimes trifid. Remainder characters as described for the nominate 

subspecies. As no adults were available, we are also only tentatively 

identifying the Angolan larvae with subspecies gediensis. 

The larval pelt described by Pereira (94) from Mogambique (Mas- 

singa, Sul do Save Province), was certainly of the nominate subspecies, 

as no mention was made of the spine of the dorsal valve of siphon. 

122 r:stud.. Ets. e Doc.- 134


DISTRIBUTION. C. weschei seems to be a relatively rare though 

widely distributed mosquito in Tropical Africa, from the Atlantic to 

the Indian coasts. 

Map 17 shows the only two Angolan localities known for this species. 

BIOEOOLOGICAL NOTES. Larval C. weschei bred along the margins 

of a pond with water-lilies and other aquatic vegetation, and in a sunlit 

grassy ground pool with turbid water. The organic matter content of 

both breeding places was apparently high. 

Larvae of Anopheles squamosus were also found in both biotopes. 

Other mosquito larvae present in one or the other of the breeding places 

were: Aedeomyia africana, Anopheles coustani s. str., Culex grahami and 

C. inconspicuosus. 

C. weschei gediensis was reported by Someren et al. (op. cit.) as biting 

man rarely, in bush. 

7.5.34--Culex (Culex) zombaensis Theobald, 1901 

C. zombaensis was known to occur in Angola only through the larval 

records by Gfindara (10) (11), from Capelongo, Chissamba and Mulondo. 

MATERIAL EXAMINED. CUBAL. 2 9, 2 plus 5 larvae, 14.IX. 1970; 

HUAMBO, 1 larva, 25.I.1964; QUISSOL, 6 larvae, 23.I.1970. 

TXANOMIC NOTES. In our zom.baensis larvae, head seta B is almost 

always bifid, C has 3-5 branches and d is either single or bifid, the mentum 

has 7 somewhat pointed teeth each side the median tooth, the comb 

elements are either narrow scales or spines [not mentioned in Wolfs' description 

(94)] with small median teeth and well developed basal denticles, 

the siphon index is 6.5-8, the saddle hair has 3 or 4 short branches, the 

upper caudal seta has 4-6 branches, and the anal papillae are from about 

as long as saddle to twice its length. 

Larvae of C. zombaensis are very similar to those of C. theileri and 

C. univittatus. In the larval key given under 6, the separation of these 

species was based on the character of the comb elements, this being the 

criterium already adopted by Mattingly (27) in his correction to the key 

by Hopkins (24). In the writers' experience, C. zombaensis can be sepa- 


RIBEIR0, H. & RAMOS. H. Cunha -- <strong>Research</strong> o. the mosquitoes of A.gola -- X 

rated from C. univittalus also by the shape and denticulation of pecten 

spines, as it was presumed by Mattingly & Lips (42) and confirmed by 

Someren (80) (p. l 1). On the other hand, the antennae of zombaensis are 

darkened throughout also in Angolan specimens, while the antennae of 

theileri and uivittatus are infuscate only at base and beyond tuft, a 

difference already pointed out by Ovazza et al. (81). 

DISTRIBUTION. C. zombaensis is widely distributed throughout East 

Africa. In Angola, according to the available information (Map 14), it 

would seem that this mosquito is mainly associated with Mountain and 

Highland biomes (cf. Map 3). Anyhow, all the six Angolan localities 

known for C. zombaensis are at altitudes from about 900 to 1,700 meters. 

BIOEOOLOGICAL NOTES. All our adults specimens were reared in the 

camp laboratory. 

4 breeding places of C. aombaensis were recorded: two niches in the 

margins of slow-flowing rivers (one of them with much Pistia sp.), a pond 

and a borrow-pit. The pH of the breeding water was 6.8 in one instance, 

while on other larval biotope the organic matter content was presumably 

high owing to the presence of many decaying vegetable debris. Larvae 

of the following species were also found as associates of C. aombaensis: 

Anopheles coustani s. str. and Culex antennatus (twice each), and An. 

funestus, An. gambiae, An. squamosus and C. univittatus (once). A typical 

larval biotope of C. zombaensis is shown in Fig. 14. 

C. zombaensis is not known to bite man. 

124 Estud., E,s. e Doc.- 134

8-- RELATION TO DISEASE 

Genus Culex as a whole seems to be mainly associated with birds, 

though species within the genus range in host preferences from those 

feeding on cold-blooded vertebrates (amphibians) to that species which 

are persistent man-biters. On the other hand, members of genus Culex 

are quite often involved in the transmission of arthropod-borne animal 

viruses, which seems to be a particular trait of mosquitoes in general 

when compared with other arthropods (95). 

According to their position in the antropophily-zoophily gradient, 

the Angolan species of Culex may be divided into four main groups: 

(a) the small group of the markedly antropophilic species, with C. p. quinque/asciatus, 

C. thaiassists, and C. poicilipes,' (b) the group of the indifferent 

species, that is, those which easily bite both man and animals, including 

C. rubinotus, C. theileri, C. aurantapex jinjaensis, C. simpsoni (form A) 

and C. bitaeniorhynchus; (c) those species which undoubtly prefer animal 

blood but have also been recorded as biting man occasionally, with C. annulioris, 

C. antennatus, C. argenteopunctatus kingi, C. cinereus, C. duttoni, 

C. guiarti, C. horridus, C. inconspicuosus, C. ingrami, C. invidiosus, 

C. ornatothoracis, C. per[uscus, C. p. pipiens, C. pruina, sspp., C. t. tritaeniorhynchus, 

C. univittatus, C. watti and C. weschei gediensis,' (d) the 

group of those zoophilic species which were never recorded as biting man, 

even occasionally, including the remaining ten species of subgenus Culex 

and most (nine out of thirteen) species of the other minor subgenera. 

The first group of species includes some of the best known nuisance 

mosquitoes in Tropical Africa, often becoming true pests, at least in 

certain areas. Besides, C. p. quinquefasciatus is also one of the mosquitoes 

most often associated with the circulation of arthropod-borne animal 


RIBEIRO, H. & RAMOS. H. Cunha -- <strong>Research</strong> o the mosquitoes of Agola -- X 

viruses in nature (96) (97). In Africa, this was shown to be the case of the 

Germiston, Sindbis, Wesselsbron, West Nile, Banzi, Ingwavuma and 

Olifantsvlei arboviruses, the first four ones known to occur also among 

human populations in Angola (98). C. thalassius, in turn, was shown by 

Kerr (99) to be a potencial vector of the Yellow Fever virus. 

Species of the second group may be particularly important in conveying 

viruses from the wild animal reservoir into human communities. 

Banzi, Germiston and Witwatersrand arboviruses were isolated from 

C. rubinotus in South Africa, while C. theileri is a proven carrier of Germiston, 

Rift Valley Fever, Shuni, Sindbis and West Nile viruses. 

AB viruses are also associated with some species in the third group. 

Acado virus was isolated from a mixed pool of C. antennatus and C. univittatus 

in Ethiopia, while univittatus was found to be naturally infected 

in Africa with the viruses Ingwavuma, Mossuril, Sindbis, Spondweni, 

Usutu, Wesselsbron and West Nile and was shown to be an experimental 

vector of Banzi, Sindbis and West Nile arboviruses (100). C. p. pipiens, in 

turn, was found naturally infected with Olifantsvlei virus in South Africa, 

Sudan and Ethiopia, with Tahjna virus in Kenya, Moambique and 

Europe and, quite recently, with viruses Sindbis and West Nile, also in 

South Africa (101). 

In the fourth group of species, Sindbis arbovirus was isolated from 

wild C. tigripes females in the Transvaal. 

Lastly, let us mention that in the Oriental Region C. p. quinque/asciatus, 

C. bitaeniorhynchus and C. tritaeniorhynchus are known carriers 

of other AB viruses, as well as vectors of human filariasis (36) (97) (102). 

On the other hand, the role of C. p. quinque/asciatus as a secondary 

vector of bancroftian filariasis in Tropical Africa is also now well established 

(103). 

126 Eslud., Ens. e 1)oc. - 134

ACKNOWLEDGEMENTS 

The writers are greatly endebted to the Medical Entomology team 

of the former Instituto Provincial de Safide Pfiblica, Angola, namely to 

Dr. C. A. Pires, Dr. R. A. Capela, Mrs. M. T. Casaca, Mr. E. Vasconcelos, 

Mr. Chipindo Bernardo, Mr. M. Vassuavava and Mr. E. Chivimbe, 

as well as to Mr. L. Madruga and Mr. A. Gomes. 

The multiple assistance then received from the late Director of the 

Instituto, Dr. V. M. R. Casaca, also made possible this paper. 

We are very grateful to Dr. A. de Barros Machado, then still the 

Director of the Laborat6rio de Biologia do Dundo (Diamang, Lunda), 

for his kind cooperation and helpful advices during our stay at Dundo. 


SUMMARY 

In this paper, the writers revise our present knowledge on the genus 

Cule¾ in Angola. Other than the available information from the literature, 

are included all the unpublished data gathered by the writers and 

their team during several years of entomological work throughout the 

country and based upon the identification of more than 8,400 Culex specimens. 

The previous list of the Angolan Culex is now increased from 

34 presumed members to 47, with the addiction of the following species, 

subspecies and forms here recorded for the first time in Angola: C. (M.) 

salisburiensis salisburiensis, C. (E.) lingianus, C. (E.) wigglesworthi, 

C. (C.) aurantapex }in}aensis, C. (C.) chorleyi, C. (C.) ethiopicus, C. (C.) 

invidiosus type form, C. (C.) per/idiosus, C. (C.) ?er/uscus, C. (C.) 

pruina pruina, C. (C.) pruina eschirasi, C. (C.) toroensis, and C. (C.) 

weschei ? gediensis. Our knowledge on the geographical distribution of 

the species of Culex in Angola is also considerably increased with, on 

the whole, 447 new locality records. 

Keys to females and to fourth instar larvae of the Angolan representatives 

of the genus are also presented, which are intended to account 

for the variations found in our series. 

Under each Culex taxon, its previously known distribution, the material 

examined by the writers and the Angolan localities from which the 

mosquito is now known are indicated, and some notes on taxonomy and 

bioecology are added. Distribution maps for each taxon are also presented 

and considered from a zoogeographical viewpoint. 

Lastly, the medical importance of Angolan Culex, especially as 

vectors of arthropod-borne animal viruses, is briefly revised. 


RSUM 

Dans ce travail, les auteurs font une rvision de nos connaissances 

actuelles concernant le genre Culex h l'Angola. Outre l'information 

bibliographique disponible, sont ici incluses toutes les donnes encore 

indites obtenues par les auteurs et leur quipe pendant quelques 

annes de travail entomologique au travers du pays et bases sur 

l'identification de plus de 8400 specimens de Culex. La liste des 34 

Culex angolais jusqu'ici connus est maintenant passe / 47 lbments 

avec l'addition des espces, sousespces et formes ici repres pour la 

premiere lois h l'Angola: C. (M.) salisburiensis salisburiensis, C. (E.) 

kingianus, C. (E.) wigglesworthi, C. (C.) aurantapex }injaensis, C. (C.) 

chorleyi, C. (C.) ethiopicus, C. (C.) invidiosus type form, C. (C.) per- 

[idiosus, C. (C.) perfuscus, C. (C.) pruina pruina, C. (C.) pruina 

eschirasi, C. (C.) toroensis and C. (C.) weschei ? gediensis. Notre connaissance 

de la distribution gographique des Culex angolais est aussi 

de beaucoup amliore avec 447 nouvelles localit,s pour l'ensemble 

de toutes les espces. 

Des clefs des femelles et des larves au quatrime stade de tous 

les Culex de l'Angola sout aussi present,es, en essayant d'y inclure 

les variations trouves dans nos sries. 

Pour chaque taxon, on indique sa distribution jusqu'ici connue 

/ l'Angola, le matriel examin6 par les auteurs et les localitfis oh il 

est maintenant connu, en ajoutant aussi quelques notes taxonomiques 

et biocologiques. Sont aussi pr6sentes des cartes de distribution pour 

chaque taxon, envisages d'un point de vue zoog6ographique. 

3, la fin, on essaye un bref aperqu de l'importance m6dicale des 

Culex angolais, particulirement comme vecteurs de virus de l'homme 

et des animaux transmis par des arthropodes. 

Estud., Ens. e Doc.- 13. 131

RESUMO 

Neste trabalho, os autores fazem uma revisfio dos nossos conhecimentos 

actuais sobre o gnero Culex em Angola. Juntamente corn 

a informa;io disponivel na literatura, silo incluidos todos os dados 

ainda inditos obtidos pelos autores e sua equipa durante vfirios anos 

de trabalho entomo16gico atravs do Pais. As 152 localidades onde foi 

colhido o material aqui estudado estendem-se desde o Norte de Cabinda 

fronteira corn a Namibia e o Caprivi, no Sul, e desde o nivel 

do mar at cerca de 2000 metros de altitude, incluindo a faixa litoral, 

a montanha marginal, a escarpa ao Iongo desta e o planalto. Nio foi 

possivel, no entanto, prospectar certas fireas significativas do Norte 

e Leste de Angola, particularmente face fi fronteira corn a Zfimbia. 

Corn base na identificaio de mais de 8400 espcimes colhidos 

pertencentes a este gnero, a lista dos 34 Culex angolanos previamente 

conhecidos aumentada para 47 elementos, corn a adiio das seguintes 

espcies, subespcies e formas aqui assinaladas pela primeira vez em 

Angola: C. (M.) salisburienses salisburiensis, C. (E.) kingJanus, C. (E.) 

wigglesworthi, C. (C.) aurantapex jinjaensis, C. (C.) chorleyi, C. (C.) 

ethiopicus, C. (C.) invidiosus forma tipo, C. (C.) perfidiosus, C. (C.) 

perfuscus, C. (C.) pruina pruina, C. (C.) pruina eschirasi, C. (C.) toroensis 

and C. (C.) weschei ? gediensis. 0 nosso conhecimento da distribuiio 

geogrfifica das espcies de Culex em Angola tambm consideravelmente 

aumentado com 447 novas localidades para o conjunto 

de todas as esp6cies. 

Sio tambm apresentadas chaves de identifica;io das fSmeas e 

larvas em quarto estfidio de todos os representantes angolanos do 


RIBEIR0, H. & RAMOS, H. Cunha -- <strong>Research</strong> on the mosquitoes of Angola X 

g6nero Culex, tendo-se procurado dar conta das variables presentes 

nas nossas s6ries. 

Em Angola, estio representados os cinco subg6neros de Culex 

caracteristicos da Africa ao sul do Sara: subg6nero Lutzia Theobald 

(com 1 esp6cie, a finica esp6cie africana de Lutzia), subg6nero Maillotia 

Theo. (com 1 espficie, a mais generalizada das 4 espficies do subg6nero 

que ocorrem no continente africano), subg6nero Eumelanomyia 

Theo. (com as 8 esp6cies africanas mais comuns), subg6nero Culiciomyia 

Theo. (3 esp6cies, tamb6m das mais largamente distribuidas em 

Africa) e subgfinero Culex L. (representado por 31 das 57 esp6cies 

conhecidas da Africa ao sul do Sara). 

Para cada taxon indicam-se a sua distribuifio previamente conhecida 

em Angola, o material examinado pelos autores e as localidades 

donde 6 actualmente conhecido, juntando-se ainda algumas notas taxon6micas 

e bioeco16gicas. S5o tamb6m apresentados mapas de distribui5o 

para cada taxon, considerados em relaio com um esboco da 

zoogeografia de Angola. A sinonimia e dados relativos ao material-tipo 

de cada taxon s5o apenas referidos quando e na reedida em que o 

imponha a anfilise sistemfitica, podendo ser consultada para esse efeito, 

a16m da bibliografia particular eventualmente citada, a filtima ediio 

(1977) do excelente Catfilogo Mundial de K. Knight & A. Stone. 

Nas notas taxon6micas silo tratados alguns dos problemas sistemtticos 

mais evidentes postos pelo estudo do material de Angola. 

Assim, admite-se que as formas angolanas de Culex (E.) horridus, C. 

(E.) inconspicuosus, C. (E.) rubinotus, C. (C.) chorleyi (da area de 

Cariango) e C. (C.) toroensis sejam na realidade distintas, ao nivel 

subespecifico, das respectivas forrnas-tipo. De modo semelhante, o C. 

(C.) simpsoni apresenta em Angola duas formas (designadas A e B no 

texto) que devem ser tratadas como duas subespficies diferentes, enquanto 

se julga que o C. (C.) ingrami angolano 6 na realidade uma 

nova esp6cie, ainda no descrita, muito pr6xima daquele. Um interesse 

particular oferece tamb6m o estudo sistemfitico das duas formas de 

C. (C.) pipiens presentes em Angola, as quais, de acordo corn os dados 

de vfiria ordem disponiveis, deveriam ser tratadas como espficies distintas. 

Por outro lado, o estudo sistemb, tico do C. (C.) annutioris, 

C. (C.) invidiosus e C. (C.) univittatus em Angola leva os autores 

a nio reconhecer subespficies nestes taxa mas apenas varia5o de 

carficter polim6rfico. 

[34 Estud., Ens. e Doc. 134

RIBgIRO, H. & RAMOS, H. Cunha -- <strong>Research</strong> on the mosquitoes of ,4ngola X 

Por filtimo, rev-se sucintamente a importfincia m6dica dos Culex 

angolanos, particularmente como vectores de arbovirus patog6nicos 

para o hornera e animais. Entre os C. (C.) p. quinquefasciatus, C. (C.) 

thalassius, C. (C.) poicilipes, C. (E.) rubinotus, C. (C.) theileri, C. (C.) 

aurantapex }in}aensis, C. (C.) simpsoni (forma A) e C. (C.) bitaeniorhynchus 

contam-se muito provavelmente os principals vectores dos 

arbovirus (entre vfirios outros) Germiston, Sindbis, Wesselsbron e West 

Nile, os quais se sabe circularera entre as populaq&s humanas de 


Eslud., Ens. c Doc.- 134 ]½,5

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77--MOUCHET, J., GARIOU, J. & HAMON, J.-->. Encyc. Ent. (A), 37: 1-383, illus.. 1959. 

Estud., Errs. e Doc.- 134 ]41

RIBgIR0, H'. & RAMOS, H. Cunha -- <strong>Research</strong> ot the mosquitoes of Atgola -- X 

80--SOMEREN, E. C. C. Van -- >. Proc. R. Ent. Soc. Lond. (B), 215 (1-2): 3-12, 1956. 

81--OVAZZA, M., HAMON, J. & NERI, P. -- . An. Inst. Hig. Med. Trop. In press. 

new 

93 -- PEREIRA, M. C.--. 

An. Inst. Med. Trop. (Lisbon), 3: 365-372, 1946. 

94--WOLFS, J.--. Rev. Zool. Bot. Afr., 4'1 (1): 87-90, 1948. 

95--MATTINGLY, P. F.-- The Biology of Masquito-Borne Disease. The 

Science of Biology. Series: 1. American Elsevier Publishing Company, 

Inc., New York, 1969. 

96 --ANONYMOUS-- Working Catalogue of Arthropod-borne viruses. The 

American Committee on Arthropod-borne viruses. U.S. A., 1977. 

97MCINTOSH, B. M. -- . Dept. Agr. Techn. Services, Entomology Memoir n ø 43, Pretoria, 

1975. 


RIBIIRO, H. & RAMOS, H. Cunha -- <strong>Research</strong> on the mosquitoes o1' Angola -- X 

98 -- KOKERNOT, R. H., CASACA, V. M. R., WEINBREN, M.P. & MCIN- 

TOSH, B. M.--rrSurvey for antibodies against Arthropod-borne viruses 

in the sera of indigenous residents of Angola>>. Trans. R. Soc. Trop. 

Med. Hyg., 50 (5): 563-570, 1965. 

99--KERR, J. A.-->. Ann. Trop. Med. Parasit., 

2t5: 119-127, 1932. 

100--JUPP, P. G. & MCINTOSH, B. M.--ttQuantitative experiments on the 

vector capability of Culex (Culex) univittatus Theobald with West 

Nile and Sindbis viruses>>. J. Med. Ent., 7 (3): 371-373, 1970. 

101--MCINTOSH, B. M., JUPP, P. G. & DOS SANTOS, I.-->. J. ent. 

Soc. Sth. Afr., 41 (1): 57-61, 1978. 

102--RODRIGUEZ, F. M. et a/. -- . Ind. J. Med. Res., 1515 (5): 719-725, 1977. 

103--HAMON, J. et al.--

ILLUSTRATIONS

RIIIIRO, H. & RAMOS, H. Cunha -- <strong>Research</strong> on the mosquitoes o1' ,4ngola X 

4t 0 0 

SOUTH WEST 

12e l&e $e 

o 

AFRICA 

2 

22 e 2& ß 

MAP 1--Main collecting localities. Localities marked with squares were not sur- 

veyed by the writers. 


RIBtoRO,' H.' &'RMOS, 'H. Cunha2 " "ResearCh on"tti md$qubes ':o[)Whkota ':: 

ß 

Conic 

SOUTH t$T 

ARiD 

MAP 2--Main biotic zones in Africa 

+. +. +- Mountain biomes. 

--Limit of e West Mfican subregion 

lf18 tstud., Ens. e Doc. ' 134 

// 

.. _ 

,

RIBgIRO, H. & RAMOS, H. Cunha -- <strong>Research</strong> on the mosquitoes o! Angola 

-IO 

-12 

West African Subregion 

Guinean Forest Zone 

Escarpment Zone 

Southern Congo Savanna Zone 

! 

East and South African Subregion 

Humid Montane Zone 

Rhodesian Highland Zone 

:_.X Southwest Arid Zone 

..' Northern Strip of the 

Southwest Arid Zone 

MAP 3 -- Tentative zoogeographical division of Angola. 


X

RIBEIRO, H. & RAMOS, H. Cunha -- <strong>Research</strong> on the mosquitoes o! Angola X 

ONGO 

12 ß 

.% 

SOUTH WEST AFRICA 

20 © 

LEGEND 

(LuLzlo) t,gr*.9e, ß 

MAP 4--Known distribution in Angola of Culex (Lutzia) tigripes. 

150 Eslud., Ens. e Doc. - 134

RIBEIR0, H. & RAMOS, H. Cunha -- <strong>Research</strong> on the mosquitoes of Angola X 

.SOUTH i 

MAP 5--Known distribution in Angola of species of subgenus Maillotia and of 

the eumelanomyia and rubinotus-rima groups of subgenus Eumelanomyia. 



ß 

ß 

ß ß 

ß 

ß 

cjOUTH WEST A F R I CA 

22 ø 24 ß 

MAP 6- Known distribution in Angola of species of the subgenera Eumelanomyia 

(mochthogenes group). 


RIBEIRO, H: & RAMOS, H., Cunha -- ResEarch on.,the mosquiJoes o[ lngala X 

%OUT H 

16ø / 

LEGEND 

c_ LL.uS i nor 

œ1ntru$ 

nebgtoss 

w œ b T '. .A [_ 

16o 18 -0 2 ø 22 ø . . 

MAP 7 .--Known distribution in,Angola of species of the su. bge. n, us Culiciomyia. 


6 ø 

_

RIBSIR0, H. & RAMOS, H. Cunha -- <strong>Research</strong> on the mosquitoes of Angola -- X 

12 e 

CONGO 

Io 

o o o 

o 

oOO 

o 

o 

o o 

LEqEND 

aurontope 


A I & I 

14, e 15 e 1 e 20 e 22 ß 2& e 

MAP 8- Known distribution in Angola of species of the subgenus Culex, group A, 

bitaenniorhynchus subgroup. 1- C. annulioris and C. aurantapex jinjaensis. 


RIBEIRO, H. & RAMOS, H. Cunha -- <strong>Research</strong> on the mosquitoes of ,4ngola X 

o 

ß ß 

ß 

ß 

12 ø l& e 

o 

LEGEND 

Z 

X 

bttoniorhynchu s 

ethiopicus 

I p_o,cJl;pe 

oo 

o 

WEST AFRICA 

i 

1,' 

i 

15' 20' ;212 ß 2' 

MAP 9--Known distribution in Angola of species of the subgenus Culex, group A, 

bitaeniorhynchus subgroup. 2--C. bitaeniorhynchus, C. ethiopicus, and C. poicilipes. 

Estud., E/rs. e Doc.- 134 155 

o

RIBEIR0 , H. & RA. MOS , H. Cunha -- Res.e.:arch on the mosquitoes o[ Angola X 

12e 1/* ø 16 ø 18 ø 20 ø 2:2 ø 

ONGO 

o 

6 6 

c A 

A 


duttoni 

thalosslus 

LEGEND, 

/ : !,ritocn[orh),nc hus 

E ti 

12 ø l& ø 18 ø 18 ø 20 ø ;2 ø 2/, ø 

MAP 10- Known distribution in Angola of species of the subgenus Culex, group A. 

3- C. duttoni, C. thalassius, C. tritaeniorhynchus and C. watti. 

1.56 Estud., Ens. e Doc.- 134

RIBEIrO, H. & RAMOS, H. Cunha -- <strong>Research</strong> on the mosquitoes o[ Angola 

120 14 o 16 o 11o 20 ø . 22o 2&o 

0 ' ' ' ' L 

œ,END: 

orqznteopunctQU 

form A 

6 ø 6e rm B 

o ø 

, 16 

MAP 11 --Known distribution in Angola of species of the subgenus Culex, group A. 

4 C. argenteopunctatus kingi and C. simpsoni. 


RIBIiIIRO, H. & RAMOS, H. Cunha -- <strong>Research</strong> on the mosquitoes o[ 4ngola -- X 

A 


l& ø 16 ø 18 ø 2 ß 2 ß 2t, o 

1 6 ø 

MAP 12 -- Known distribution in Angola of species of the subgenus Culex, group A. 

5- C. theileri and C. univittatus. 

158 Estud., Ens. e Doc. - 134 

8 ø 

o ø


CONGO 

A V ?? ?v A 

Y 

18 e 20 ø 22 ß 2;, 

LE.&END 

pipinns ptplens 

/ mote records 

) fcmotc and/n- ',,'vt records 

? 

?pin qulnquefosciot 

H WEST AFRICA 

i d 

12 o 14 ß '16 ß 1 $ o 2 0 ø 22 ø 24 ß 

mate records 

ttmoe and/or rtcords arvo 

MAP 13--Known distribution in Angolan of species of the sugbenus Culex, 

group B. I- C. pipiens piplens and C. p. quinque/ascia/us. 


RIBEIRO, H. & RAMOS, H. Cunha -- <strong>Research</strong> on the mosquiioes o/ Angola -- X 

ONGO ' 

SOUTH WEST 

12 ø 11' ø 716o 

16 ø It ø 20 ø 22 ø .21' ø 

AFRICA 

LEGEND 

chortc, 

torocns;s 

trifdatus 

zombaens s 

20 ø 22 o 21, o 

MAP 14--Known distribution in Angola of species of the subgenus Culex, 

group B. 2- C. chorleyi, C. toroensis, C. triiilatus and and C. zombaensis. 


RIBIIRO, H.'& RAMOS, H. Cunha -- <strong>Research</strong> on the mosquitoes o!'Angofa X 

12 ø 14 ø 

SOUTH WEST 

? 

12 ø 14 ø 16 ø 18 ø 

? 

A F R C A 

LEGEND. 

ontcnnotus 

½cccns 

0 ø 2 ø 24 ø 


group B. 3- C. antennatus and C. decens. 


RIBgIRO, H, & R.MOS, H. Cunha -- <strong>Research</strong> on the mosquitoes o[ Angola X 

SbUTH WEST 

12 e l& e 1 e 11' 

AFRICA 

20 22 e 


group B. 4- C. invidiosus, C. ornatothoracis, C. per[idiosus, C. per[uscus, C. teh,- 

silla and C. tri/oliatus. 

162 Estud., E,ts, e Doc.- 134

RIBgIRO, H. & RAMOS, H. Cunha -- <strong>Research</strong> on the mosquitoes of /lngola -- X 

12 ø 1/* ø 16* 19 ø 20 ' 22ø 2' 

CONGO LEGEND 

Z 9rahaml 

, , 

SOUTH WEST 

l& ø 16 ø 18 ø 

AF RI C,, 

prun schros 

20 ø 22 ø 24* 

MAP 17- Known distribution in Angola of species of the subgenus Culex, group B. 

5--C. grahami, C. guiarti, C. ingrami vat., C. pruina and C. weschei? gediensis. 


60

RIBEIRO, H. & RAMOS, H. Cunha -- <strong>Research</strong> on the mosquitoes o! Angola -- X 

, , , . ,,... . ..... ......., ,11. 

ß . - . ' . ?,..- , . . .. 

.'..' , .',d;.; '. ''...' . ' 

. g -', . : 

. 

': ...... *'". .. .- .> 

.;,,.. .,,: i: ., .. 

. .. 

..... 

,. . 

. 

. , 

ß . 

ß . ... ,. , ' . ... 

j.:..- ,...,': -, ' . 

.. : 

..q 

:... ., " ' ß . ' :.' ,x 

FIG. 1- Pipetfing laae of C (L.) tigripes, C. (Culic.) cinerellus and C. 

ingrami from a niche in the margins of e Lulondo ver, at Tando Zinze (Cabinda 

District). Note e cleared lowland Congo forest. 

FIG. 2--Collecting larvae of C. (E.) inconspicuosus and C. (C.) per[idiosus (as 

well as of the associated An. [unestus) from rock-pools in the margins of the 

Luachimo River, at Dundo. 


RIBgIRO, H. & RAMOS, H. Cunha -- <strong>Research</strong> on the mosquitoes o! 4ngola 

. . -' . 

FIG. 3Muembege River, at la Tando. ae of C (L.) tigripes, C (E.) 

kingius, C (E.) wigglesworthi and C (Culic.) cinerdlus (as well as of e associat 

An. demeilloni d Uranotaenia mhonaensis) were caught in e niches 

at margins and mong e veble deb. 

Estud., En. e Doc.-,-134 165 

. 

X


FIG. 4- Pipetting larvae of C. (Culic.) nebulosus from bamboo stumps, at Dala 

166 

Tando. 

Estud., Ens. e Doc. -- 134


FIG. 5- Collecting larvae of C. (C.) annulloris, C. (C.) slmpsonl form A and 

C. (C.) univittatus (as well as of the associated An. Iongipalpis) from grassy earth 

drains, at Bocoio. 


RIBEIR0, H. & RAMOS, H. Cunha -- <strong>Research</strong> on the mosquitoes o[ lngola 

FIG. 6--Pipetting larval C. (L.) tigripes, C. (C.) antennatus, C. (C.) poict7ipes, 

C. (C.) telesilla and C. (C.) univittatus (as well as 4n. gambiae, 4n. argenteolobatus, 

,4n. pharoensis and ,4n. squamosus) in the overflown grassy margins of the receding 

Cubang River, at Calai (South-West Africa in the background). 

168 œstud., Ens. e Doc. 134 

X

RIBEIR0, H. & RAMOS, H. Cunha -- <strong>Research</strong> on the mosquitoes o] Angola M X 

- .' ; . ... 

., .. .. 

. "'* ' ...' 

.. . ,' ..:.s - 

s'j?. 

FIG. 7--Rocky niche in the margins of the Husolo River, at Ebanga, breeding 

place of C. (C.) chorleyi and C. (C.) univittatus, as well as of the associated An. 

marshalii. 


, ß


.......... ...; ..: ? 

. 

.: 

ß '%4, .'. 

ß '-? ":-. 

iv:" *: ..... : 

. ' 

FIG. 8 Rock-pools covered with fallen leaves along the margins of the Caringa 

River, at Dala Tando, typical larval biotopes of C. (Culic.) cinerellus and C. (C.) 

decens. Uranotaenia mashonaensis was found breeding among the rocks. 

170 Estud., Ens. e Doc. 134 

. .


FIG. 9- Larvae of C. (C.) duttoni and C. (C.) pruina pruina were collected from 

holes in these rocks of the Capecha River, at Dala Tando. 


Rmv. mo, H. & RAM08, H. Cunha -- <strong>Research</strong> on the mosquitoes o] lngola . 

FIG. 10--Lagoa dos Paralelos pond, in the Momedes desert, north of Virei, 

typical breeding site of C. (C.) ethiopicus, C. (C.) simpsoni form A and the asso- 

ciated An. squamosus. Paralelos Mount is seen at left, while Mount Canahuia, at 

ß far center, dominates the landscape. 

_ _ 

.. . , ,% .. ,,. . ' 

%.-,.,. ".. -¾. . . . ß .:, .. 

FIG. 1 !--Small pond near Cablnda (town), in the sothern coastal area of the 

Cabinda District, brding ar of C. (C.) guiarti and of the assiated Ficalbia 

uni/ormis. 

172 Estud., Ens. e Doc. w 134

RIBgIRO, I-[. RAMOS, H. Cunha -- <strong>Research</strong> on the mosquitoes o! /tngola -- X 

ß . . . ß . .t,,a ?, ß 

ß . , .. ... ... , . .....'.'. . . ...... ...... ,.,,. .. 

%". ' ' - ._ .......... "' "",4'":? . ....... . ....,.... ..'.,:,,,i ..,'' ¾'* . ;';, -,. '';.' '";..%' . . .,. ..:. 

'" ..:., . .- ....,,. ß :..' ......:: ..... 

.. ,,., ...... ,.... .. .... 

FIG. 12--Pipetting larvae of C. fL.) tigripes, C. (C.) decens, C. (C.) invidiosus 

type form, C. (C.) perIidiosus (and of the associated Uranotaenia [usca) from 

rockholes in the Luachimo River margins, at Dundo. Note the gallery forest. 


RIBEIRO, H. & RAMOS, H. Cunha -- <strong>Research</strong> on the mosquitoes o! .4ngola 

FIG. 13- Sunlit ground poll of clear water, at Calai, typical breeding place of 

C. (C.) poicilipes and C. (C.) theilerœ .4n. squamosus was found as an associate 

breeder. 

174 Estud., Ens. e Doc. 134 

X

RIBEIRO, H. & RAMOS, H. Cunha -- <strong>Research</strong> on the mosquitoes o! Angola m X 

. , . .... . . . ß 

FIG. 14 Grassy niche in the tourgin of c Cubul d Gundu rivcr, t Cubul, 

wcll of iat . m6[ nd n. corsl . sir. 

Esmd., Ens. e Doc.- 134

COMPOSTO E IMPRESSO NA: 

PRINTIPO--IndGstrias Gr&flca$, Ld. 

R. MOUSINHO DE ALBUOUEROUE, 6-C 

TELEFS. 97 05 33 - 97 92 43 -- DAMAIA

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