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.
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.
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.
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); Sá et al., 2020 (Atratus Group, taxonomy, morphology, descriptions, bionomics, distributions, keys); Sá 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 Sá 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 Sá et al., 2020) comatus Senevet & Abonnenc, 1939 commevynensis Bonne-Wepster & Bonne, 1920 comminutor Dyar, 1920 comptus Sá & Sallum, 2020 (in Sá 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 Sá et al.,2020) longistriatus Sá & Hutchings, 2022 (in Sá et al., 2022) longistylus Sá & Sallum, 2020 (in Sá 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 Sá et al., 2020) spiniformis Sá & Hutchings, 2022 (in Sá 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