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
<strong>Research</strong> on the Mosquitoes of
Angola (Diptera, Culicidae)
X THE GENUS CULEX L., 1758. CHECK-LIST
WITH NEW RECORDS, KEYS TO FEMALES AND
LARVAE, DISTRIBUTION, AND TAXONOMIC AND
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
<strong>Research</strong> on the Mosquitoes of
Angola (Diptera,, Culicida 0
X--THE GENUS CULEX L., 1758. CHECK-LIST
WITH NEW RECORDS, KEYS TO FEMALES AND
LARVAE, DISTRIBUTION, AND TAXONOMIC AND
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
Estud., Ens. e Doc.- 134 19

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
Estud., Ens. e Doc.- 134 23

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.
Estud., Ens. e Doc.- 134 27

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.
Estud., Ens. e Doc.- 134 31

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.
Estud., Ens. e Doc.- 134 35

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

RIBEIRO, H. & RAMOS, H. Cunha -- <strong>Research</strong> on the mosquitoes of Angola -- X
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
Estud., Ens. e Doc.- 134 37

RIBgIRO, H. & RAMOS, H. Cunha -- <strong>Research</strong> on the mosquitoes of Angola -- X
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,
Estud., Ens. e Doc. 134 41
40
49
41
42
43
47

RIBEIRO, H. & RAMOS, H. Cunha -- <strong>Research</strong> on the mosquitoes of Angola -- X
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.
Estud., Ens. e Doc.- 134 43

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
Estud., Ens. e Doc.- 134 45

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
48 Estud., Ens. e Doc. 134

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.
Estud., Ens. e Doc.- 134 49

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
50 Estud., Ens. e Doc.- 134

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

RIBEIRO, H. & RAMOS, H. Cunha -- <strong>Research</strong> on the mosquitoes of Angola X
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
52 Estud., Ens. e Doc.- 134

RIBEIRO, H. & RAMOS, H. Cunha -- <strong>Research</strong> on the mosquitoes of Angola -- X
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).
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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.
54 Estud., Ens. e Doc.- 134

RIBEIRO, H. & RAMOS, H. Cunha -- <strong>Research</strong> on the mosquitoes of Angola X
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
Estud.. Ens. e Doc.- 134 55

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

RIBEIRO, H. & RAMOS, H. Cunha -- <strong>Research</strong> on the mosquitoes of Angola X
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).
60 Estud., Ens. e Doc. 134

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
New record for Angola.
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).
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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-
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RIBgIRO, H. & RAMOS, H. Cunha -- <strong>Research</strong> on the mosquitoes of Angola X
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)
Estud., Ens. e Doc.- 134 63

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
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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
Estud., Ens. e Doc.- 134 65

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-
Estud., Ens. e Doc.- 134 67

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

RIBEIRO, H. & RAMOS, H. Cunha -- <strong>Research</strong> on the mosquitoes of Angola -- 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
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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
72 Estud., Ens. e Doc.- 134

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.
Estud., Ens. e Doc.- 134 73

RIBEIRO, H. & RAMOS, H. Cunha -- <strong>Research</strong> on the mosquitoes o[ Angola -- X
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
New record for Angola.
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;
Estud., Ens. e Doc.- 134 75

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

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
78 Estud., Ens. e Doc.- 134

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

RIBgIRO, H. & RAMOS, H. Cunha -- <strong>Research</strong> on the mosquitoes of Angola -- X
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
82 Estud., Ens. e Doc.- 134

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

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

RIBgIRO, H. & RAMOS, H. Cunha -- <strong>Research</strong> on the mosquitoes of Angola X
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.
86 Estud., Ens. e Doc.- 134

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
New record for Angola.
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
Angola.
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
New record for Angola.
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.
88 Estud., Ens. e Doc.- 134

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
Estud., Ens. e Doc.- 134 89

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

RIBgIRO, H. & RAMOS, H. Cunha -- <strong>Research</strong> on the mosquitoes of Angola -- X
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 &
Estud., Ens. e Doc.- 134 93

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
94 Estud., Ens. e Doc.- 134

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

RIBEIRO, H. & RAMOS, H. Cunha -- <strong>Research</strong> on the mosquitoes of Angola X
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
Estud., Ens. e Doc.---134 97

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
98 Estud., Ens. e Doc.- 134

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

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

RIBEIRO, H. & RAMOS, H. Cunha -- <strong>Research</strong> on the mosquitoes of Angola X
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
New record for Angola.
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
Estud., Ens. e Doc.---134 101

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.
Estud., Ens. e Doc.- 134 105

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

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

RIBgIRO, H. & RAMOS, H. Cunha -- <strong>Research</strong> on the mosquitoes of Angola X
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
112 Estud., Ens. e Doc.- 134

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

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

RIBEIRO, H. & RAMOS, H. Cunha -- <strong>Research</strong> on the mosquitoes of Angola X
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-
Estud., Ens. e Doc.- 134 115

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;
116 Estud., Ens. e Doc. 134

RIBgIRO, H. & RAMOS, H. Cunha -- <strong>Research</strong> on the mosquitoes of Angola X
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
Estud., Ens. e Doc.- 134 117

RIBEIRO, H. & RAMOS, H. Cunha -- <strong>Research</strong> on the mosquitoes of Angola -- X
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.
118 Estud., Ens. e Doc.- 134

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

RIBEIRO, H. & RAMOS, H. Cunha -- <strong>Research</strong> on the mosquitoes of Angola -- X
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-
Estud.. Ens. e Doc.- 134 123

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
Estud., Ens. e Doc.- 134 125

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.
Estud., Ens. e Doc.- 134 127

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.
Estud., Ens. e Doc.- 134 129

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
Estud., Ens. e Doc.- 134 133

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
Angola.
Eslud., Ens. c Doc.- 134 ]½,5

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(Diptera: Culicidae)>>. Proc. Ent. Soc. Wash., 63: 29-52, 1961.
19- STONE, A.- ((A Synoptic Catalog of
(Diptera: Culicidae)>>. Proc. Ent.
20- STONE, A.- ((A Synoptic Catalog of
(Diptera: Culicidae)>>. Proc. Ent.
the Mosquitoes of the World, suppl. I1
Soc. Wash., Iõ: 117-140, 1963.
the Mosquitoes of the World, suppl. III
Soc. Wash., 19: 197-224, 1967.
21 --STONE, A.- ((A Synoptic Catalog of the Mosquitoes of the World, suppl. IV
(Diptera: Culicidae)>>, Proc. Ent. Soc. Wash., 72: 137-171, 1970.
22--STONE, A. -- ((Corrections to mosquito catalog supplements III and IV>>.
Proc. Ent. Soc. Wash., 73: 180, 1971.
23--KNIGHT, K. L. & STONE, A.--((A Catalog of the Mosquitoes of the
World (Diptera: Culicidae)>>. The Thomas Say Foundation. Ent. Soc.
Am., 6:611 pp., 1977.
24- HOPKINS, G. H. E.- ((Mosquitoes of the Ethiopian Region. I- Larval bio-
nomics of mosquitoes and taxonomy of Culicine larvae>>. London, 1952.
25- SOMEREN, E. C. C. Van- ((Ethiopian Culicidae: a description of the early
stages of Culex (CuIex) terzii Edwards>>. Proc. R. Ent. Soc. Lond.
(B), 27 (5-6): 84-88, 1958.
26 SOMEREN, E. C. C. Van- ((The female and early stages of Culex (Culex)
nakuriensis Mattingly, with a description of a new subspecies of Culex
(CuIex) shoae Hamon & Ovazza>>. Proc. R. Ent. Soc. Lond. (B), 36 (1-2):
11-16, 1967.
27- MATTINGLY,
dix II.
F. P. In MATTINGLY, F. P. & BROWN, E. S., 1965. Appen-
28- De MEILLON, B., PARENT, M. & BLACK O'C.- ((Descriptions of new
larvae and pupae of Ethiopian Culicini>>. Bull. Ent. Res., 36 (1): 85-101,
1945.
138 Estud., Ens. e Doc.- 134

RnmI}O, H. & R.4M08, H. Cunha -- <strong>Research</strong> on the mosquitoes o[ Angola
29 -- SIRIVANAKARN, S.--. Con>.
Ann. Soc. ent. France, ]3:>: 85-144, 1963.
38- KNIGHT, K. L.- . Proc. Ent. Soc. Wash.,
39-
(5): 212-234, 1953.
MUSPRATT J. .
J. ent. Soc. Sth Air., 24 (1): 92-103, 1961.
40--HAMON, J. & RICKENBACH, A.- . Bull. Soc. Path. Ex., 48 (6): 848-859, 1955.
41--CARTER, H. F.. Bull Ent. Res., 2:
37-38, 1911.
42--MATTINGLY, P. F. & LIPS, M. -- . Rev. Zool. Bot. air., 47 (3-4): 311-343, 1953.
43--CORBET, P.S., WILLIAMS, M. C. & GILLETT, J. D.--

RIBIIRO, H. & RAMOS, H. Cunha -- <strong>Research</strong> on the mosquitoes o/ Angola X
46--EDWARDS, F. W. -- Diptera Faro. Culicidae, in P. Wytsman 'Genera !nsectorurn>>,
194 ..... Fas. Diptera, Bruxelles, 1932.
47- HAMON, J. & OVAZZA, M.- . Bull. Soc.
Path. Ex., 49 (i): 89-99, 1956.
48- MATTINGLY, P. F.- . Ann. trop.
Med. Parasit., 41: 239-252, 1947.
49--SOMEREN, E. C. C. Van, TEESDALE, C. & FURLONG, M.- . Bull. Ent. Res., 46: 463-493, 1955.
50--RIBEIRO, H. -- . Anais Inst. Med. Trop., :>3 (i-2):
157-161, 1966.
51 --LARIVIRE, M. & ABONNENC, E.- . Bull.
Inst. fran'. Afr. noire, 18 (4): 1191-1199, 1956.
52--SIRIVANAKARN, S.-

RIBIglRO, H. & RAMOS. H. Cunha -- <strong>Research</strong> on the mosquitoes of Angola -- X
63--CHAPIN, J. P.--. Bull. Amer.
Mus. Nat. H., 65: 1-203, 1932.
64--SUNDARARAMAN, S.-->. Am. J. Hyg., 150: 307-314, 1949.
65--BARR, A. R.--. Am. J. Trop. Med. Hyg., 6 (1): 153-165,
1957.
66--KNIGHT, K. L. & MALEK, A. A.-->.
Bull. Soc. Fouad 1 e Entom., Cairo, 35: 175-185, 1951.
67--KNIGHT, K. L. -- >. U. S. Naval Medical <strong>Research</strong> Unit n ø 3,
Cairo, Egypt: 1-3, 1951.
68 -- MATTlNGLY, P. F.--((The distribution of some African mosquitoes>>.
Proc. Linn. Soc. Lond., Sess. 165, 1952-53 (1): 49-61, 1954.
69--RIBEIRO, H. -- . -- Mosquito News, 33 (4): 568-572, 1973.
70--EDWARDS, F. W. -- . Bull. Ent. Res.,
4: 47-59, 1913-1914.
71--BELKIN, J. N.--. 2 vols., 608 and 412 pp., illus., University of California Press.
1962.
72--DOBROTWORSKY, N. V.--(>. Bull. Org. mond. Santd, 37:
251-255, 1967.
73- MUSPRATT, J.- . J. ent. Soc. Sth. Afr., '18 (2):
149-207, 1955.
74--THEOBALD, F. V.--. Ann. & Mag.
of Nat. Hist., 8 th series, 5: 373-378, 1910.
75--MATTINGLY, P. F. & BROWN, E. S.- . Bull Ent. Res., 46: 69-110, 1955.
76--HAMON, J., ABONNENC, E. & NOEL, E. -- . Ann. Parasit. hum. comp., 30 (3):
278-308, 1955.
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.
142 Estud., Ens. e Doc.- 134

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.
Estud., Ens. e Doc. 134 147

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.
Estud., Ens. e Doc. 134 149
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.
Estud., Ens. e Doc.- 134 151

RIBEIRO, H. & RAMOS, H. Cunha -- <strong>Research</strong> on the mosquitoes of Angola X
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).
152 Estud., Ens. e Doc.- 134

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.
Estud., Ens. e Doc. 134 153
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
SOUTH WEST AFRICA
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.
154 Estud., Ens. e Doc.- 134

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
SOUTH WEST AFRICA
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.
Estud., Ens. e Doc. 134 157

RIBIiIIRO, H. & RAMOS, H. Cunha -- <strong>Research</strong> on the mosquitoes o[ 4ngola -- X
A
SOUTH WEST AFRICA
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

RIBEIRO, H. & RAMOS, H. Cunha -- <strong>Research</strong> on the mosquitoes o[ Angola -- X
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.
Estud., Ens. e Doc.- 134 159

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

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
MAP 15--Known distribution in Angola of species of the subgenus Culex,
group B. 3- C. antennatus and C. decens.
Estud., Ens. e Doc.- 134 161

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
MAP 16--Known distribution in Angola of species of the subgenus Culex,
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.
Estud., Ens. e Doc.- 134 163
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.
164 Estud., Ens. e Doc.- 134

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

RIBEIRO, H. & RAMOS, H. Cunha -- <strong>Research</strong> on the mosquitoes o! Angola X
FIG. 4- Pipetting larvae of C. (Culic.) nebulosus from bamboo stumps, at Dala
166
Tando.
Estud., Ens. e Doc. -- 134

RIBEIRO, H. & RAMOS, H. Cunha -- <strong>Research</strong> on the mosquitoes o[ Angola X
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.
Estud., Ens. e Doc.- 134 167

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.
Estud., Ens. e Doc.- 134 169
, ß

RIBEIRO, H. & RAMOS, H. Cunha -- <strong>Research</strong> on the mosquitoes o[ Angola X
.......... ...; ..: ?
.
.:
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
. .

RIBEIRO, H. & RAMOS, H. Cunha -- <strong>Research</strong> on the mosquitoes o! Angola X
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.
Estud., Ens. e Doc.- 134 171

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.
Estud., Ens. e Doc.- 134 173

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