Subgenus Nyssorhynchus Blanchard, 1902
Anopheles argyritarsis Robineau-Desvoidy, 1827.
Subfamily Anophelinae, genus Anopheles. Subgenus Nyssorhynchus includes 45 formally recognised extant species and one extinct (fossil) species. The extant species are divided between three Sections, the two largest of which are further subdivided into Series and Groups that are believed to represent phylogenetically related assemblages based principally on morphological similarity (Harbach, 2004). Subgenus abbreviation – Nys.
Subgenus Nyssorhynchus is most closely related to subgenus Kerteszia. The two subgenera are very similar in overall morphology, but the following features easily distinguish the adults, larvae ana pupae of Nyssorhynchus. The scutum of adults lacks dark longitudinal lines and has scales on the acrostichal and dorsocentral areas in addition to a few scales in patches on the lateral and/or anterior areas. The scutellum has numerous scattered white, yellow or silver scales along the posterior border. The wings differ in having crossveins sc-r and r1-rs closer together (distance about 0.30 or less the length of the sector dark spot on the costa and radius-one), no spot of dark scales where the humeral crossvein intersects the radius, the subcosta terminates at the proximal end or the middle of the subcostal pale spot, vein R4+5 is narrowly pale-scaled apically with a corresponding isolated apical pale fringe spot, vein CuA is mostly pale-scaled or equally pale- and dark-scaled without a basal dark spot and the anal vein has three pale-scaled areas setting off two dark areas between the basal and an apical pale spot (a greater amount of dark scaling occurs in species of the Myzorhynchella Section). The gonocoxite, claspette and aedeagus of the male genitalia exhibit a number of distinctions. The gonocoxite has a subapical seta, two subequal accessory setae are inserted before mid-length (about 0.4 from base), the internal seta is as long but much weaker than the accessory setae, and the parabasal seta is shorter than the internal seta (about 0.6 its length). The ventral lobes of the claspette are fused to form a single structure, the dorsal lobes each bear a single group of two or three specialised setae, and the aedeagus is broad and not noticeably tapered (at least in the basal 0.8). Larvae exhibit differential and diagnostic features, which include, but are not limited to, seta 1-A branched and inserted dorsomesally, seta 2-C inserted anterior to seta 3-C, seta 4-C inserted posteromesal to seta 3-C, hypostomal suture well developed (but not reaching the posterior tentorial pit), presence of a sclerotised tubercle at the bases of setae 1,2-P, short seta 4-M (not more than half as long as seta 6-M), strongly plumose setae 8-M,T and 6,7-I,II, palmate or fanlike seta 3-T, multiply branched seta 2-I–IV, presence of seta 14-III, seta 2-IV inserted mesad to seta 4-IV, well developed seta 1-S (with multiple branches) and eight pairs of seta 4-X. Salient features of Nyssorhynchus pupae include a trumpet with a long pinna (more than half as long as the trumpet) and a meatal cleft, seta 9-I more than half the length of seta 6-I, well developed setae 1-I–VII and 5-V–VII (nearly as long or longer than the segment), seta 14-III present, seta 2 on segments IV and V inserted mesad of seta 3, seta 3-VI inserted mesad to or in line with seta 1-VI, seta 10-VI absent, and seta 1-Pa slender, similar to seta 2-Pa or sinuous and apically hooked. See genus Anopheles.
With exception of the molecular study by Freitas et al. (2015), phylogenetic studies of anopheline mosquitoes based on both morphological and molecular data support for the monophyly of subgenus Nyssorhynchus (Krzywinski et al., 2001, 2001; Sallum et al., 2000, 2002; Collucci & Sallum, 2003; Harbach & Kitching, 2005, 2016; Foster et al., 2017). Except for the study of Foster et al., all of the studies that support the monophyly of the subgenus also support its sister relationship with subgenus Kerteszia. Nyssorhynchus was recovered as the sister to a clade comprised of subgenera Kerteszia and Stethomyia in the analyses of COI and COII mtDNA and 2.5S rDNA sequences conducted by Freitas et al., and was recovered in a sister relationship with Cellia + Anopheles in the study of Foster et al. based on analyses of mitochondrial protein coding genes. The analysis of morphological data by Harbach & Kitching (2005) indicated subgenus Cellia (Old World) is sister to Nyssorhynchus + Kerteszia, but this relationship was not recovered in their later study (Harbach & Kitching, 2015). It is noteworthy, however, that a molecular phylogeny of 18 species constructed from aligned protein sequences of 1,085 single-copy orthologs indicates that the Nyssorhynchus lineage diverged prior to the lineages that gave rise to subgenera Anopheles and Cellia (Neafsey et al., 2015). Phylogenetic relationships among species of subgenus Nyssorhynchus based on nuclear and mitochondrial gene sequences were elucidated by Bourke et al. (2010). Similarly, relationships among five of the six species belonging to the Myzorhynchella Section of subgenus Nyssorhynchus were investigated by Bourke et al. (2011) based on the ITS2 region of rDNA and the nuclear white gene. Foster et al. (2013) analysed combined data for the nuclear white and CAD genes and the COI barcode region of mtDNA obtained from representatives of the Albimanus, Argyritarsis and Myzorhynchella Sections and only found support of the monophyly of the last section. Similarly, their analyses did not support the monophyly of the Strodei Subgroup of the Oswaldoi Series or the monophyly of the Albitarsis Series of the Argyritarsis Section. The analyses of Foster et al. (2017) did not support the monophyly of the Myzorhynchella Section, which excluded An. parvus, and the Argyritarsis Series, which excluded An. atacamensis.
The immature stages of Nyssorhynchus species occur in a variety of sunlit or partially shaded habitats, including ponds, lakes, swamps, stream and river margins, canals, ditches, flood land, ground pools, borrow pits, spring water, wheel ruts and animal tracks. The immature stages of An. aquasalis mainly occur in brackish water habitats such as in mangrove swamps and coastal ground pools, and those of An. albimanus, in addition to ground pools and stream margins, have also been found in crab holes, tree holes and artificial containers. Females of most Nyssorhynchus species feed at dusk or during the night. They feed on a variety of large mammals and infrequently on birds. Some species readily attach humans.
Anopheles albimanus, An. aquasalis, An. argyritarsis, An. darlingi, An. nuneztovari and An. oswaldoi s.l. are vectors of malarial protozoa. Molecularly identified An. dunhami (as Nyssorynchus dunhami) was found recently to harbour natural infections of Plasmodium falciparum in the peri-Iquitos region of Amazonian Peru (Prussing et al., 2018). Anopheles albitarsis and An. aquasalis also transmit arboviruses, and some species also transmit Wuchereria bancrofti.
Species of subgenus Nyssorhynchus are restricted to the Neotropical Region, except for An. albimanus, which extends into the Nearctic Region (northern Mexico and along the Rio Grande River in Texas, USA).
Lane, 1953 (taxonomy); Cova-Garcia, 1961 (Venezuela); Forattini, 1962 (taxonomy); Faran, 1980 (Albimanus Section); Faran & Linthicum, 1981 (Amazonia); Clark-Gil & Darsie, 1983 (Guatemala); Linthicum, 1988 (Argyritarsis Section); Wilkerson & Strickman, 1990 (keys, Central America and Mexico); Peyton et al., 1992 (taxonomy); Bourke et al., 2011 (Myzorhynchella Section, phylogeny); Foster et al., 2017 (phylogenetic relationships).
albertoi Unti, 1941
albimanus Wiedemann, 1820
albitarsis Lynch Arribálzaga, 1878
anomalophyllus Komp, 1936
antunesi Galvão & Franco do Amaral, 1938
aquasalis Curry, 1932
argyritarsis Robineau-Desvoidy, 1827
arthuri Unti, 1941
atacamensis González & Sallum, 2010
benarrochi Gabaldon, Cova-Garcia & Lopez, 1941
braziliensis (Chagas, 1907)
darlingi Root, 1926
dominicanus Zavortink & Poinar, 2000 (fossil species, 33.9–40.4 Mya, Cenozoic, Eocene)
deaneorum Rosa-Freitas, 1989
dunhami Causey, 1945
evansae (Brèthes, 1926)
galvaoi Causey, Deane & Deane, 1943
goeldii Rozeboom & Gabaldon, 1941
guarani Shannon, 1928 (in Dyar, 1928)
halophylus Silva do Nascimento & Lourenço-de-Oliveira, 2002
ibiapabaensis (Sant’Ana & Sallum, 2023)
ininii Senevet & Abonnenc, 1938
jamariensis (Sant'Ana & Sallum, 2022)
janconnae Wilkerson & Sallum, 2009 (in Motoki et al., 2009)
konderi Galvão & Damasceno,1942
lanei Galvão & Franco do Amaral, 1938
lutzii Cruz, 1901
marajoara Galvão & Damasceno, 1942
nigritarsis (Chagas, 1907)
nuneztovari Gabaldon, 1940
oryzalimnetes Wilkerson & Motoki, 2009 (in Motoki et al., 2009)
oswaldoi (Peryassú, 1922)
parvus (Chagas, 1907)
pictipennis (Philippi, 1865)
pristinus Nagaki & Sallum, 2010 (in Nagaki et al., 2010)
rangeli Gabaldon, Cova-Garcia & Lopez, 1940
rondoni (Neiva & Pinto, 1922)
rondoniensis (Sant’Ana & Sallum, 2022)
sanctielii Senevet & Abonnenc, 1938
sawyeri Causey, Deane, Deane & Sampaio, 1943
striatus Sant'Ana & Sallum, 2016
strodei Root, 1926
tadei Saraiva & Scarpassa, 2021
triannulatus (Neiva & Pinto, 1922
trinkae Faran, 1979
untii (Sant’Ana & Sallum, 2023)