Subgenus Melanoconion Theobald, 1903

Type species: 

Culex atratus Theobald, 1901.


Subfamily Culicinae, genus Culex. Subgenus Melanoconion includes 177 species. See Culex classification, Subgenus Melanoconion). Subgenus abbreviation – Mel.


ADULTS – Small or medium-sized mosquitoes; antenna as long as proboscis; decumbent scales of vertex varied, narrow, linear or falcate in center to entirely broad; maxillary palpus and proboscis entirely dark-scaled; palpus with 3 or 4 palpomeres in females, palpomere 4 vestigial or absent; palpus of males with 5 palpomeres, usually longer than proboscis, sometimes slightly shorter, palpomeres 4 and 5 usually strongly setose; scales on anterior half of scutum usually entirely dark, infrequently partially pale, producing a distinct pattern; acrostichal setae absent [except Cx. spissipes]; antepronotum with row of setae only, scales absent; scales present on lower posterior surface of mesokatepisternum, present or absent on upper corner and upper median area of mesanepimeron; legs usually dark-scaled, femora sometimes with pale knee spots and/or pale rings at joints of hindtarsomeres; wing scales varied in shape; abdominal terga usually with distinct basolateral pale spots, sometimes with complete basal pale bands, rarely entirely dark-scaled. MALE GENITALIA Ninth tergal lobes varied in size and form, narrowly to rather widely separated; gonocoxite varied in shape, with scales; subapical lobe subapical in position, divided, proximal and distal divisions strongly developed, widely separated, each with basal columnar stem of varied length, stem of proximal division distally divided or undivided, bearing 2 apical rod-like setae, stem of distal division shorter or as long as stem of proximal division, bearing 1 hooked rod, 2 blade-like setae and 3 or 4 appressed foliform setae and/or 1 or more variously modified setae; gonostylus at least half length of gonocoxite, varied in form; phallosome not columnar, aedeagus usually without upper tergal bridge connecting lateral plates; lateral plate varied in development, with prominent basal hook; paraproct with comb-like crown of 5–12 blunt or pointed blades; cercal setae present or absent. LARVAE – Head broader than long, usually with prominent ocular bulge; antenna as long or slightly shorter than head; seta 1-A usually large, rarely reduced, inserted beyond mid-length of antenna; seta 2-C usually absent, sometimes present; seta 5-C weakly to strongly developed, single or branched; seta 6-C long, always single; seta 3-P much shorter then setae 1,2-P; seta 7-I large, similar to seta 6-I, seta 7-II small, similar to seta 7-III–VI; comb usually comprised of numerous fringed scales in patch, sometimes relatively few rather large spine-like scales in patch or single row; setae 1,2-VIII usually with basal sclerotised plate; siphon tubular, distally tapering, length varied; pecten spines in close-set row, rarely widely spaced, usually with fine barbs of numerous denticles; seta 1-S in 3–8 posterolateral pairs, length varied, usually inserted beyond pecten, sometimes placed within pecten, 2 (sometimes 3 or 4) dorsolateral setae; seta 2-S usually well developed, hooked and usually with recurved tooth; saddle complete, without acus, usually without spines, inconspicuous spicules usually present; ventral brush (seta 4-X) with 5 or 6 pairs of setae on grid. PUPAE – Trumpet with distinct meatal slit extending from pinna, rarely absent; setae 1,5-III–VI usually with several branches; seta 9-VIII placed at or removed cephalad of caudolateral angle of segment; caudal lobe of tergum VIII not overlying lateral part of tergum IX; paddle broad, ovate, outer margin without spicules; seta 1-Pa and seta 2-Pa present. See genus Culex.

Phylogenetic relationships: 

Sirivanakarn (1983) suggested that the Melanoconion lineage may have given rise to Aedinus, Anoedioporpa, Belkinomyia, Carrollia, Micraedes, Microculex and Tinolestes based on morphological similarities shared with each of these subgenera. The results of a cladistic analysis based on implied weighting of morphological data by St John (2007) supported this hypothesis. The cladistic analyses of Harbach et al. (2012) supported the monophyly of the Melanoconion Group of subgenera, i.e. Aedinus, Anoedioporpa, Belkinomyia, Carrollia, Melanoconion, Micraedes, Microculex and Tinolestes, and indicated, as was suggested by Berlin (1969), Adames & Galindo (1973) and Sirivanakarn (1983), that it and genera Deinocerites and Galindomyia comprise the most derived mosquitoes of tribe Culicini. Torres-Gutierrez et al. (2018) investigated the phylogenetic relationships of 43 species of the subgenus based on fragments of two single-copy nuclear genes (CAD, HB) and a mitochondrial gene (COI). The results supported the monophyly of the Melanoconion and Spissipes Sections and a few groups and subgroups, but the monophyly of the Vomerifer and Crybda Groups of the latter section was doubtful. The authors emphasised the need for further investigation based on additional genes and species represented by accurately identified specimens, i.e. males and females with associated larval and pupal exuviae.

Bionomics and disease relations: 

The immature stages are found in a variety of ground-water habitats in heavy or partial shade or full sunlight, including forest pools, jungle swamps, ponds, marshes, lakes, margins of streams and canals containing plants or decaying plant material. Eggs are laid individually or in a batch. Adults are active at night, and females apparently feed on a variety of vertebrates, including humans.

A number of Melanoconion species are important actual or potential vectors of Venezuelan equine encephalitis virus and other arboviruses.


Melanoconion ranges from the southern part of the United States southward though most of Brazil, Paraguay and northern Argentina, and to Peru on the western side of the Andes Mountains.

Principal references: 

Sirivanakarn, 1983 (taxonomy, classification, morphology, bionomics, medical importance, distribution); Pecor et al., 1992 (illustrated catalogue, species listings, type data, synonyms, literature, distributions); Sallum & Forattini, 1996 (Spissipes Section, classification, keys, morphology, species descriptions, distributions, bionomics); Navarro & Weaver, 2004 (Vomerifer and Pedroi Groups, molecular phylogeny); Torres-Gutierrez & Sallum, 2015 (catalogue of South American species, classification, species listings, type data, synonyms, distributions, literature); et al., 2020 (Atratus Group, taxonomy, morphology, descriptions, bionomics, distributions, keys); et al., 2022 (Educator Group, taxonomy morphology, descriptions, distributions, keys); González et al., 2023 (Neotropical subgenera of Culex, comparative morphology, molecular associations).

abominator Dyar & Knab, 1909
abonnenci Clastrier, 1970
adamesi Sirivanakarn & Galindo, 1980
akritos Forattini & Sallum, 1995
albinensis Bonne-Wepster & Bonne, 1920
alcocki Bonne-Wepster & Bonne, 1920
aliciae Duret, 1953
alinkios Sallum & Hutchings, 2003
alogistus Dyar, 1918
amitis Komp, 1936
andricus Root, 1927
aneles Dyar & Ludlow, 1922
angularis Sá & Sallum, 2022 (in et al., 2022)
anips Dyar, 1916
anoplicitus Forattini & Sallum, 1989
apeteticus Howard, Dyar & Knab, 1913
aphyllus Talaga, 2020 (in Talaga et al., 2020)
arboricolus Galindo &  Mendez, 1961
atratus Theobald, 1901
aureonotatus Duret & Barreto, 1956
bahiensis Duret, 1969
bastagarius Dyar & Knab, 1906
batesi Rozeboom & Komp, 1948
bejaranoi Duret, 1953
bequaerti Dyar & Shannon, 1925
bibulus Dyar, 1920
bifoliolatus Duret & Barreto, 1956
brachiatus Hutchings & Sallum, 2008
carcinophilus Dyar & Knab, 1906
caribeanus Galindo & Blanton, 1954
caudatus Clastrier, 1970
caudelli (Dyar & Knab, 1906)
cedecei Stone &  Hair, 1968
changuinolae Galindo & Blanton, 1954
clarki Evans, 1924
columnaris Sá & Hutchings, 2020 (in et al., 2020)
comatus Senevet & Abonnenc, 1939
commevynensis Bonne-Wepster &  Bonne, 1920
comminutor Dyar, 1920
comptus Sá & Sallum, 2020 (in et al., 2020)
confundior Komp &  Rozeboom, 1951
conspirator Dyar &  Knab, 1906
contei Duret, 1968
coppenamensis Bonne-Wepster & Bonne, 1920
corentynensis Dyar, 1920
creole Anduze, 1949
cristovaoi Duret, 1968
crybda Dyar, 1924
delpontei Duret, 1969
diamphidius Peyton & Harbach, 1991
distinguendus Dyar, 1928
dolichophyllus Clastrier, 1970
dunni Dyar, 1918
dureti Casal & García, 1968
dyius Root, 1927
eastor Dyar, 1920
educator Dyar & Knab, 1906
egcymon Dyar, 1923
eknomios Forattini & Sallum 1992
elephas Komp, 1936
elevator Dyar & Knab, 1906
ensiformis Bonne-Wepster & Bonne, 1920
epanastasis Dyar, 1922
equinoxialis Floch & Abonnenc, 1945
ernanii Duret, 1968
ernsti Anduze, 1949
erraticus (Dyar & Knab, 1906)
evansae Root, 1927
exedrus Root, 1927
fairchildi Galindo & Blanton, 1954
faurani Duret, 1968
ferreri Duret, 1968
flabellifer Komp, 1936
foliafer Komp & Rozeboom, 1951
galindoi Komp & Rozeboom, 1951
galvaoi Duret, 1968
garcesi Duret, 1968
glyptosalpinx Harbach, Peyton & Harrison, 1984
gnomatos Sallum, Hutchings, Leila & Ferreira, 1997
guedesi da Silva Mattos & Xavier, 1991
herrerai Sutil Oramas, Pulido Florenzano & Amarista Meneses, 1987
hutchingsae Talaga, 2022 (in Talaga & Gendrin, 2022)
idottus Dyar, 1920
ikelos Forattini & Sallum, 1995
inadmirabilis Dyar, 1928
inextricabilis Talaga, 2020 (in Talaga et al., 2020)
inhibitator Dyar & Knab, 1906
innovator Evans, 1924
intonsus Galindo & Blanton, 1954
intrincatus Brèthes, 1916
invocator Pazos, 1908
iolambdis Dyar, 1918
isabelae Duret, 1968
jocasta Komp & Rozeboom, 1951
johnnyi  Duret, 1968
johnsoni Galindo & Mendez, 1961
jubifer Komp & Brown, 1935
kazanjii Talaga, 2020 (in Talaga et al., 2020)
kummi Komp & Rozeboom, 1951
lacertosus Komp & Rozeboom, 1951
limacifer Komp, 1936
longisetosus Sá & Sallum, 2020 (in et al.,2020)
longistriatus Sá & Hutchings, 2022 (in et al., 2022)
longistylus Sá & Sallum, 2020 (in et al., 2020)
lopesi Sirivanakarn & Jakob, 1979
loturus Dyar, 1925
lucackermanni Talaga 2022 (in Talaga & Gendrin, 2022)
lucifugus Komp, 1936
madininensis Senevet, 1936
martinezi Casal & García, 1968
maxinocca Dyar, 1920
mesodenticulatus Galindo & Mendez, 1961
milwardi Xavier & da Silva Mattos, 1972
misionensis Duret, 1953
mulrennani Basham, 1948
mutator Dyar & Knab, 1906
nicceriensis Bonne-Wepster & Bonne, 1920
ocossa Dyar & Knab, 1919
oedipus Root, 1927
olimpioi Xavier, da Silva & da Silva Mattos, 1970
orfilai Duret, 1953
palaciosi Duret, 1968
panocossa Dyar, 1923
paracrybda Komp, 1936
pavlovskyi Casal & García, 1967
peccator Dyar & Knab, 1909
pedroi Sirivanakarn & Belkin, 1980
penai Sirivanakarn, 1979
pereyrai Duret, 1967
phlabistus Dyar, 1920
phlogistus Dyar, 1920
phyllados Hutchings & Sallum, 2008
pifanoi Anduze, 1949
pilosus (Dyar & Knab, 1906)
plectoporpe Root, 1927
portesi Senevet & Abonnenc, 1941
productus Senevet & Abonnenc, 1939
psatharus Dyar, 1920
putumayensis Matheson, 1934
quadrifoliatus Komp, 1936
quasihibridus Galindo & Blanton, 1954
rabanicolus Floch & Abonnenc, 1946
rabelloi Forattini & Sallum, 1987
rachoui Duret, 1968
ribeirensis Forattini & Sallum, 1985
ronderosi de Linero, 1967
rooti Rozeboom, 1935
rorotaensis Floch & Abonnenc, 1946
sallumae Talaga, 2022 (in Talaga & Gendrin, 2022)
sacchettae Sirivanakarn & Jakob, 1982
saramaccensis Bonne-Wepster & Bonne, 1920
sardinerae Fox, 1953
serratimarge Root, 1927
silvai Duret, 1968
simulator Dyar & Knab, 1906
spathulatus Forattini & Sallum, 1987
spinifer Sá & Sallum, 2020 (in et al., 2020)
spiniformis Sá & Hutchings, 2022 (in et al., 2022)
spissipes (Theobald, 1903)
sursumptor Dyar, 1924
symbletos Sallum & Hutchings, 2003
taeniopus Dyar & Knab, 1907
tecmarsis Dyar, 1918
terebor Dyar, 1920
theobaldi (Lutz, 1904) (in Bourroul, 1904)
tournieri Senevet & Abonnenc, 1939
trifidus Dyar, 1921
trigeminatus Clastrier, 1970
trilobulatus Duret & Barreto, 1956
trisetosus Fauran, 1961
unicornis Root, 1928 (in Dyar, 1928)
vaxus Dyar, 1920
vexillifer Komp, 1936
vomerifer Komp, 1932
wepsterae Komp & Rozeboom, 1951
ybarmis Dyar, 1920
zeteki Dyar, 1918

Scratchpads developed and conceived by (alphabetical): Ed Baker, Katherine Bouton Alice Heaton Dimitris Koureas, Laurence Livermore, Dave Roberts, Simon Rycroft, Ben Scott, Vince Smith