Classification of genus Anopheles began more than 100 years ago with the works of Theobald, who proposed numerous genera based on the character and distribution of thoracic and abdominal scales. These genera were ‘ill-defined and fancifully differentiated’, and this led to considerable dissatisfaction with Theobald’s system (Knab, 1913). The current system of subgeneric classification is based primarily on the number and positions of specialised setae on the gonocoxites of the male genitalia, and this basis of classification has been accepted since it was introduced by Christophers (1915). Christophers proposed three generic subdivisions which Edwards (1921) and Root (1923) formally recognised as the subgenera Anopheles, Myzomyia (=Cellia) and Nyssorhynchus. Edwards (1932) adopted this system and added subgenus Stethomyia in his classical treatise on family Culicidae. This system recognised Kerteszia as an informal group within subgenus Nyssorhynchus. Kerteszia was elevated to subgeneric status by Komp (1937). Subgenus Lophopodomyia was proposed by Antunes (1937) and subgenus Baimaia by Harbach et al. (2005). Subgenus Christya was resurrected from synonymy with Anopheles Meigen for An. implexus (Theobald) and An. okuensis Brunhes, Le Goff & Geoffroy by Harbach & Kitching (2015).
The internal classification of genus Anopheles (see attachment below) is based primarily on the schemes proposed by Edwards (1932), Reid & Knight (1961), Grjebine (1966), Gillies & de Meillon (1968), Reid (1968), Faran (1980) and Linthicum (1988). These schemes were amalgamated and updated by Harbach (1994, 2004, 2013). The classification presented here reflects all taxonomic changes and species recognised to date.
Infrasubgeneric categories have no formal status under the International Code of Zoological Nomenclature (1999). They are convenience categories only, often based on superficial similarities that may not indicate natural relationships. The informal categories now generally accepted by mosquito systematists include Sections, Series, Groups, Subgroups and Complexes. These are the categories of informal group taxa recognised here. The practice of constructing group names by placing the term denoting the level of classification after the specific name of the group (Reid & Knight, 1961), i.e. Arribalzagia Series rather than Series Arribalzagia, is followed. Furthermore, since informal group names are not regulated by the Code, they are treated as vernacular names in the manner promulgated by Belkin (1962) and explained by Peyton (1990). These names are printed in Roman type with the first letter capitalised even though the name of a nominal species or other formal taxon precedes the term (capitalised) denoting the level of classification, e.g. Maculipennis Group and Gambiae Complex. Alternatively, in situations where this practice might be unacceptable, an italicized binomen or other scientific name may be used in combination with the term (not capitalised) denoting the level of classification, e.g. Anopheles maculipennis group and Myzomyia series. In the case of binomina, the generic name or its abbreviation should always be used in the combination as specific names should not stand alone, except in unusual situations such as the listing of specific epithets assigned to a specified genus, e.g. the classification of Anopheles species presented here.
Taxonomic categories are basically subjective groupings of subordinate taxa that are defined by the included species. Which species are included in individual groups, and which morphological and biological characteristics of these species are used to define the groups, depends entirely upon the judgement and experience of the taxonomist. For this reason it is not possible to provide objective definitions for the various informal categories of classification now recognised within the genus.
Only extant species of Anopheles are included in the classification. Taxa are arranged alphabetically within groups, and the groupings at each level of classification are believed to represent phylogenetically related assemblages of species based principally on morphological similarity. However, some groupings contain one or more species of uncertain relationship with members of subordinate groups. These species are listed before the subordinate groups as unassigned members of the higher taxon. The authorities who first introduced or most recently redefined the informal taxonomic groups are indicated by literature citations. References for species complexes are those that include the first mention or treatment of all the species currently recognized within the group.
Reid & Knight (1961) revised Edwards’s (1932) infrasubgeneric classification of subgenus Anopheles. These authors divided the subgenus into two sections based on the shape of the pupal trumpet. The Laticorn Section was created for species with a wide funnel-shaped trumpet having the longest axis transverse to the stem, and the Angusticorn Section for species with a semi-tubular trumpet having the longest axis vertical more or less in line with the stem. These Sections were subsequently divided into Series, and the larger Series were divided into species Groups. The term ‘section’ was substituted for the ‘group’ category of Edwards to avoid confusion with ‘species group’ (which is equivalent to Group of current convention). The new classification retained the ‘Anopheles series’ of Edwards, but the original Arribalzagia and Christya ‘groups’ were reduced to ‘series’. Except for a few minor changes made at or below the ‘group’ level (Reid, 1968; Harrison, 1972; Vu et al., 1991; changes noted by Harbach, 2004), and the removal of An. kyondawensis to subgenus Baimaia (Harbach et al., 2005), the internal classification of subgenus Anopheles has changed relatively little since it was introduced by Reid & Knight (1961).
Grjebine (1966), Reid (1968) and Gillies & de Meillon (1968) are largely responsible for the existing internal classification of subgenus Cellia, although this system is based on Edwards (1932). For the most part, these authors dealt with the Anopheles of Madagascar, Southeast Asia and Africa, respectively. Reid (1968) and Gillies & de Meillon (1968) independently substituted the term ‘series’ for the ‘group’ categories in Edwards’ classification of Cellia (as Myzomyia). As Harrison (1980) noted, Gillies & de Meillon also introduced the term ‘section’ as a category below series level even though ‘species group’ was already in use for this level of taxon and ‘section’ as in use as a category above the series level (Reid & Knight, 1961). Inasmuch as Gillies & de Meillon interchanged the terms section and group, e.g., Funestus Section (pp. 2, 17) and Funestus Group (p. 128), Harbach (1994) replaced their ‘section’ with ‘group’ to correspond with the convention followed by other authors. This action added uniformity to the internal classification of genus Anopheles.
The internal classification of subgenus Nyssorhynchus has a complicated history. Christophers (1924) included Kerteszia in subgenus Nyssorhynchus, and Edwards (1932) followed this concept rather than the schemes proposed by either da [no-lexicon]Costa[/no-lexicon] Lima (1928), who recognised Kerteszia and Nyssorhynchus as subgenera of genus Nyssorhynchus, or Dyar (1928), who treated these taxa as distinct subgenera of genus Anopheles. Edwards (1932) divided subgenus Nyssorhynchus into three groups. Groups A (Nyssorhynchus) and B (Myzorhynchella) together equate the current concept of subgenus Nyssorhynchus, and Group C (Kerteszia) equates subgenus Kerteszia. Kerteszia was again treated as a subgenus by Komp (1937, 1942), but its independent status remained in doubt (Zavortink, 1973) until Peyton et al. (1992) firmly established the uniqueness of this group.
Edwards (1932) subdivided his Group A (Nyssorhynchus) into the Argyritarsis, Tarsimaculatus and Rondoni series. This system remained unchanged until Gabaldon (1940) referred to the Tarsimaculatus Series as the Albimanus Series and divided it into the Albimanus, Oswaldoi and Triannulatus subseries. These categories were accepted by Gabaldon & Cova Garcia (1952) who placed additional species in the Triannulatus and Oswaldoi subseries.
Da Costa Lima (1928) split subgenus Nyssorhynchus into two groups, Groups A and B, which partially correspond to the contemporary Argyritarsis and Albimanus sections introduced by Faran (1980). In another scheme, Galvão (1943) divided the subgenus into two series, the Argyritarsis and Tarsimaculatus series, which correspond to Groups A and B of da Costa Lima (1928). Galvão further divided the Argyritarsis Series into two complexes (the albitarsis and argyritarsis complexes), and recognised a monotypic group (albimanus) and three complexes within the Tarsimaculatus Series (the rondoni, tarsimaculatus and triannulatus complexes). Levi Castillo (1949) divided the subgenus into the Albimanus and Argyritarsis groups, and included three series within the former. These series, the Albimanus, Oswaldoi and Triannulatus series, corresponded with the subseries of Gabaldon (1940), except that An. galvaoi and An. nuneztovari were shifted from the Oswaldoi Subseries to the Triannulatus Series.
The most recent treatments of subgenus Nyssorhynchus include Faran (1980), Linthicum (1988) and Peyton et al. (1992). Faran divided the subgenus into the contemporary Albimanus and Argyritarsis Sections and recognised the ‘Myzorhynchella group’. The latter was originally described as a genus by Theobald (1907), recognised as a group (Group B) by Edwards (1932), treated as a subgenus by Galvão (1941) and recently defined as a Section of coequal rank with the Albimanus and Argyritarsis Sections by Peyton et al. (1992). Faran (1980) and Linthicum (1988) revised the Albimanus and Argyritarsis Sections, respectively. Faran (1980) divided the Albimanus Section into the Albimanus (monotypic) and Oswaldoi Groups, separated the Oswaldoi Group into the Triannulatus (monotypic) and Oswaldoi Subgroups, and split the Oswaldoi Subgroup into the Oswaldoi and Strodei Complexes (later called the Oswaldoi and Strodei Series by Faran & Linthicum (1981). Linthicum’s (1988) classification of the Argyritarsis Section was first presented in Faran & Linthicum (1981). These authors divided the Argyritarsis Section into the Argyritarsis and Albitarsis Groups, subdivided the Argyritarsis Group into the Argyritarsis, Lanei (monotypic), Pictipennis (monotypic) and Darlingi (monotypic) Subgroups, and recognised the Albitarsis and Braziliensis (monotypic) Subgroups within the Albitarsis Group.
In view of the previous terms applied to different levels of classification within subgenus Nyssorhynchus, Harbach (1994) recognised the ‘groups’ and ‘subgroups’ of Faran (1980) and Linthicum (1988) and the ‘complexes’ of Faran (1980) as Series, Groups and Subgroups, respectively, to bring the informal classification of subgenus Nyssorhynchus more in line with that established for subgenera Anopheles and Cellia.