Abstract

The phylogenetic relationships and generic assignments of ‘Finlaya’ and related taxa of uncertain taxonomic position in the classification of Aedini proposed by Reinert et al. (2004) are explored using 232 characters from eggs, fourth-instar larvae, pupae, adults and immature habitat coded for 116 exemplar species. The ingroup comprises 74 species, including 41 currently classified as ‘Finlaya’, 25 formerly placed in this group and eight related species. The outgroup includes four non-aedine species and 38 aedine species representing all generic-level taxa of the polyphyletic ‘Ochlerotatus’ and major clades within the polyphyletic ‘Aedes’ in figure 4 of Reinert et al. (2004). Data are analysed in a total-evidence approach using implied weighting. The analysis produced four most parsimonious cladograms and ambiguity among the four is relatively minor concerning just four parts of the cladograms, two of which are linked. The strict consensus tree corroborates the monophyly of the 13 genera proposed by Reinert et al. (2004) that are included in the analysis. Overall, the results show remarkable congruence with those of Reinert et al. (2004) despite differences in the taxa included and morphological characters examined in the two studies. Seventeen genera, of which 11 are new, are proposed for monophyletic clades based on the strength of character support and application of the principle of equivalent rank. Bruceharrisonius stat. nov., Macleaya stat. nov., Molpemyia stat. nov. and Pseudoskusea stat. nov. are raised from subgeneric rank within ‘Ochlerotatus’; Hulecoeteomyia stat. nov. and Phagomyia stat. nov. are resurrected from synonymy with Finlaya; and Collessius gen. nov. (Ochlerotatus macfarlanei Edwards), Dahliana gen. nov. (Culex geniculatus Olivier), Dobrotworskyius gen. nov. (Aedes tubbutiensis Dobrotworsky), Georgecraigius gen. nov. (Culex atropalpus Coquillett), Gilesius gen. nov. (Culex pulchriventer Giles), Himalaius gen. nov. (Finlaya gilli Barraud), Jarnellius gen. nov. (Culex varipalpus Coquillett), Jihlienius gen. nov. (Aedes chungi Lien), Patmarksia gen. nov. (Leucomyia australiensis variety papuensis Taylor), Rampamyia gen. nov. (Culex notoscriptus Skuse) and Vansomerenis gen. nov. (Aedes pulchrithorax Edwards) are introduced for the type species indicated in parentheses, and their allies. Additionally, Horsfallius subgen. nov. (Culex fluviatilis Lutz) and Lewnielsenius subgen. nov. (Aedes muelleri Dyar) are introduced as subgenera of Georgecraigius and Jarnellius, respectively. As is usual with generic-level groups of Aedini, these genera are polythetic taxa that are diagnosed by unique combinations of characters. The analysis also shows that Oc. (Protomacleaya) is a polyphyletic assemblage of species, which is retained as a non-monophyletic taxon until the included species can be classified into monophyletic groups.

INTRODUCTION

Aedini (sensuBelkin, 1962) is the largest tribe in family Culicidae (Diptera). It includes 1235 currently recognized species, 32 subspecies, nine infrasubspecific forms/varieties and nine nomina dubia. Reinert, Harbach & Kitching (2004) conducted a phylogenetic analysis of higher-level relationships within the tribe using 172 morphological characters from all life stages and 119 exemplar species representing the 12 genera and 56 subgenera that were recognized at the time. All of the genera except Aedes and Ochlerotatus were recovered as monophyletic. (The authors of all aedine taxa are cited in Appendix 4.) In addition to 12 formerly recognized genera, Reinert et al. (2004) proposed generic status for 32 subgenera of Aedes and Ochlerotatus, raised a clade consisting of Oc. (Finlaya) and its relatives (Kochi Group) to generic rank as genus Finlaya, elevated Downsiomyia from synonymy with Finlaya as the generic name for a clade comprising Oc. (Fin.) niveus and its relatives (Niveus Group), and described a new genus, Tanakaius, to accommodate Oc. (Fin.) togoi and Oc. (Fin.) savoryi (Togoi Group). Genus Aedes was restricted to the former subgenus Aedes and genus Ochlerotatus was limited to a clade consisting of the type species, six related species and members of subgenus Rusticoidus. These actions left species previously included in subgenera Aedimorphus and Cancraedes of Aedes and subgenera Bruceharrisonius, Chaetocruiomyia, Finlaya, Macleaya, Molpemyia, Ochlerotatus, Protomacleaya and Pseudoskusea of Ochlerotatus without generic assignments. The present study is the first step toward resolving the relationships and generic placements of these taxa. It treats species and species-groups of uncertain taxonomic position (incertae sedis) that were previously included in subgenus Finlaya of Ochlerotatus, and species previously removed from Finlaya by Zavortink (1972) and placed in subgenera Protomacleaya and Ochlerotatus of Ochlerotatus.

Edwards (1932) provided the first comprehensive classification of Finlaya Theobald (as a subgenus of Aedes). He recognized eight groups within Finlaya (designated groups A–H) based on a few morphological characters of adults. Barraud (1934) followed Edwards' arrangement of genera and subgenera. Knight (1948) recognized that Edwards' Group F contained more or less unrelated forms and therefore divided it into eight divisions (denoted by Roman numerals). Knight & Marks (1952) revised and expanded the classification of Edwards and added numerous subgroup categories. They also based their classification almost exclusively on adult characters and indicated that a number of the groups were unnatural assemblages. Whereas Knight & Hull (1951), when examining the Philippine fauna, utilized the group categories of Knight & Marks (1952) (they noted this paper was in press), subsequent authors, i.e. Bohart (1957), Marks & Hodgkin (1958), Macdonald & Traub (1960), Lee et al. (1982) and Lu & Ji (1997), provided some modifications to their classification, again based primarily on adult characters. Dobrotworsky (1965) followed Knight & Marks' arrangement and applied it to the ten species occurring in Victoria, Australia. Marks (in Lee et al., 1982) stated that Knight & Marks' ‘ … subdivision of the groups of subgenus Finlaya was based on adult characters only. These groups and subgroups are by no means sacrosanct and we already know, from life history studies, that reassessment of the present placing of some species will be needed. This applies particularly to rockpool species.’Belkin (1962) followed the classification of Knight & Marks (1952) but stated that Finlaya as understood at present is a large heterogeneous assemblage of species … and ‘I believe that a natural classification of Finlaya cannot be developed until the immature stages are carefully studied.’Tanaka, Mizusawa & Saugstad (1979) recognized that previous attempts at classification of Finlaya did not reflect a natural classification and proposed nine groups for species of Finlaya occurring in the Japanese and Korean area based primarily on characters of the male genitalia and larvae. Schick (1970) revised the Terrens Group and Arnell & Nielsen (1972) revised the Varipalpus Group based on characters of the adult and immature stages. Zavortink (1972) removed all New World species from Finlaya and placed them in subgenera Aztecaedes, Gymnometopa, Protomacleaya and Ochlerotatus of Aedes and subgenus Conopostegus of Haemagogus based on features of the adults, pupae and larvae. Reinert (1990, 1993, 1999b, 2003) transferred some species to other subgenera, i.e. Kenknightia, Molpemyia, Zavortinkius and Bruceharrisonius. Reinert (2000a) elevated Ochlerotatus to generic rank and transferred subgenus Finlaya to that genus. Reinert (2002e) separated subgenus Finlaya and related subgenera of Ochlerotatus into Assemblages and Subassemblages based on features of the female genitalia. He mentioned that ‘Subgenus Finlaya, as currently configured, contains several diverse groups that are markedly different in the female genitalia; therefore, the above definition of the subgenus is somewhat imprecise and indicates the need for a revision of the entire taxon. However, this revision should include all life stages as suggested by Belkin (1962). Finally, Reinert et al. (2004), as noted above, formally limited genus Finlaya to the Kochi Group.

Few molecular phylogenetic studies of aedine taxa have been published. Wesson, Porter & Collins (1992) examined the relationships of six aedine species (four ‘Aedes’, one Haemagogus, one Psorophora) based on rDNA ITS2 sequence data. Besansky & Fahey (1997) included three ‘Aedes’ and one Haemagogus among 13 mosquito species in a study of relationships based on sequence data for the nuclear protein-coding white gene. Kumar, Black & Rai (1998) investigated the relationships of 15 culicine species representing six ‘genera’, including nine species of ‘Aedes’ and one each of Armigeres and Haemagogus. More recently, Cook et al. (2005) investigated the relationships among 20 species of ‘Aedes’ representing six traditionally recognized subgenera, including Aedes (1 species), Aedimorphus (1), Diceromyia (2), Halaedes (3), Ochlerotatus (10) and Stegomyia (3), based on sequence data for the cytochrome oxidase c subunits I and II (COI, COII) of mtDNA. Finally, Behbahani et al. (2005) used ITS2 sequences to construct a phylogenetic tree for five species of Stegomyia. The results of Wesson et al. (1992), Besansky & Fahey (1997) and Kumar et al. (1998) indicated a paraphyletic Aedini, but these studies provided little insight into the phylogeny and classification of the tribe because they included very few generic-level taxa and few species. For example, whereas St. aegypti and St. albopicta formed a strongly supported sister pair in the white-gene phylogeny of Besansky & Fahey (1997), they were not placed in a congeneric relationship in the ITS2 phylogeny of Wesson et al. (1992) or the RFLP phylogeny of Kumar et al. (1998). All three of these studies included Ochlerotatus (Protomacleaya)’triseriatus, which was strongly paired with Psorophora in the phylogeny of Wesson et al. (1992) and with Haemagogus in the phylogenies of Besansky & Fahey (1997) and Kumar et al. (1998). It is interesting to note that Wesson et al. (1992) suggested that Aedes sensu auctorum should be split into two genera, one including Oc. (Pro.)’triseriatus [as Ae. (Pro.) triseriatus] along with species of Haemagogus and Psorophora. The COI and COII phylogenies of Cook et al. (2005) agreed with respect to species-level groupings, but because sequences for neither gene were available for all the species analysed, it was ‘difficult to draw firm conclusions regarding the interspecies relationships’. Support for supraspecific clades was either very low or meaningless in cases where higher-lever taxa were represented by several samples of a single species. In general, clade support, assessed by bootstrap proportions, was slightly higher for the COII phylogeny. Whereas the monophyly of Di. taylori+Di. furcifer (as Ae. taylori and Ae. furcifer) was weakly supported in the COI phylogeny, it was very strongly supported (bootstrap value = 99) in the COII phylogeny. Likewise, the monophyly of St. luteocephala+ (St. albopicta+St. aegypti) (as species of Aedes) received low support in the COI phylogeny and modest support (bootstrap value = 53) in the maximum-likelihood tree of COII sequences. Eight species of Ochlerotatus included in the analysis of COI data (COII data were not available) were placed in a sister-group relationship to Oc. (Rusticoidus) rusticus (as Ae. rusticus), which was sister to Ae. cinereus, but all of the internal branches were only weakly supported. The three species of Halaedes (not included in the COI phylogeny) formed a strongly supported clade in the COII phylogeny (bootstrap value = 99) comprising ashworthi + (wardangensis+australis) (as Aedes species). Overall, the results of Cook et al. (2005) are not inconsistent with those of Reinert et al. (2004).

As noted above, Reinert et al. (2004) formally recognized 46 genera for most of the species previously included in 12 genera. Many species previously included in two subgenera of former genus Aedes and eight subgenera of former genus Ochlerotatus, including subgenus Finlaya, could not be placed in any of the 46 genera, and are regarded as incertae sedis (i.e. of uncertain placement) within Aedini. To avoid confusion, species of uncertain generic and subgeneric placement, and generic-level names used in the previous sense that include these species, are enclosed within single quotation marks herein to distinguish them from formally recognized monophyletic taxa. The objective of the present study is to clarify the relationships and generic assignments of ‘Finlaya’ and related taxa of uncertain taxonomic position in the classification proposed by Reinert et al. (2004). Herein we present the results of cladistic analysis of these taxa and propose further changes to the classification of Aedini that more accurately reflect natural affinities.

MATERIAL AND METHODS

Taxa examined

The study taxa (Appendix 1; see also for their authorship and geographical distribution) include an ingroup of 74 species. This group comprises species of ‘Finlaya’ and Protomacleaya that were placed within the polyphyletic assemblage of Ochlerotatus basal to ‘Aedes’ in figure 4 of Reinert et al. (2004), i.e. species included in the clades between Oc. (Fin.)’chrysolineatus and Oc. (Fin.)’poicilius (now Finlaya poicilia), and incertae sedis species listed in their appendix 4. The ingroup also includes species that were classified as ‘Finlaya’ prior to Zavortink (1972). The outgroup taxa include both aedine and non-aedine species. The ‘aedine outgroup includes representatives of all generic-level taxa arrayed in the basal polyphyletic assemblage of ‘Ochlerotatus’ in figure 4 of Reinert et al. (2004), as well as species selected from major clades within the distal polyphyletic assemblage of ‘Aedes’. The ‘non-aedine outgroup comprises the four outgroup taxa used in the analyses of Reinert et al. (2004), i.e. Culex quinquefasciatus, Culiseta inornata, Mansonia titillans and Orthopodomyia signifera.

We initially intended to include several additional species of ‘Finlaya’ in the study but this was not possible because specimens were unavailable, several life stages were unknown, and/or published descriptions and illustrations were inadequate. Taxa examined (Appendix 1) include type species of most genus-level names (abbreviations for genera and subgenera follow Reinert, 2001a; Reinert et al., 2004). However, we did not include type species if they had several missing life stages or inadequate specimens for detailed study.

Generally, we examined 3–6 specimens (range 1–20) of each life stage and structure for each species. However, for some life stages of a few species no material was available and literature descriptions were either inadequate or lacking this information, e.g. pupae of Oc.echinus, Oc.pulchriventer and Oc.rubrithorax, and eggs of many species. We also studied individually reared, pin-mounted adults with associated slide-mounted fourth-instar larval and pupal exuviae where available. A phase contrast or differential interference contrast microscope with 400× magnification is needed to observe the very slender distal parts of many setae for the measurement of total length. Lesser magnification normally results in incomplete measurements, as noted in many published illustrations of larval and pupal stages. Pinned adults were examined with a binocular stereomicroscope utilizing cold white light delivered by a fibre-optic illumination system. An adjustable examination stage with biaxial rotation capability (see figure 125.3 in Russell et al., 1963) allowed observation and lighting of specimens at any angle.

A few characters used in the analysis of Reinert et al. (2004) are excluded from the present study because all the study taxa exhibit the same character state. Some characters were modified by adding or deleting character states or altering the range of continuous variables to score the study taxa more accurately. A number of new characters are included that were not used by Reinert et al. (2004). Character states corrected in the present study from those incorrectly entered in Appendix 2 of Reinert et al. (2004) (in parentheses) are: Oc. (Brh.)’greenii = 25 (19), 67 (48), 68 (49); Oc. (Mac.)’tremulus= 81 (59), 114 (88), 183 (137); Oc. (Fin.)’chrysolineatus= 188 (140). The discussion following each character deals primarily with taxa treated in the present study.

Specimens examined were from the following collections: National Museum of Natural History, Smithsonian Institution, Washington, DC; The Natural History Museum, London, UK; Department of Zoology and Entomology, University of Queensland, Brisbane, Australia; Australian National Insect Collection, CSIRO Division of Entomology, Canberra, Australia; Florida State Collection of Arthropods, Division of Plant Industry, Gainesville, Florida; Laboratoire/Cellule Entomologie, EID Méditerranée, Montpellier, France; Bohart Museum, University of California, Davis, California; National Institute of Malariology, Parasitology and Entomology, Hanoi, Vietnam; Florida Medical Entomology Laboratory, University of Florida, Vero Beach, Florida; and the first author.

CHARACTER DESCRIPTIONS

The data (Appendix 2) comprise 232 characters from eggs (3), fourth-instar larvae (74), pupae (31), females (71), males (11), female genitalia (16), male genitalia (25) and habitat of immature stages (1). Characters from Reinert et al. (2004) are included in parentheses following current character numbers. Data for the characters were derived from direct observations except where life stages were not available or structures were missing from available specimens. We coded some such missing data from literature sources (e.g. characteristics of most eggs) or collection data sheets. The remaining missing data were denoted by a ‘?’. States in continuous characters were determined either by clear gaps in the observed counts or measurements (e.g. character 28), or by reference to observed intraspecific variation (e.g. character 38). Characters that could not be scored owing to the absence of homologous structures (‘dependent characters’) are indicated by a dash (–), e.g. character 102 in Mansonia. All multistate characters were treated as unordered. Polymorphic characters were explicitly coded as exhibiting only those states observed.

Adult characters were derived from females unless otherwise noted. Males of many species often have fewer setae and scales than females, and would skew the coding of some character states. We have reinterpreted the homologies of several structures from those used in some published works.

Anatomical nomenclature and chaetotaxy follow Harbach & Knight (1980, 1982) except for terms proposed by Reinert (1990, 1999b, 2000b, 2002a). Many structures of characters used in the present study are illustrated in these references. However, references to character states illustrated in other publications are provided in the explanations of the characters listed below.

Eggs

Larvae (fourth-instars)

  • 4(4). Labiogula: (0) short, length < width; (1) moderate to long, length ≥ width. The width of the labiogula is measured from the outer margins of the posterior tentorial pits and its length is measured from the caudolateral angle of the dorsomentum to the posterior margin of the collar. See Belkin (1962: figs 218, 243, Ge. longiforceps and Fl. franclemonti) for examples of state (0) and his figures 234, 277, Oc. (Fin.) argyronotum and Mu. alternans, for state (1).

  • 5(5). Antenna, length: (0) short, ≤ 0.40 median length of dorsal apotome; (1) moderate to long, ≥ 0.42 median length of dorsal apotome. Antennal length is determined by dividing the antennal shaft length by the dorsal apotome median length (preferably in exuviae). Antennae vary from short (state 0) (see Tanaka et al., 1979: figs 99, 101, Ta. togoi and Ta. savoryi) to long (state 1) (see Tanaka et al., 1979: figs 102, 104, Oc. (Fin.)’seoulensis and Oc. (Brh.) alektorovi (= kobayashii Nakata); Hopkins, 1952: fig. 62, Oc. (Fin.) pulchrithorax).

  • 6(new). Antenna, spicules: (0) absent; (1) few; (2) numerous. Some aedine species have antennae without spicules (state 0), e.g. Abraedes (see Zavortink, 1972: fig. 34, Ab. papago), Aztecaedes, Finlaya, Gymnometopa, Halaedes, Kompia and Stegomyia. Other species have a few, normally small and scattered spicules (state 1), e.g. Downsiomyia, Haemagogus, Howardina, Kenknightia (see Reinert, 1990: fig. 24, Ke. dissimilis) and Zavortinkius. Many species have numerous, normally well-developed spicules covering all or most of the antenna (state 2), e.g. Aedes (see Tanaka et al., 1979: fig. 134, Ae. yamadai), Mucidus, Psorophora, Tanakaius and Oc.(Rusticoidus).

  • 7(7). Seta 1-A, length: (0) short, ≤ 3.0 times antennal width at point of attachment; (1) longer, ≥ 3.1 times antennal width at point of attachment. The usual condition of seta 1-S is moderately long to long (state 1) in most Aedini, e.g. Oc.(Protomacleaya) (see figures in Zavortink, 1972) and Zavortinkius, but this seta is short in some taxa (state 0), e.g. Opifex (Opi.) fuscus and most Stegomyia (see Belkin, 1962: figs 208, 314, Op. (Opi.) fuscus and St. aegypti).

  • 8(6). Seta 1-A, development: (0) single or 2-branched; (1) ≥ 3 branches. Many species of Aedini have seta 1-A single (state 0) (see Belkin, 1962: fig. 231, Oc. (Fin.) albilabris) whereas others are multiple-branched (state 1) (see Belkin, 1962: fig. 234, Oc. (Fin.) argyronotum). However, this seta is single or two-branched (state 0) in some species, and it is also two-branched in some specimens of a few species in which seta 1-A is multiple-branched, which are scored (0,1).

  • 9(8). Seta 2,3-A, position: (0) insertion apical or nearly apical; (1) insertion noticeably subapical. Setae 2-A and 3-A are inserted apically on nearly apically on the antennae of most Aedini (state 0); however, they are noticeably subapical (state 1) in Rhinoskusea (see Reinert, 1976b: figs 12–14) and the outgroup species Ma. titillans.

  • 10(9). Seta 1-C, development: (0) spiniform; (1) single, thinner, distal part attenuate; (2) forked or branched, proximal part stout. Spiniforms (state 0) are single setae, thickened throughout with a bluntly pointed or rounded apex (see Zavortink, 1972: fig. 56, Hg. (Con.) leucotaeniatus). Setae exhibiting state (1) are single, moderately to very thick proximally but attenuate distally (see Belkin, 1962: fig. 218, Ge. longiforceps; Hopkins, 1952: fig. 62, Oc. (Fin.) pulchrithorax). Those exhibiting state (2) are thickened proximally and have two or more branches distally. Seta 1-C is branched in species of Finlaya and Hw. walkeri (see Berlin, 1969: fig. 18, for illustration of state 2).

  • 11(10). Seta 4-C, position: (0) insertion anterior to seta 6-C; (1) insertion at same level or posterior to seta 6-C. The position of seta 4-C appears to be constant in generic-level taxa for which the taxonomy is stable. However, both states occur in some of the larger subgenera, but in different species groups. See Belkin (1962) for illustrations of state (0) (his figs 218, 277, Ge. longiforceps and Mu. alternans) and state (1) (his fig. 211, Ha. australis). See character 15 for comment.

  • 12(11, in part). Seta 4-C, length: (0) short; (1) moderate; (2) long; (3) very long. The lengths of setae 4–7-C are compared to the median length of the dorsal apodeme (DAp), preferably in exuviae. The setae are considered short (≤ 0.19 length of DAp), moderate (0.21–0.38 length of DAp), long (0.40–0.81 length of DAp) or very long (≥ 0.90 length of DAp).

  • 13(new). Seta 5-C, length: (0) short; (1) moderate; (2) long; (3) very long. See character 12 for explanation of setal lengths.

  • 14(new). Seta 5-C, development: (0) single; (1) branched. Larval seta 5-C is single (state 0), e.g. Abraedes, Aztecaedes, Finlaya, Howardina and Zavortinkius. Other taxa have the seta branched (state 1), e.g. Aedes, Downsiomyia, Geoskusea, Levua and Tanakaius.

  • 15(12). Seta 6-C, position: (0) insertion anterior to seta 7-C; (1) insertion at same level or posterior to seta 7-C. The positions of setae 6-C and 7-C are determined on head capsules that are horizontal with the thorax and abdomen, normally exuviae. If the head is in a prognathous orientation and turned down anteriorly, these and other setae may appear to be positioned differently. This also applies to characters 11 and 17. See illustrations in Tanaka et al. (1979: figs 99, 101, Ta. togoi and Ta. savoryi) for state (0) and their fig. 128 (Ge. baisasi) for state (1).

  • 16(new). Seta 6-C, length: (0) short; (1) moderate; (2) long; (3) very long. See character 12 for explanation of setal lengths.

  • 17(13). Seta 7-C, position: (0) insertion anterior to seta 5-C; (1) insertion at approximately same level as seta 5-C; (2) insertion posterior to seta 5-C. See Belkin (1962: figs 237, 239, 241, etc., Finlaya, as Kochi Group) for examples of state (0), Zavortink (1972: fig. 34, Ab. papago) for state (1) and Tanaka et al. (1979: figs 99, 101, Ta. togoi and Ta. savoryi) for state (2). See character 15 for comment.

  • 18(new). Seta 7-C, length: (0) short; (1) moderate; (2) long. See character 12 for explanation of setal lengths.

  • 19(14). Seta 12-C, position: (0) insertion mesal to or at same level as seta 13-C; (1) insertion lateral to seta 13-C. Seta 13-C is longer and normally borne lateral to seta 12-C in Culicidae. For this reason, the longer seta in a mesal position relative to the other in some species of Aedini is interpreted as seta 13-C (see fig. 277 of Mu. alternans in Belkin, 1962; labelled as 12-C).

  • 20(15). Seta 13-C, development: (0) single; (1) branched. Seta 13-C is normally single (state 0) in many of the species examined. However, it is branched (state 1) in a number of other taxa, e.g. Aedes, Geoskusea, Kenknightia, Levua, Mucidus, Psorophora, Rhinoskusea and Zavortinkius.

  • 21(new). Seta 14-C, development: (0) single; (1) branched. Seta 14-C is often single (state 0) (see Tanaka et al., 1979: fig. 101, Ta. savoryi) and may be slender or stiff and stout. The seta is branched (state 1) (see Tanaka et al., 1979: fig. 102, Oc. (Fin.) seoulensis) and may be slender or stellate in some species.

  • 22(16). Seta 19-C: (0) absent; (1) present. The occurrence of seta 18-C and the presence or absence of seta 19-C ventrally on the cervical membrane of larvae belonging to 29 genera, 72 subgenera and 331 species of Culicidae was reported by Hochman & Reinert (1974). See illustrations in Hochman & Reinert (1974) for examples and a discussion of the occurrence of seta 19-C in the large and heterogeneous genus Ochlerotatus sensu auctorum, and its taxonomic importance and possible phylogenetic significance.

  • 23(17). Ventromedian cervical sclerite: (0) absent; (1) present. A small solid or fragmented sclerite is present on the ventromedian area of the cervix in many aedine larvae. It appears to be absent in Ko. purpureipes, Le. geoskusea, Oc.(Cha.) wattensis, Oc.(Fin.) macfarlanei, Oc. (Mac.) tremulus, Psorophora, Rhinoskusea, Zavortinkius and the non-aedine outgroup species. Reinert (1976a) provided a review and illustrations of this sclerite in Culicidae.

  • 24(18). Setae 1–3-P: (0) not attached to a common setal support plate; (1) two or three of these setae attached to a common setal support plate. See Zavortink (1972: figs 18, 28, Oc. (Pro.) kompi and Oc. (Pro.) knabi) for examples of state (0) and his figs 9, 12, Oc. (Pro.) triseriatus and Oc. (Pro.) zoosophus, for state (1).

  • 25(19). Seta 1-P, length: (0) ≤ length of seta 2-P; (1) > length of seta 2-P. Seta 1-P is normally longer (state 1) than seta 2-P in Aedini, but it is shorter than or equal in length to 2-P (state 0) in Finlaya (as Kochi Group) (see Belkin, 1962: fig. 239, Fl. burnetti), Hw. walkeri, Sc. arboricola, Oc. (Och.) muelleri and some Oc. (Finlaya) .

  • 26(21). Seta 5-P, length: (0) ≤ length of seta 6-P; (1) > length of seta 6-P. The length of seta 5-P in relation to that of seta 6-P is relatively consistent for most generic-level taxa that are well defined. See Belkin (1962) for illustrations of state (0) (his fig. 243, Fl. franclemonti) and state (1) (his fig. 231, Oc. (Fin.) albilabris).

  • 27(22). Seta 5-P, development: (0) single; (1) branched. Seta 5-P is single (state 0) in several taxa, e.g. Aedes, Geoskusea, Hg.(Conopostegus), Isoaedes, Levua, Mucidus and Rhinoskusea. In other taxa, this seta is branched (state 1), e.g. Abraedes, Aztecaedes, Gymnometopa and Kompia.

  • 28(23). Seta 8-P, length: (0) ≤ 1.2 times length of seta 4-P; (1) ≥ 1.8 times length of seta 4-P. Seta 8-P is less than or equal to 1.2 times the length of seta 4-P (state 0) in most taxa examined (see Tanaka et al., 1979: fig. 99, Ta. togoi). It is greater than or equal to 1.8 times the length of seta 4-P in Aedes, Oc. (Rusticoidus), Oc. (Fin.) albotaeniatus, Oc. (Och.) fluviatilis, Ps. (Jan.) ferox, Ps. (Gra.) jamaicensis (see Belkin, Heinemann & Page, 1970: figs 60, 67) and the outgroup species Cs. inornata.

  • 29(new). Seta 8-P, development: (0) single or occasionally 2-branched; (1) multiple-branched (occasionally 3-branched). A few taxa have seta 8-P single or occasionally 2-branched (state 0) (see Belkin, 1962: figs 211, 277, Ha. australis and Mu. alternans). Most taxa have seta 8-P multiple-branched, but it is occasionally only 3-branched (state 1) (see Belkin, 1962: figs 239, 316, Fl. burnetti and Sc. albolineata).

  • 30(new). Seta 1-M, length: (0) ≤ 2.5 times length of seta 2-M; (1) ≥ 3.5 times length of seta 2-M. See Tanaka et al. (1979: figs 99, 111, Ta. togoi and Oc. (Fin.) oreophilus) for examples of state (0) and their fig. 104, Oc. (Brh.) alektorovi (=kobayashii), for state (1).

  • 31(new). Seta 1-M, development: (0) single (rarely 2-branched); (1) ≥ 3 branches. Illustrations of seta 1-M single (state 0) in Rhinoskusea are found in Reinert (1976b). Examples of seta 1-M with three or more branches (state 1) are found in Oc. (Protomacleaya) (see figures in Zavortink, 1972).

  • 32(25). Seta 4-M, development: (0) single; (1) branched. Many generic-level taxa of Aedini examined in the present study have seta 4-M branched (state 1) (see Schick, 1970: figs 16, 22, ‘Oc. (Pro.) ’ terrens and ‘Oc. (Pro.)’ berlini). See illustrations in Zavortink (1972: figs 32, 34, 37, 40, Ko. purpureipes, Ab. papago, Az. ramirezi and Gy. mediovittata) for examples of state (0).

  • 33(26). Seta 7-M, length: (0) < length of seta 5-M; (1) ≥ length of seta 5-M. Seta 7-M is normally shorter than seta 5-M (see Tanaka et al., 1979: fig. 98, Oc. (Fin.) hatorii) (state 0) in most Aedini; however, it is longer than seta 5-M (state 1) in Geoskusea, Kompia, Tanakaius (see Tanaka et al., 1979: fig. 99, Ta. togoi), Ha. australis, Oc.(Cha.) wattensis, Oc.(Fin.) albocinctus, Oc. (Fin.) gilli, Oc. (Mac.) tremulus, Ps. (Jan.) ferox and the outgroup species Cx. quinquefasciatus.

  • 34(27). Seta 2-T, development: (0) single; (1) branched. See Zavortink (1972: fig. 44, Oc. (Och.) muelleri) for an example of single seta 2-T (state 0) and Belkin (1962: fig. 211, Ha. australis) for an example of branched 2-T (state 1).

  • 35(new). Seta 4-T, development: (0) single; (1) ≥ 2 branches, not stellate; (2) ≥ 3 branches, stellate. Stellate setae have three to numerous, moderately stout to stout, stiff branches that are truncate or bluntly pointed (not attenuate) and radiate in different directions from or near the base (starburst-shaped) (see Belkin, 1962: fig. 237, Fl. bougainvillensis). Branches of stellate setae often are of different lengths. In some species, e.g. Az. ramirezi and Oc. (Pro.) zoosophus, this and some other setae approach the stellate condition except the branches are somewhat narrower and tend to be in one plane (fan-shaped). These are scored (1).

  • 36(28). Seta 3-I, development: (0) single; (1) branched. The development of seta 3-I is consistent for generic-level taxa (except some of the large ones), with few exceptions. Some specimens of a few species show an overlap of the two states.

  • 37(29). Seta 7-I, length: (0) < 0.45 length of seta 6-I; (1) ≥ 0.55 length of seta 6-I. Seta 7-I is normally moderately long to long (state 1) (≥ 0.55 length of seta 6-I, see Belkin, 1962: fig. 274, Le. geoskusea) in Aedini, but it is shorter in some species (state 0) (≤ 0.45 length of seta 6-I, see Belkin, 1962: fig. 208, Op.(Opi.) fuscus).

  • 38(30). Seta 7-I, development: (0) single to 3-branched; (1) ≥ 4 branches. Seta 7-I is single or occasionally with 2 or 3 branches (state 0) in most Aedini (see Belkin, 1962: fig. 227, Oc. (Fin.) notoscriptus). Seta 7-I has more than three branches (state 1) in Mucidus (see Belkin, 1962: fig. 277, Mu. alternans). This seta has overlapping states in Oc. (Fin.) keefei and Op. (Opi.) fuscus, and is scored (0,1).

  • 39(31). Seta 12-I: (0) absent; (1) present. Seta 12-I is absent (state 0) in Abraedes, Aedes, Aztecaedes, Gymnometopa, Howardina, Isoaedes, Kompia, Scutomyia, Stegomyia, Oc.(Cha.) wattensis, Oc. (Och.) muelleri, Oc. (Och.) varipalpus and Oc. (Pro.) zoosophus. Reinert (2000a) discussed the presence and absence of seta 12-I in generic-level taxa previously included in Aedes and Ochlerotatus. Most taxa examined in the present study have seta 12-I (state 1) (see Tanaka et al., 1979: fig. 86, Oc. communis).

  • 40(new). Seta 1-II, development: (0) single; (1) ≥ 2 branches, not stellate; (2) ≥ 3 branches, stellate. Illustrations in Belkin (1962) show the various states of seta 1-II, i.e. state (0) in fig. 234 (‘Oc. (Fin.) argyronotum), state (1) in fig. 231 (‘Oc. (Fin.) albilabris) and state (2) in fig. 237 (Fl. bougainvillensis).

  • 41(new). Seta 2-II, development: (0) single; (1) branched. Examples of seta 2-II development are illustrated in Belkin (1962: fig. 218, Ge. longiforceps) for state (0) and his fig. 237 (Fl. bougainvillensis) for state (1).

  • 42(new). Seta 5-II, development: (0) single; (1) ≥ 2 branches, not stellate; (2) ≥ 3 branches, stellate. Seta 5-II is branched in most Aedini, either not stellate (state 1) (see Belkin, 1962: figs 227–229, Oc. (Fin.) notoscriptus) or stellate (state 2) (see Belkin, 1962: fig. 237, Fl. bougainvillensis). This seta is single in a few taxa (state 0) (see Reinert, 1979: fig. 5, Ia. cavaticus).

  • 43(new). Seta 6-II, length: (0) < length of seta 6-III; (1) ≥ length of seta 6-III. See Tanaka et al. (1979: figs 102, 104, Oc. (Fin.) seoulensis and Oc. (Brh.) alektorovi (=kobayashii)) for state (0) and their figs 98, 99, Oc. (Fin.) hatorii and Ta. togoi, for state (1).

  • 44(32). Seta 6-II, development: (0) single; (1) branched. Seta 6-II is normally branched in Aedini, but it is single in Levua, Mucidus and Oc. (Molpemyia) of the ingroup, and also in other aedines, e.g. Alanstonea, Eretmapodites, subgenus Harbachius of Verrallina and Ar. (Arm.) breinli.

  • 45(33). Seta 7-II, development: (0) similar to seta 7-I; (1) different than seta 7-I. The length, thickness and branching of seta 7-II are normally consistent in generic-level taxa of Aedini, but in some of the larger taxa this character is consistent only at the species-group level. Length of seta 7-II is often slightly less than that of seta 7-I. See Belkin (1962: fig. 208, Op. (Opi.) fuscus) for an example of state (0) and his fig. 218 (Ge. longiforceps) for state (1). Seta 7-II is short, slender and 2-branched in Oc. (Och.) fluviatilis and differs from seta 7-I, which is long, stout, aciculate and single (see fig. 53 in Zavortink, 1972), and is scored (1).

  • 46(34). Seta 8-II, development: (0) single; (1) branched. See Zavortink (1972: figs 32, 34, 40, Ko. purpureipes, Ab. papago and Gy. mediovittata) for examples of single seta 8-II (state 0) and his figs 4, 7, 9 (‘Oc. (Pro.) hendersoni, Oc. (Pro.) brelandi and Oc. (Pro.) triseriatus) for examples of state (1).

  • 47(new). Seta 6-III, development: (0) single; (1) branched. Seta 6-III is single (state 0) is a few taxa, e.g. Aedes, Isoaedes, Mucidus, Ge. longiforceps, Oc. (Fin.) candidoscutellum, Oc. (Fin.) rubrithorax, Oc. (Mol.) pecuniosus, Oc. (Bruceharrisonius)’, Oc. (Ochlerotatus) (several species) and the outgroup species Ma. titillans and Or. signifera. This seta is branched in most taxa examined.

  • 48(35). Seta 3-V, length: (0) ≤ 1.55 times length of seta 5-V; (1) ≥ 1.90 times length of seta 5-V. Seta 3-V is at least two times longer (often much longer) than seta 5-V in species of Section I of Ochlerotatus (as defined by Reinert, 2000a) whereas it is shorter than 1.5 times the length of seta 5-V in his Section II. These measurements are slightly modified here. See Appendix 2 for other taxa.

  • 49(new). Seta 1-VII, length: (0) short; (1) moderately long; (2) long to very long. The length of seta 1-VII is compared to the length of segment X. Segment X length is the middorsal length measured along a straight line parallel to the longitudinal axis from the posterior margin of the saddle (minus marginal spicules) anteriorly to the first annulation in the membrane cephalad of the saddle. Fourth-instar larvae, not exuviae, should be used for measurements of segment X. Seta 1-VII is considered short (state 0) if it is less than or equal to 0.42 times the length of segment X, moderate (state 1) if it is 0.48–0.85 times the length of segment X and long to very long (state 2) if it is greater than or equal to 0.94 times the length of segment X.

  • 50(new). Seta 2-VII, position: (0) insertion near seta 1-VII; (1) insertion far anterior to seta 1-VII. Seta 2-VII is inserted near seta 1-VII (state 0) in most species examined (see Tanaka et al., 1979: figs 95, 102, Oc. (Fin.) japonicus and Oc. (Fin.) seoulensis). See Berlin (1969: figs 18, 44, Hw. walkeri and Hw. sexlineata; Belkin, 1962: figs 237, 251, Fl. bougainvillensis, Fl. hollingsheadi) for examples of seta 2-VII inserted far anterior to seta 1-VII (state 1). Seta 2-VII is normally stellate and located on the anterior half of segment VII in state (1).

  • 51(36). Seta 2-VII, development: (0) single; (1) branched. Seta 2-VII is normally single and short (state 0) in most Aedini (see Belkin, 1962: figs 218, 274, Ge. longiforceps and Le. geoskusea). This seta has two or more branches (state 1) in a few taxa, e.g. Downsiomyia, Finlaya (see Belkin, 1962: figs 237, 239, 241, Fl. bouganivillensis, Fl. burnetti and Fl. fijiensis), Haemagogus, Howardina and Scutomyia.

  • 52(new). Seta 3-VII, position: (0) insertion anterior to seta 1-VII; (1) insertion at same level or posterior to seta 1-VII. Seta 3-VII is inserted anterior to seta 1-VII (state 0) in most Aedini (see Belkin, 1962: figs 218, 243, Ge. longiforceps and Fl. franclemonti). This seta is inserted at the same level or posterior to seta 1-VII (state 1) in a few taxa, e.g. Kenknightia, Levua, Opifex, Tanakaius and Ha. australis (see Belkin, 1962: figs 208, 211, Op. (Opi.) fuscus and Ha. australis).

  • 53(new). Seta 3-VII, development: (0) single; (1) branched. See Zavortink (1972: figs 32, 34, 37, 40, Ko. purpureipes, Ab. papago, Az. ramirezi and Gy. mediovittata) for examples of state (0) and Reinert (1990: figs 24–32, Kenknightia) for examples of state (1).

  • 54(37, in part). Seta 3-VII, length: (0) short; (1) moderate; (2) long to very long. See character 49 for the method used to measure the length of this seta. The relatively long seta 3-VII in the outgroup species Ma. titillans is moderated by the very long dorsal length of segment X resulting in a moderate length (state 1), but this condition does not apply to any of the aedine species examined.

  • 55(new). Seta 10-VII, development: (0) single; (1) branched. Seta 10-VII is single (state 0) in many aedine taxa, e.g. Downsiomyia, Haemagogus (see figures in Arnell, 1973), Scutomyia and Tanakaius. This seta is branched (state 1) in other taxa, e.g. Finlaya, Opifex and Ha. australis (see Belkin, 1962: figs 208, 211).

  • 56(38). Seta 12-VII, position: (0) insertion anterior to seta 13-VII; (1) insertion at approximately same level as seta 13-VII; (2) insertion posterior to seta 13-VII. Seta 12-VII is inserted anterior to seta 13-VII (state 0) in a few taxa, e.g. Hg. (Haemagogus) (see figures in Arnell, 1973), most Howardina, Kompia, Scutomyia and St. africana. It is inserted at approximately the same level as seta 13-VII (state 1) in a few taxa, e.g. Abraedes, Gymnometopa and Hg. (Conopostegus) (see figures in Zavortink, 1972). Most aedine taxa have seta 12-VII posterior to seta 13-VII (state 2) (see Reinert, 1976b: figs 12, 13, Rhinoskusea).

  • 57(39). Seta 12-VII, development: (0) single; (1) branched. Seta 12-VIII is single (state 0) in many Aedini, e.g. Downsiomyia, Finlaya, Howardina (see figures in Berlin, 1969) and Stegomyia. It is branched (state 1) in other taxa, e.g. Aedes, Levua, Opifex and Zavortinkius (see Reinert, 1999b: fig. 14).

  • 58(40). Setae 1,2-VIII, insertion: (0) not attached to common setal support plate; (1) both attached to common setal support plate. Setae 1,2-VIII are not attached to a common setal support plate (state 0) in most aedine taxa. Both setae are attached to a common setal support plate (state 1) in Mucidus (see Belkin, 1962: figs 277, 279, Mu. alternans and Mu. painei).

  • 59(new). Seta 1-VIII, length: (0) short; (1) moderate; (2) long. Seta 1-VIII is considered short (state 0) if it is less than or equal to 0.38 times the length of seta 2-VIII, moderate (state 1) if it is 0.50–0.95 times the length of seta 2-VIII and long (state 2) if it is greater than or equal to 1.10 times the length of seta 2-VIII. See Belkin (1962) for examples of state (0) (fig. 211, Ha. australis), state (1) (fig. 234, Oc. (Fin.) argyronotum) and state (2) (fig. 231, Oc. (Fin.) albilabris).

  • 60(41). Seta 2-VIII, development: (0) single; (1) branched. Seta 2-VIII is single (state 0) in most Aedini. It is branched (state 1) in a few taxa, e.g. Opifex (see Belkin, 1962: fig. 208, Op. (Opi.) fuscus), Psorophora, Zavortinkius and Oc. (Bruceharrisonius)’.

  • 61(42). Seta 4-VIII, development: (0) single; (1) branched. Seta 4-VIII, like seta 2-VIII, is single (state 0) in most Aedini. This seta is branched (state 1) in many of the species that have seta 2-VIII also branched (see Belkin et al., 1970: figs 56, 60, Ps. (Pso.) ciliata and Ps. (Jan.) ferox).

  • 62(43). Comb: (0) few to several scales in one or two irregular rows; (1) numerous scales in a patch. The development of the comb is consistent for most generic-level taxa; however, in some large taxa it is normally consistent at the species-group level. See examples in Belkin (1962: fig. 316, Sc. albolineata) for state (0) and his fig. 243 (Fl. franclemonti) for state (1).

  • 63(44). Comb plate: (0) absent; (1) present. A comb plate is normally present in Psorophora (see Belkin et al., 1970: fig. 67, Ps. (Gra.) jamaicensis) and usually absent in most other Aedini. It occurs in Stegomyia (eight species), Sc. arboricola (most specimens), Do. dorseyi, Hg. (Hag.) capricornii, Hg. (Hag.) janthinomys, Oc. (Cha.) elchoensis and Oc. (Mac.) tremulus (Reinert et al., 2004).

  • 64(45). Siphon, acus: (0) absent; (1) present. An attached or detached acus located at the base of the siphon (in line with the pecten) is found in many aedine taxa, e.g. Downsiomyia, Finlaya, and species of Ochlerotatus and Oc. (Ochlerotatus) examined in this study (see figures of Ochlerotatus in Arnell & Nielsen, 1972; Arnell, 1976). This structure is absent in other taxa, e.g. Abraedes, Gymnometopa, Halaedes, Stegomyia, Zavortinkius, Oc. (Cha.) wattensis, Oc. (Mac.) tremulus and Oc. (Molpemyia) (see Reinert, 1993: fig. 8, Oc. (Mol.) pecuniosus).

  • 65(46). Pecten: (0) absent; (1) present, spines evenly spaced; (2) present, distal one or more spines more widely spaced. Most aedine taxa have a pecten with the spines evenly spaced (state 1) (see Belkin, 1962: figs 227–229, Oc. (Fin.) notoscriptus). A few taxa have a pecten with the distal one or more spines more widely spaced (state 2) (see Tanaka et al., 1979: figs 131, 132, Ae. esoensis). The pecten is absent (state 0) in the outgroup taxa Ma. titillans and Or. signifera.

  • 66(47). Seta 1-S, development: (0) one seta at base of siphon; (1) one seta some distance distally from base of siphon; (2) two or more setae some distance from base of siphon. Seta 1-S inserted ventroposteriorly some distance from the base of the siphon, and without accessory setae (state 1) (see Tanaka et al., 1979: fig. 101, Ta. savoryi), is the usual condition in Aedini. Aedes (see Tanaka et al., 1979: fig. 131, Ae. esoensis) and Oc. (Rusticoidus) possess accessory setae in addition to the non-basal seta 1-S (state 2). Reinert (1999a, 2000a) discussed the presence of accessory setae in these two taxa and noted their occurrence in Oc. (Och.) crinifer, Oc. (Och.) hexodontus, Oc. (Och.) hokkaidensis and Oc. (Och.) punctor (see Tanaka et al., 1979). Accessory setae on the siphon (state 2) are typical of genus Culex whereas seta 1-S at the base of the siphon (state 0) is characteristic of many species of Culiseta Felt.

  • 67(48). Seta 6-S, development: (0) absent; (1) short; (2) long. Setae 6-S and 9-S are absent (state 0) from the highly modified siphon of Ma. titillans (see Belkin et al., 1970: fig. 50). See illustrations in Belkin (1962: fig. 227, Oc. (Fin.) notoscriptus) for state (1) and his fig. 237 (Fl. bougainvillensis) for state (2).

  • 68(49). Seta 8-S, development: (0) short; (1) long. Seta 8-S is normally short (state 0) in Aedini. It is notably long (state 1) in genus Finlaya (see illustrations in Belkin, 1962; as Kochi Group), Oc. (Fin.) papuensis and the outgroup species Ma. titillans.

  • 69(new). Seta 9-S, development: (0) absent; (1) short, slender, nearly straight or slightly curved; (2) stouter, hooklike. Most Aedini have seta 9-S relatively short and nearly straight or gently curved (state 1) (see Belkin, 1962: fig. 234, Oc. (Fin.) argyronotum). Some taxa have seta 9-S short and hooklike (state 2) (see Belkin, 1962: fig. 274, Le. geoskusea). See explanation for Ma. titillans (state 0) under character 67.

  • 70(51). Sclerotization of segment X: (0) dorsal saddle incomplete ventrally; (1) saddle completely encircles segment X. All species of Aedini have either a small or large dorsal saddle or sclerotization that completely encircles segment X. A saddle completely encircling segment X is found in Oc. (Ochlerotatus) (some species groups), other Ochlerotatus (see illustrations in Arnell, 1976; Scapularis Group), Psorophora, the non-aedine outgroup species, and some other aedine outgroup species, e.g. Edwardsaedes imprimens and Huaedes wauensis.Reinert (2002a) provided illustrations of different developmental states of this character in Aedini.

  • 71(50). Saddle, acus: (0) absent; (1) present. Most species examined in this study do not have a saddle acus (state 0). However, within genus Ochlerotatus an acus is present (state 1) in Oc. communis (see Wood, Dang & Ellis, 1979: pl. 20), Oc. infirmatus and Oc. scapularis, Oc. (Rus.) refiki and Oc. (Rus.) rusticus. It is also present in Oc. (Fin.) albilabris, Oc. (Fin.) notoscriptus, Oc. (Psk.) bancroftianus and Oc. (Psk.) postspiraculosus. The acus is attached to the base of the saddle in these species except in Oc. (Fin.) albilabris and Oc. (Fin.) notoscriptus in which the acus is a small oval sclerite located in a similar position but detached from the base of the saddle. Orthopodomyia signifera has a narrow, basal, sclerotized band that is separated from the saddle. Because of its structure and location, it is not considered to be homologous with the acus in Aedini. An acus is present in the outgroup species Cs. inornata.

  • 72(new). Saddle, moderate to well-developed spicules on posterior margin: (0) absent; (1) present. See Belkin (1962: fig. 314, St. aegypti) for an example of the saddle without well-developed spicules on the posterior margin and his fig. 237 (Fl. bougainvillensis) for an example of well-developed spicules on this area.

  • 73(52). Seta 1-X, position: (0) inserted on saddle; (1) inserted ventral to saddle. See Belkin (1962: fig. 227, Oc. (Fin.) notoscriptus) for an example of state (0) and his fig. 211 (Ha. australis) for an example of state (1). Seta 1-X is inserted on the ventral or ventroposterior margin of the saddle in Ge. baisasi, Oc. (Mac.) tremulus and Oc. (Och.) muelleri, and is scored (0).

  • 74(53). Seta 3-X, development: (0) single; (1) branched. Seta 3-X is normally single in Aedini (see Zavortink, 1972: fig. 9, Oc. (Pro.) triseriatus) but it is branched in Az. ramirezi, Gy. mediovittata, Ko. purpureipes, Hw. sexlineata (see Zavortink, 1972: figs 32, 37, 40 for the first three species) and the outgroup species Cs. inornata and Ma. titillans (see Belkin et al., 1970: fig. 50).

  • 75(54). Sclerotization supporting seta 4-X (ventral bush): (0) absent; (1) grid with only transverse bars; (2) grid with both lateral and transverse bars; (3) boss. Reinert (2002a) conducted a comparative analysis of the ventral brush and its attachment to segment X in Aedini. Sclerotization supporting the ventral brush was divided into four types and 18 subtypes. The ancestral form of the ventral brush was hypothesized to have consisted of numerous, multiple-branched setae with short, stout stems attached to a well-developed grid consisting of both transverse and lateral grid bars. This brush also had numerous multiple-branched, precratal setae with short stems inserted on the ventral midline of a complete saddle. The derived condition was described as being a brush consisting of a few, simple, single setae on a poorly developed grid with only transverse grid bars. This brush lacked precratal setae, and the saddle was incomplete ventrally. Reinert further proposed that the condition in Christophersiomyia (as a subgenus of Aedes) with 6–8 moderately long, 2–7-branched setae with short stems inserted in an irregular row without both lateral and transverse grid bars could represent the most derived state because of the occurrence of a similar condition in most sabethine mosquitoes (see discussion in Harbach & Kitching, 1998). The ventral brush in sabethines, however, is normally represented by a single pair of setae (species of subgenus Sabethinus of Sabethes have an auxiliary pair of setae 4-X). The well-developed boss of Oc. (Mol.) pecuniosus (state 3) is shown in Reinert (1993: fig. 8) and that of Mu. laniger is illustrated in Tyson (1970: fig. 17).

  • 76(55). Precratal/preboss setae (i.e. two or more setae anterior to grid or boss): (0) absent; (1) present. Setae of the ventral brush are normally paired but an occasional specimen may possess an odd number of setae. Some species, e.g. Hg. (Hag.) splendens, have a single unpaired preboss seta whereas other species have a single unpaired precratal seta (these are scored as 0). Psorophora have numerous precratal setae that are inserted in the ventral margin of the saddle (see Belkin et al., 1970: figs 56, 60, 67).

  • 77(56). Seta 4a-X of ventral brush, length: (0) short; (1) long. Reinert (2002a), following Belkin (1962) and Knight & Laffoon (1971), defined seta 4-X as ‘Any seta of the ventral brush, the variable number of paired or unpaired setae arising from the ventral longitudinal midline of abdominal segment X. The most caudal (posterior) seta is designated as seta 4a-X, the next cephalad (anterior) seta as 4b-X, and so on. Seta 4a-X is normally long, rarely moderately long, and similar in length to seta 4c-X in Aedini. Abraedes, Aztecaedes, Kompia, Oc. (Och.) muelleri and Oc. (Fin.) biocellatus have a short seta 4a-X (for first four taxa see Zavortink, 1972: figs 32, 34, 37, 44).

  • 78(new). Seta 4a-X of ventral brush, development: (0) single to 3-branched (occasionally 4-branched); (1) ≥ 5-branched. See Belkin (1962: fig. 314, St. aegypti) for an example of state (0) and his fig. 316 (Sc. albolineata) for an example of state (1).

Pupae

  • 79(57). Cephalothorax, with clear unpigmented spots: (0) absent; (1) present. Clear unpigmented areas of the cephalothorax (scutum and metanotum) (state 1), and also the abdominal terga in some species, of pupal exuviae are characteristic of Finlaya (see illustrations in Belkin, 1962; as Kochi Group). Belkin (1962) reported that in living pupae of these mosquitoes, which breed almost entirely in water in plant axils, the unpigmented areas are iridescent blue and are very similar to those found in pupae of many New World sabethines and species of Cx. (Microculex), which also inhabit water held in leaf axils. Mattingly (1969) also reported that ‘Some South American sabethines and culicine pupae, occurring in bromeliad axils, are provided with luminescent integumentary patches which persist for some time after emergence of the adult’. Berlin's (1969) illustrations of the pupal metanotum of Hw. inaequalis and Hw. stenei show an unpigmented lateral spot. Because no specimens of these two species were available for examination, it is uncertain if these spots are similar to those reported above. Alanstonea brevitibia and Belkinius aurotaeniatus have similar clear unpigmented spots. A small clear spot dorsal to the trumpet base was observed in the two available specimens of Oc. (Fin.) loi, hence this character is scored (1) for this species.

  • 80(58). Trumpet, tracheoid area: (0) absent; (1) present at base, weakly developed; (2) present distal to base, well developed. The tracheoid area of the trumpet is well developed (state 2) in Finlaya, Mucidus, Rhinoskusea and the non-aedine outgroup species Cx. quinquefasciatus and Ma. titillans (see Belkin et al., 1970: figs 33, 51). The tracheoid area in most generic-level taxa of Aedini is weakly developed at the base of the trumpet (state 1), but it is absent (state 0) in outgroup taxa, e.g. Gymnometopa, Op. (Opi.) fuscus, Psorophora (most species), Oc. (Fin.) albotaeniatus, other aedine taxa, e.g. Alanstonea, Armigeres, Fredwardsius, Huaedes, Leptosomatomyia, Skusea, and Ae. (Adm.) alboscutellatus, as well as the non-aedine outgroup species Cs. inornata and Or. signifera.

  • 81(59). Seta 1-CT, development: (0) similar to seta 3-CT; (1) very strongly developed, considerably longer than seta 3-CT; (2) weakly developed, considerably shorter than seta 3-CT. With few exceptions, the length and development of seta 1-CT is constant within well-defined generic-level taxa. See illustrations in Belkin (1962) for examples of state (0) (fig. 233, Oc. (Fin.) argyronotum), state (1) (fig. 242, Fl. franclemonti) and state (2) (fig. 266, Oc. (Och.) edgari).

  • 82(60). Seta 5-CT, length: (0) ≤ 1.2 times length of seta 4-CT; (1) > 1.3 times length of seta 4-CT. Seta 5-CT is approximately equal to the length of seta 4-CT in many taxa. This seta is noticeably longer than seta 4-CT (state 1) in other taxa, e.g. Kenknightia (see Reinert, 1990: figs 15–23).

  • 83(61). Seta 7-CT, length: (0) ≤ length of seta 6-CT; (1) 1.2–5.0 times length of seta 6-CT; (2) ≥ 6.0 times length of seta 6-CT. The length of seta 7-CT is shorter than or equal to the length of seta 6-CT (state 0) in Stegomyia (see Belkin, 1962: fig. 313, St. aegypti; Huang, 1990: fig. 10, St. africana). For examples of state (1), see Belkin (1962: figs 207, 210, Op. (Opi.) fuscus and Ha. australis) and for state (2) see Zavortink (1972: figs 60, 62, Hg. (Con.) leucocelaenus and Hg. (Con.) clarki).

  • 84(62). Seta 11-CT, development: (0) single; (1) branched. See Belkin (1962: fig. 226, Oc. (Fin.) notoscriptus) for example of state (0) and his fig. 266 (‘Oc. (Och.) edgari) for state (1). A few species occasionally have seta 11-CT 2-forked distally and are scored (0).

  • 85(63). Seta 13-CT: (0) absent; (1) present. Among three paratypes of Ab. papago examined, a well-developed, 2-branched seta is present on both sides of one exuviae, an alveolus is present on both sides of another, and the seta and alveolus are absent on the third. Reinert (1980, 2000a) provided a review of the occurrence of seta 13-CT in Culicidae and Aedini.

  • 86(new). Seta 3-I, length: (0) < length of seta 6-I; (1) ≥ length of seta 6-I. The length of seta 3-I is shorter than the length of seta 6-I (state 0) in some taxa, e.g. Oc. scapularis (see Arnell, 1976: fig. 36). For examples of state (1), in which seta 3-I is longer than seta 6-I, see Schick (1970: figs 15, 31, Oc. (Pro.) terrens and Oc. (Pro.) homoeopus).

  • 87(new). Seta 3-I, development: (0) single, rarely split apically; (1) branched. Seta 3-I is often single (rarely with apex split) (state 0) in numerous taxa, e.g. Abraedes, Aedes, Aztecaedes, Downsiomyia (see Tanaka, 2002: fig. 8, Do. nipponica), Finlaya and Oc. (Protomacleaya) . See Belkin (1962: figs 210, 217, Ha. australis and Ge. longiforceps) for examples of state (1).

  • 88(64). Seta 6-I, length: (0) ≤ length of seta 7-I; (1) > length of seta 7-I. The length of seta 6-I is shorter than or equal to the length of seta 7-I (state 0) in some taxa, e.g. Op. (Opi.) fuscus and Ha. australis (see Belkin, 1962: figs 207, 210). Examples of taxa with seta 6-I longer than seta 7-I (state 1) are Ps. (Jan.) ferox (see Belkin et al., 1970: fig. 59) and Za. longipalpis (see Reinert, 1999b: fig. 11).

  • 89(65). Seta 2-II, position: (0) insertion mesal to or at same level as seta 1-II; (1) insertion lateral to seta 1-II. Seta 2-II is normally lateral to seta 1-II (state 1) in Aedini, however, it is mesal to or at the same level as seta 1-II (state 0) in Finlaya, Oc. (Fin.) chungi and Oc. (Mol.) pecuniosus (see Reinert, 1993: fig. 7).

  • 90(66). Seta 2-II, length: (0) < length of seta 1-II; (1) ≥ length of seta 1-II. Seta 2-II is normally shorter than seta 1-II in Aedini, however, it is as long or longer and often stouter than seta 1-II (state 1) in some taxa, e.g. Rhinoskusea, Scutomyia and Ha. australis.

  • 91(67). Seta 3-II, position: (0) insertion mesal to or at same level as seta 2-II; (1) insertion lateral to seta 2-II. The position of seta 3-II in relation to seta 2-II is constant for most generic-level taxa, with few exceptions. However, in Oc. (Finlaya) and Oc. (Protomacleaya) this character appears to be constant at the species-group level. Examples of state (0) are found in Aztecaedes and Kompia (see Zavortink, 1972: figs 31, 36), and state (1) in Abraedes and Zavortinkius (see Zavortink, 1972: fig. 33, Ab. papago; Reinert, 1999b: fig. 11, Za. longipalpis).

  • 92(68). Seta 3-II, length: (0) ≤ length of seta 6-II; (1) > length of seta 6-II. State (0) is found in a number of aedine taxa, e.g. Aedes, Geoskusea, Isoaedes and Levua. Numerous taxa have seta 3-II longer than seta 6-II (state 1), e.g. Abraedes, Aztecaedes, Gymnometopa and Kenknightia. Seta 3-II is normally slender when it is short to moderately long and stout when it is long.

  • 93(70). Seta 5-II, position: (0) insertion lateral to or at same level as seta 4-II; (1) insertion mesal to seta 4-II. Seta 5-II is longer and normally lateral to seta 4-II (state 0) (see Belkin, 1962: fig. 233, Oc. (Fin.) argyronotum) in most Aedini. See Belkin (1962: fig. 207, Op. (Opi.) fuscus) for an example of state (1). This seta is normally lateral to seta 4-II, rarely positioned mesally, in Oc. (Pro.) hendersoni.

  • 94(new). Seta 5-II, length: (0) ≤ length of seta 3-II; (1) > length of seta 3-II. Most taxa examined have seta 5-II shorter than or equal to the length of seta 3-II (state 0), e.g. Haemagogus (see Arnell, 1973: figs 34, 39). Examples of taxa with seta 5-II longer than seta 3-II are Oc. (Fin.) albilabris and usually Oc. (Fin.) notoscriptus (see Belkin, 1962: figs 226, 230).

  • 95(71). Seta 6-II, length: (0) ≤ length of seta 7-II; (1) > length of seta 7-II. Seta 6-II is normally longer than seta 7-II (state 1) in Aedini (see Belkin, 1962: fig. 230, Oc. (Fin.) albilabris), however, it is shorter (state 0) (see Belkin, 1962: fig. 207, Op. (Opi.) fuscus) in some generic-level taxa and species groups (see Appendix 2).

  • 96(69). Seta 3-III, development: (0) single; (1) branched. Most taxa examined have seta 3-III single, long and stout (state 0) (see Belkin, 1962: fig. 230, Oc. (Fin.) albilabris). Seta 3-III is branched and normally slender (state 1) in several taxa, e.g. Isoaedes, Kenknightia, Opifex and Rhinoskusea (see Reinert, 1976b: figs 9–11, for Rhinoskusea).

  • 97(72). Seta 6-III, development: (0) single; (1) branched. The development of seta 6-III is constant for generic-level taxa, with few exceptions. However, in some specimens of some species, e.g. Ke. harbachi, Oc. (Fin.) banksi and Oc. (Pro.) berlini, this seta usually exhibits one state but occasionally exhibits the other state, e.g. normally single but sometimes 2-branched, these are scored (0,1).

  • 98(74). Seta 5-V, length: (0) < medial length of tergum VI; (1) ≥ medial length of tergum VI. Seta 5-V is often single (rarely 2-branched), very long, stout and longer than the tergum of the following segment, but in some taxa it is branched, short to moderately long, slender and shorter than tergum VI. See Belkin (1962) for a comparison of the two character states (fig. 313, St. aegypti, state 0; fig. 207, Op. (Opi.) fuscus, state 1).

  • 99(75). Seta 2-VI, position: (0) insertion mesal to or at same level as seta 1-VI; (1) insertion lateral to seta 1-VI. The position of seta 2-VI, in relation to seta 1-VI, is normally consistent within most well-defined generic-level taxa. See examples in Belkin (1962: fig. 233, Oc. (Fin.) argyronotum, state 0; fig. 210, Ha. australis, state 1).

  • 100(76). Seta 3-VI, position: (0) insertion mesal to or at same level as seta 1-VI; (1) insertion lateral to seta 1-VI. Seta 3-VI is normally lateral to seta 1-VI (state 1) in Aedini. However, it is positioned at the same level or mesal to seta 1-VI (state 0) in Finlaya, Downsiomyia, Oc. (Cha.) wattensis, Oc. (Fin.) albilabris, Oc. (Fin.) argenteitarsis, Oc. (Fin.) assamensis, Oc. (Fin.) loi, Oc. (Fin.) subalbitarsis, Oc. (Mac.) tremulus, Op. (Opi.) fuscus (see Belkin, 1962: fig. 207) and the non-aedine outgroup species Cs. inornata and Cx. quinquefasciatus.

  • 101(73). Seta 6-VII, position: (0) insertion anterior to seta 9-VII; (1) insertion posterior to or at some level as seta 9-VII. Seta 6-VII is inserted posterior (occasionally at the same level) to seta 9-VII (state 1) in most Aedini, e.g. Oc. (Protomacleaya) (see illustrations in Schick, 1970 and Zavortink, 1972). Seta 6-II is anterior to seta 9-VII (state 0) in some species examined, e.g. Psorophora (see Belkin et al., 1970: figs 57, 59, 66), Oc. (Bruceharrisonius)’, Oc. (Fin.) gilli, Oc. (Fin.) loi and Za. fulgens.

  • 102(77). Seta 9-VII, length: (0) ≤ length of seta 6-VII; (1) > length of seta 6-VII. Seta 9-VII is normally longer and often stouter than seta 6-VII (state 1) in Aedini, but it is occasionally shorter than or equal to the length of seta 6-VII (state 0). Seta 6-VII does not occur in pupae of Ma. titillans, nor in Ma. dyari and Ma. flaveola, therefore this character could not be scored for this species and is entered as missing (–) in the data matrix.

  • 103(78). Seta 9-VIII, development: (0) single or 2-branched; (1) ≥ 3 branches. Many aedine taxa have seta 9-VIII multiple-branched, often aciculate and often strongly developed (see Belkin, 1962: fig. 233, Oc. (Fin.) argyronotum). See Reinert (1993: fig. 7, Oc. (Mol.) pecuniosus) for example of state (0).

  • 104(79). Paddle, midrib: (0) weakly developed, not reaching apex of paddle; (1) well developed, extending to or near apex of paddle. Most generic-level taxa of Aedini have the midrib well developed and extending to or near the apex of the paddle (state 1). The midrib, however, is weakly developed to nearly indistinct in some taxa, e.g. Udaya and Zeugnomyia. The midrib is shortened and often restricted to about the basal 0.60 of the paddle (state 0) in other taxa, e.g. Abraedes, Aztecaedes, Gymnometopa, Kompia, as well as Oc.deserticola, Oc.muelleri, Oc.sierrensis and Oc.varipalpus (see Arnell & Nielsen, 1972: fig. 8) of subgenus Ochlerotatus, these are scored (0).

  • 105(80). Paddle, fringe of hairlike spicules: (0) absent; (1) present. Reinert (2000a) listed the generic-level taxa of Aedini with a fringe of hairlike spicules on the pupal paddle. These are present or absent in generic-level taxa, with few exceptions. Most species of Stegomyia have well-developed hairlike spicules on the paddle margin, but there are notable exceptions, e.g. St. aegypti and the Dendrophilus Group (see illustrations in Huang, 1997).

  • 106(81, in part). Paddle, apical margin: (0) sharply or broadly rounded or flat; (1) with moderate to deep emargination at or near apex of midrib. Most Aedini have the apical margin of the paddle rounded, but some have this area flattened or with a slight emargination, e.g. Oc. (Och.) deserticola and Oc. (Och.) varipalpus, and are scored (0). A few taxa have a deeper emargination on the apical margin (state 1), e.g. Abraedes, Aztecaedes, Kompia, Fl. kochi, Ha. australis, Hw. walkeri, Oc. (Och.) atropalpus and Oc. (Och.) epactius (see Zavortink, 1972: fig. 46).

  • 107(new). Seta 1-Pa, length: (0) short to moderately long, ≤ 0.33 length of paddle; (1) long, ≥ 0.40 length of paddle. Downsiomyia and Finlaya are examples of state (0) (see illustrations of Finlaya, as Kochi Group, in Belkin, 1962). Levua geoskusea and Oc. (Brh.) greenii are examples of state (1) (see Reinert, 2003: fig. 2E).

  • 108(82, in part). Seta 1-Pa, development: (0) absent; (1) present, single, rarely 2-branched or 2-forked; (2) present, ≥ 3 branches. Seta 1-Pa is absent (state 0) in Ma. titillans (see Belkin et al., 1970: fig. 51). See Reinert (1990: figs 15–23, Kenknightia) for state (1) and Reinert (1976b: figs 9–11, Rhinoskusea) for state (2). Some species show an overlap between the two character states, e.g. Do. leonis (2 or 3 branches), Ps. (Jan.) ferox (2 or 3 branches) and Cs. inornata (2 or 3 branches), and are scored (1,2). However, most specimens of these species have three or more branches.

  • 109(83). Seta 2-Pa: (0) absent; (1) present. Absence of seta 2-Pa (state 0) is the usual condition in Aedini, however, seta 2-Pa is present (state 1) in subgenera Grabhamia (except Ps. infinis) and Psorophora of Psorophora (see Belkin et al., 1970: figs 57, 64–66), Ta. savoryi (rarely absent on one side of a few specimens) (see Tanaka, 2002: fig. 3) and the outgroup species Cx. quinquefasciatus. One of six pupal exuviae of Oc. (Och.) fluviatilis examined has the apex of one paddle with two setae and the other with two setae and a setal alveolus, which was considered an anomaly.

Adults (females except where otherwise noted)

  • 110(84). Erect scales of head: (0) absent; (1) restricted to occiput; (2) on occiput and vertex. See Harbach & Kitching (1998) for examples of state (1) (their fig. 6B) and state (2) (their fig. 6A). Erect scales in Aedini are either restricted to the occiput or borne on both the occiput and vertex. In some species, several semierect, narrow, forked scales similar to erect scales on the occiput occur on the vertex (often posterior to the ocular setae), and these are scored (2). Erect scales are absent in Op. (Opi.) fuscus, however, numerous short, curved, dark setae occur on the occiput and vertex.

  • 111(85). Decumbent scales of vertex: (0) broad; (1) narrow; (2) both broad and narrow. Harbach & Knight (1980) illustrated examples of scales that are narrow and broad. In species with both broad and narrow scales on the vertex (state 2), the narrow scales are usually on the median area and may be restricted to a double row along the coronal suture (see Tanaka et al., 1979: fig. 228, Oc. (Fin.) oreophilus). In Scutomyia, a patch of narrow scales occurs on the anterior median area (state 2). Examples of state (0) are shown in Tanaka et al. (1979: fig. 222, Oc. (Fin.) seoulensis) and state (1) in their fig. 221 (Ta. savoryi).

  • 112(86). Ocular line, width: (0) narrow; (1) broad. The ocular line is interpreted here as the area anterior to the ocular setae and posterior to the dorsal margin of the eye. Examples of a narrow ocular line (state 0) are found in Tanaka et al. (1979: figs 220, 222, Ta. togoi and Oc. (Fin.) seoulensis) and a broad one (state 1) in Reinert (1990: fig. 3, Ke. harbachi). The ocular line of Cs. inornata is moderate in width, and is here scored as (0). See discussion of scale types under character 111.

  • 113(87, in part). Ocular scales: (0) narrow; (1) broad; (2) both narrow and broad. Most species have either narrow or broad scales on the ocular line but a few have both narrow and broad scales (state 2), e.g. Howardina, Mu. laniger and Oc. communis. See discussion of scale types under character 111.

  • 114(88). Eyes, immediately above antennal pedicels: (0) contiguous; (1) narrowly to moderately separated; (2) broadly to very broadly separated. The distance separating the eyes is determined on the area immediately above the antennal pedicels and is measured in number of eye facets (diameter). Species with eyes that touch or are separated by less than or equal to one eye facet are considered contiguous and are scored (0), e.g. Oc. (Fin.) assamensis, those separated by 2–4 (rarely 5) facets are scored (1), e.g. Oc. (Fin.) geniculatus, and those separated by 6 or more facets are scored (2), e.g. Op. (Opi.) fuscus. Scales protruding from the median, anterior area of the vertex or upper part of the interocular space may obscure this area, and in some cases, e.g. Howardina, the scales must be removed to determine the distance between the eyes. Gutsevich (1974a, b, 1975a, b) used the diameter of eye facets compared to the distance between the eyes as a means of distinguishing several genera and subgenera, however, he used a different criterion than the number of facets used here.

  • 115(90). Interocular space, scales: (0) absent; (1) narrow; (2) broad; (3) both narrow and broad. In Geoskusea, Levua and Rhinoskusea, the interocular space is reduced to a very small triangle and is apparently without scales (state 0). In these subgenera, the eyes are contiguous for nearly the entire mesal length, resulting in a very small interocular space. Most taxa examined in the present study have narrow scales on this area (state 1). Broad scales (state 2) on the interocular space occur in Abraedes, Downsiomyia, Gymnometopa, Haemagogus, Kenknightia, Kompia, Stegomyia, Zavortinkius, Ps. (Jan.) ferox and five species of Oc. (Finlaya)’. Opifex (Opi.) fuscus has only a few scattered small broad scales on this area. Both narrow and broad scales (state 3) occur on the interocular space of Oc. (Cha.) wattensis.

  • 116(89). Interocular space, setae: (0) absent; (1) present. The interocular space is defined here as the area between the compound eyes extending dorsad from the postfrontal suture to a point level with the dorsal margins of the eyes. Setae are normally present on this area in Aedini and the outgroup species. They are absent in Gymnometopa and Kompia. They are also absent in the aedine genera Ayurakitia and Udaya.

  • 117(91). Antennal pedicel, mesal surface: (0) with few to numerous scattered scales (not overlapping to only slightly overlapping and not silvery) and/or setae; (1) with patch of broad, overlapping, silvery scales; (2) bare. State (0) occurs in most Aedini and the outgroup species. State (1) occurs in Abraedes, Aztecaedes, Gymnometopa and Kompia, Oc. (Fin.) albilabris and Oc. (Fin.) notoscriptus, as well as Albuginosus, Bothaella, Scutomyia, Stegomyia and Zeugnomyia. In state (0), the mesal surface of the pedicel has few to numerous, small to large, scattered to slightly overlapping, pale (white, cream-coloured, golden brown, etc.) or dark scales and/or setae (see Tanaka et al., 1979: fig. 222, Oc. (Fin.) seoulensis), whereas in state (1) a moderately large to large patch of tightly overlapping, silvery scales is present (see Tanaka et al., 1979: fig. 234, St. aegypti). The mesal surface of Oc. (Och.) deserticola has a moderate-sized patch of broad, overlapping, white scales above and a few scattered, small, white scales below, and is scored (0). State (2) occurs in two Afrotropical species examined, Oc. (Fin.) luteostriatus and Oc. (Fin.) pulchrithorax.

  • 118(92). Antennal pedicel, lateral surface: (0) bare; (1) with scales. The outer surface of the antennal pedicel is bare (state 0) (see Tanaka et al., 1979: fig. 228, Oc. (Fin.) oreophilus) in most Aedini. Scales are present on this area (state 1) in Oc. (Rusticoidus), Ochlerotatus (some species) (see Tanaka et al., 1979: fig. 212, Oc. dorsalis) and Oc. (Ochlerotatus) (some species), Stegomyia and the outgroup species Ma. titillans.

  • 119(93). Apical flagellomeres (males): (0) both apical flagellomeres disproportionately long compared with other flagellomeres; (1) these flagellomeres not disproportionately long in comparison with others. Antennae with the two distal flagellomeres approximately equal in length and much longer than the proximal flagellomeres is the usual condition in Aedini (state 0) (see Tanaka et al., 1979: fig. 228, Oc. (Fin.) oreophilus). However, the two apical flagellomeres are approximately the same length as the other flagellomeres in both subgenera of Opifex (see Belkin, 1968: figs 2, 19).

  • 120(94). Antenna, development of flagellar whorls (males): (0) few short setae, dispersed more or less around flagellomeres; (1) several moderately long to long setae, directed more or less dorsally and ventrally; (2) numerous long setae, normally directed dorsally and ventrally. Opifex is notable in having only a few short setae on each flagellomere (state 0) (see Belkin, 1968: figs 2, 19, Op. (Opi.) fuscus and Op. (Not.) chathamicus). Levua, Rhinoskusea and Ge. longiforceps have several setae that are moderately long to long and normally directed dorsally and ventrally (state 1) (see Reinert, 1976b: fig. 2, Rh. longirostris). Antennal whorls comprising numerous long setae normally directed dorsally and ventrally (state 2) (see Tanaka et al., 1979: fig. 222, Oc. (Fin.) seoulensis) are usually present in Aedini.

  • 121(new). Maxillary palpus, pale scales: (0) absent; (1) present. Maxillary palpi may be entirely dark-scaled (state 0) or have pale scales intermixed with or forming bands or patches in the dark-scaled areas (state 1).

  • 122(95, in part). Maxillary palpomeres (males): (0) five, palpomeres 2 and 3 fused/ankylosed; (1) three, fourth absent or vestigial. Palpomeres 2 and 3 exhibiting state (0) have a paler, somewhat distinct ring where they are fused/ankylosed whereas no such area is apparent when palpomeres exhibit state (1). Very short palpi are found in males of the following Aedini: Aedes, Ae. (Cancraedes)’, Belkinius, Bothaella, Christophersiomyia, Geoskusea (most species), Haemagogus (few species), Heizmannia, Huaedes, Levua, Leptosomatomyia, Opifex (Nothoskusea), Paraedes, Rhinoskusea, Verrallina and Zeugnomyia (Reinert, 2000a). There is no way of knowing whether palpomere 2 of males with short palpi actually consists of palpomeres 2 and 3 that are completely and unrecognizably fused. The number of palpomeres is consistent for generic-level taxa, with few exceptions, e.g. Ge. baisasi and Ge. longiforceps.

  • 123(new). Maxillary palpus, length (males): (0) short, ≤ 0.25 length of proboscis; (1) moderate, 0.48–0.80 length of proboscis; (2) long, ≥ 0.84 length of proboscis. The maxillary palpus is noticeably short in males of some aedine taxa (state 0), e.g. Levua and Rhinoskusea (see Reinert, 1976b: fig. 2, Rh. longirostris), moderately long (state 1) in some, e.g. Scutomyia albolineata and Sc. arboricola (see Huang, 1979: fig. 6) and long, often as long as or longer than the proboscis (state 2) in others (see Tanaka et al., 1979: figs 212, 213, 222, Oc. dorsalis, Oc. (Och.) excrucians and Oc. (Fin.) seoulensis).

  • 124(new). Maxillary palpus, long lateroventral setae on distal part of palpomere 3 and entire length of palpomere 4 (males): (0) absent or few; (1) numerous. Many aedines have numerous long setae on the lateroventral or ventral margins distally on palpomere 3 and the entire length of palpomere 4 (state 1) (see Tanaka et al., 1979: figs 213, 215, Oc. (Och.) excrucians and Oc. (Och.) sticticus), whereas other species have no or few setae on the distal part of palpomere 3 and few to several on palpomere 4 (state 0) (see Tanaka et al., 1979: figs 220, 221, Ta. togoi and Ta. savoryi). Species with numerous long setae on the distal part of palpomere 3 often also have the distal part of the palpomere somewhat swollen.

  • 125(new). Proboscis, pale scales: (0) absent; (1) present. The proboscis may be entirely dark-scaled (state 0) (see Russell, 1996: 81, Oc. (Fin.) rubrithorax) or have pale scales intermixed with or forming bands or patches in the dark-scaled areas (state 1) (see Russell, 1996: 61, 73, Fl. kochi and Oc. (Fin.) notoscriptus).

  • 126(96). Antepronota: (0) approximated; (1) widely separated. The antepronota are nearly touching dorsally (state 0) in Hg. (Hag.) equinus and Hg. (Hag.) splendens, and in some other taxa studied previously (Reinert et al., 2004), whereas they are widely separated (state 1) in most Aedini.

  • 127(97, in part). Anterior acrostichal setae: (0) absent; (1) present. Anterior acrostichal setae are defined as those on the acrostichal area cephalad of the scutal angle whereas those caudad of this point are the posterior acrostichal setae. Setae on the anterior promontory (the broad median area of the mesonotum at the anterior end of the acrostichal area that projects more or less cephalad over the cervix) should not be confused with the presence or absence of setae on the anterior end of the acrostichal area. Arnell (1973: 8) reported the presence of minute dorsocentral and acrostichal setae (visible only in slide preparations) in Haemagogus. These setae were not seen in pinned specimens of Hg. equinus and Hg. splendens examined, and are scored as (0).

  • 128(97, in part). Posterior acrostichal setae: (0) absent; (1) present. See discussion under character 127.

  • 129(98, in part). Anterior dorsocentral setae: (0) absent; (1) present. The dorsocentral area is defined as the longitudinal area on either side of the acrostichal area extending caudally from the antedorsocentral area (see Reinert, 1999b) to the prescutellar area. Setae on the area cephalad of the juncture of the prescutal suture are referred to as ‘anterior dorsocentral setae’ and those caudad as ‘posterior dorsocentral setae.’ Also see discussion under character 127.

  • 130(98, in part). Posterior dorsocentral setae: (0) absent; (1) present. See discussion under character 129.

  • 131(new). Acrostichal and dorsocentral areas, anterior 0.30–0.70 covered with pale scales: (0) absent; (1) present. Females of some species have the combined acrostichal and dorsocentral areas dark-scaled (state 0), e.g. Ge. baisasi (see Tanaka et al., 1979: fig. 239) and Ke. dissimilis (see Reinert, 1990: fig. 1) whereas other species have this area, and often the entire anterior 0.30–0.70 (acrostichal, dorsocentral and scutal fossal areas) of the scutum, covered with pale (normally white or silvery) scales (state 1) (see Barraud, 1923: figs 13, 17–19, Oc. (Fin.) deccanus, Oc. (Fin.) cacharanus, Oc. (Fin.) lophoventralis and Oc. (Fin.) cogilli).

  • 132(new). Anterior acrostichal area, with narrow, pale-scaled stripe: (0) absent; (1) present. When present (state 1), the pale-scaled stripe normally consists of narrow scales, but these are broad in some species, e.g. Oc. (Mol.) pecuniosus (see Reinert, 1993: fig. 3), Za. fulgens and Za. longipalpis (see Reinert, 1999b: fig. 1). The pale-scaled stripe is normally narrow as in Oc. (Fin.) oreophilus but may be moderate in width and slightly separated by the acrostichal setae as in Ta. togoi (see Tanaka et al., 1979: figs 228, 220, respectively). In St. aegypti, a narrow white-scaled stripe occurs on either side of the scutum on the outer edge of the acrostichal area at the interface with the dorsocentral area (see Tanaka et al., 1979: fig. 234). Belkin et al. (1970) referred to this stripe as the ‘outer acrostichal line’. It is included here as an acrostichal stripe (state 1). In female paratypes of Oc. (Fin.) candidoscutellum, the acrostichal, dorsocentral, scutal fossal, prescutellar and supraalar areas have indistinct, narrow, stripes of whitish scales on a background of golden-brown scales. These are scored as present (1). See discussion under character 127.

  • 133(new). Posterior acrostichal area, with narrow, pale-scaled stripe: (0) absent; (1) present. See discussion under character 127.

  • 134(new). Anterior dorsocentral area, with narrow, pale-scaled stripe: (0) absent; (1) present. A stripe normally extends over the central part of the dorsocentral area (see Tanaka et al., 1979: fig. 217, Oc. (Fin.) japonicus), however, it may be on the ‘inner dorsocentral area (see Berlin, 1969: figs 8, 9) as in Ab. papago and Hw. walkeri. See discussion of character 132 for Oc. (Fin.) candidoscutellum. Also see discussion under character 129.

  • 135(new). Posterior dorsocentral area, with narrow, pale-scaled stripe: (0) absent; (1) present. See discussion under character 129.

  • 136(new). Scutal fossa, scales: (0) dark; (1) contrasting pale scales in large patch; (2) contrasting pale-scaled lines or small patches on lateral and/or mesal and/or posterior margins. The scutal fossa may be entirely dark-scaled (state 0) as in Ge. baisasi (see Tanaka et al., 1979: fig. 239), have pale scales in a large patch or be completely pale-scaled (state 1) as in Oc. (Fin.) seoulensis and Do. nishikawai (see Tanaka et al., 1979: figs 222, 227) or bear small pale patches or narrow pale-scaled lines on the lateral and/or mesal and/or posterior margins (state 2) (see Tanaka et al., 1979: fig. 221, Ta. savoryi). Pale-scaled areas in states (1) and (2) contrast (normally white or silvery) with background scales that are normally dark but may be a darker shade or colour of pale scales in some species, e.g. Oc. (Fin.) pulchriventer and Oc. (Fin.) tubbutiensis. The scutal fossa of Oc. communis normally has a pale-scaled margin with contrasting darker scales on the median area (see Carpenter & LaCasse, 1955: pl. 54), however, some specimens have the scutal fossa nearly covered with pale scales and only a few darker scales medially and is therefore scored (1,2). Some species, e.g. Ps. (Jan.) ferox (see Carpenter & LaCasse, 1955: pl. 35), have the scutal fossa covered with intermixed broad pale and dark scales in no defined pattern, scored here as state (2).

  • 137(99). Prescutellar area, median and/or posterior parts: (0) bare; (1) with scales and/or setae. The median part of the prescutellar area is bare (state 0) in most aedine species. This area in Oc. (Fin.) jugraensis (see Knight, 1968: fig. 1), Oc. (Fin.) rizali and Oc. (Fin.) sherki has a median, longitudinal, narrow, pale-scaled stripe, and the scales mesad of the prescutellar setae are dark. Some other taxa have state (1), e.g. Aztecaedes, Hg. (Haemagogus), Mucidus, Oc. (Rusticoidus), Oc. (Ochlerotatus) (few species), Psorophora (some), Rhinoskusea, Hw. walkeri and Za. longipalpis. Finlaya have scales on the anterior area and/or in transverse bands on the posterior area.

  • 138(100). Prescutellar setae: (0) absent; (1) present. Prescutellar setae are normally present in Aedini, but are absent in Hg. (Haemagogus) as well as in Alanstonea brevitibia, Pseudarmigeres argenteoventralis dunni, Hz. (Heizmannia) and Hz. (Mattinglyia) achaetae (sometimes present in the last species).

  • 139(new). Prescutellar area, pale scales on outer margin mesal to setae: (0) absent; (1) present. Many aedine species have a narrow to moderately wide stripe of pale scales mesal to the prescutellar setae (state 1) (see Tanaka et al., 1979: figs 217, 219, 220, Oc. (Fin.) japonicus, Oc. (Fin.) hatorii and Ta. togoi) but other species have dark scales on this area (state 0) (see Tanaka et al., 1979: fig. 229, Oc. (Fin.) watasei).

  • 140(new). Antealar area, anterior part, scales: (0) absent; (1) present, dark; (2) present, pale. This character refers to the part of the antealar area anterior to the forward edge of the paratergite. This area may lack scales (state 0) (see Zavortink, 1972: fig. 35, Az. ramirezi) or bear dark scales (state 1) (see Tanaka et al., 1979: fig. 239, Ge. baisasi) or pale scales (state 2) (see Tanaka et al., 1979: fig. 222, Oc. (Fin.) seoulensis).

  • 141(new). Supraalar area, scales: (0) all dark; (1) with longitudinal patch or stripe of pale scales; (2) with transverse patch of pale scales. The supraalar area may be entirely dark-scaled (state 0) (see Tanaka et al., 1979: fig. 227, Do. nishikawai), bear a longitudinal pale-scaled patch (state 1) (see Tanaka et al., 1979: fig. 217, Oc. (Fin.) japonicus) or have a transverse pale-scaled patch (state 2) (see Tanaka et al., 1979: fig. 229, Oc. (Fin.) watasei; Barraud, 1923: figs 11–13, 17, Oc. (Fin.) gubernatoris, Oc. (Fin.) deccanus and Oc. (Fin.) cacharanus).

  • 142(101). Scutellum, scales on midlobe: (0) narrow; (1) broad; (2) both narrow and broad. Scales on the midlobe (as well as lateral lobes) of the scutellum may be all narrow (state 0) (see Tanaka et al., 1979: fig. 219, Oc. (Fin.) hatorii), all broad (state 1) (see Tanaka et al., 1979: fig. 227, Do. nishikawai) or both narrow and broad (state 2) (see Knight, 1968: fig. 1, Oc. (Fin.) jugraensis). Mucidus alternans and Mu. langer have the lobes bearing several narrow and a few moderately broad, decumbent, falcate scales and numerous long, twisted, erect scales (scored 0). Long, twisted, erect scales are also present on other body areas, e.g. head, thorax, abdomen, etc. (see illustrations in Tyson, 1970). See discussion of scale types under character 111.

  • 143(102). Scutellum, scales on lateral lobes: (0) narrow; (1) broad; (2) both narrow and broad. See discussion under characters 111 and 142.

  • 144(103). Paratergal scales: (0) absent; (1) present. Generic-level taxa normally either have paratergal scales or lack them, with few exceptions. The paratergite may be bare (state 0) as in Oc. (Fin.) japonicus japonicus and Ge. baisasi (see Tanaka et al., 1979: figs 217, 239) or have several to numerous scales (state 1) as in Oc. (Fin.) seoulensis and St. aegypti (see Tanaka et al., 1979: figs 222, 234).

  • 145(104). Parascutellar scales: (0) absent; (1) present. Parascutellar scales in Aedini were first noted in species of Oc. (Rusticoidus) (Reinert, 1999a) and a few species of Oc. (Ochlerotatus)’.

  • 146(105). Antepronotal scales: (0) absent; (1) present. Scales are present on the antepronotum (state 1) (see Arnell, 1973: fig. 38, Hg. (Hag.) splendens) in the majority of species examined in the present study and may be narrow, broad or both narrow and broad. Scales are absent (state 0) in Geoskusea (see Tanaka et al., 1979: fig. 239, Ge. baisasi), Levua, Rhinoskusea, Oc. (Fin.) candidoscutellum and Ps. (Pso.) ciliata.

  • 147(106). Postpronotal scales: (0) absent; (1) present. Broad, narrow or both broad and narrow scales are normally present on the postpronotum. They are absent from this structure in Rhinoskusea (see Reinert, 1976b: fig. 2, Rh. longirostris), Hw. walkeri, Oc. (Fin.) candidoscutellum and Ps. (Pso.) ciliata.

  • 148(107). Prespiracular setae: (0) absent; (1) present. Setae are normally absent from the prespiracular area in aedines, however they are present in Psorophora and the outgroup species Cs. inornata.

  • 149(108). Postspiracular setae: (0) absent; (1) present. Postspiracular setae are present in most Aedini but are absent in some taxa (see Reinert et al., 2004). They are absent in Ko. purpureipes and the outgroup species Cs. inornata, Cx. quinquefasciatus and Or. signifera.

  • 150(109). Postspiracular scales: (0) absent; (1) present. Presence (state 1) (see Tanaka et al., 1979: fig. 220, Ta. togoi) or absence (state 0) (see Arnell, 1976: fig. 35, Oc. scapularis) of scales on the postspiracular area is normally constant for generic- and species-group taxa in Aedini.

  • 151(110). Hypostigmal scales: (0) absent; (1) present. Scales on the hypostigmal area should not be confused with those on the dorsal part of the subspiracular area when two patches are present on the latter structure, one immediately dorsal to the other. Scales are normally absent on the hypostigmal area of Aedini (state 0) (see Zavortink, 1972: fig. 5, Oc. (Pro.) brelandi). A small patch of scales is present in some species (state 1) (see Zavortink, 1972: fig. 35, Az. ramirezi).

  • 152(111). Subspiracular area: (0) bare; (1) with scales and/or setae. Presence or absence of scales on the subspiracular area is normally constant for genera and species groups of Aedini, with few exceptions. Scales and setae are present on this area, e.g. Oc. (Och.) varipalpus (see Arnell & Nielsen, 1972: fig. 7) and Ps. (Pso.) ciliata. Also see discussion under character 151.

  • 153(112). Upper proepisternal setae: (0) 1–4; (1) 5–19; (2) ≥ 20. Some specimens of a few species examined have a slight overlap (usually of one seta) of the states, e.g. Oc. (Fin.) assamensis, Oc. (Fin.) chrysolineatus, Ta. savoryi, Ge. longiforceps, Ia. cavaticus and Ko. purpureipes, whereas most specimens of these species have setal numbers of a single character state.

  • 154(113). Upper proepisternum, scales: (0) absent; (1) present. Aedine species normally have scales on the upper proepisternum (state 1) (see Tanaka et al., 1979: fig. 218, Oc. (Fin.) japonicus). These scales are absent in Geoskusea, Levua and Rhinoskusea (see Reinert, 1976b: fig. 2, Rh. longirostris).

  • 155(114). Lower proepisternum, scales: (0) absent; (1) present. When scales are present on the lower proepisternum they usually cover all or much of the surface and are normally broad and silvery or white. Scales are present in the following taxa of the present study: Abraedes, Haemagogus, Kompia, Scutomyia, Stegomyia, Zavortinkius, Oc. communis, Oc. infirmatus, Oc. scapularis, Oc. (Och.) muelleri, Oc. (Pro.) burgeri, Oc. (Pro.) terrens, Oc. (Rusticoidus) (2 species) and Ps. (Jan.) ferox.

  • 156(115). Upper mesokatepisternal setae: (0) absent; (1) present. One or more upper mesokatepisternal setae are normally present in Aedini. Reinert et al. (2004) noted taxa without these setae. These setae are absent in Hg. (Haemagogus), Mucidus (present or absent), Scutomyia and Oc. (Fin.) harperi.

  • 157(116). Prealar setae: (0) ≤ 20; (1) ≥ 21. A few specimens of some species exhibit a slight overlap of one or two setae between the two states, and are scored (0,1).

  • 158(118). Upper prealar area, scales: (0) absent; (1) present. Scales on the prealar area may be interspersed among the setae on the prealar knob or present on the lower part of the knob (this character) and contiguous with the patch of scales on the lower prealar area (character 159, see Arnell, 1976: fig. 35, Oc. scapularis).

  • 159(117). Lower prealar area, scales: (0) absent; (1) present. When scales are present (state 1) on the lower prealar area (below knob) they form a small to moderately large patch (see Tanaka et al., 1979: figs 217, 220, 222, Oc. (Fin.) japonicus, Ta. togoi and Oc. (Fin.) seoulensis). Lower prealar scales are absent in some species (state 0) (see Tanaka et al., 1979: fig. 239, Ge. baisasi). See discussion under character 158.

  • 160(119). Mesepimeron, scales: (0) absent; (1) present. Scales are normally present (state 1) (see Tanaka et al., 1979: figs 217, 218, Oc. (Fin.) japonicus and Oc. (Fin.) koreicus) in one or two patches on the mesepimeron in Aedini. However, they are absent in Levua and Rhinoskusea (see Reinert, 1976b: fig. 2, Rh. longirostris). See Reinert et al. (2004) for other Aedini without scales on the mesepimeron.

  • 161(120). Lower anterior mesepimeral setae: (0) absent; (1) present. Most taxa examined in the present study have no setae on the lower anterior area of the mesepimeron. These setae are, however, present in Ha. australis, Ia. cavaticus, Mucidus (some species), Oc. (Rusticoidus), Oc. communis, Op. (Opi.) fuscus, Psorophora, Ta. togoi and the outgroup species Cs. inornata, Cx. quinquefasciatus and Ma. titillans. See Reinert et al. (2004) for a discussion of this character in other Aedini and illustrations in Harbach & Kitching (1998: fig. 11H, setae absent, state (0); and 11G, setae present, state (1)).

  • 162(121). Mesepimeron, fine setae on ventral area: (0) absent; (1) present. Setae are absent in all taxa examined in the present study except Geoskusea (see Mattingly, 1959: fig. 2C, Ge. kabaenensis). See Reinert et al. (2004) for other aedine taxa with state (1).

  • 163(122). Metameron: (0) bare; (1) with scales; (2) with setae. The metameron is bare (state 0) in many Aedini. Scales are present (state 1) in Az. ramirezi (absent in some specimens), Oc. (Rusticoidus), Oc. (Ochlerotatus) (several species), Oc. (Fin.) echinus, Oc. (Mol.) pecuniosus (absent in some specimens), Oc. (Pro.) burgeri, Ps. (Jan.) ferox (absent in some specimens) and the outgroup species Cs. inornata. Species of Geoskusea have short, fine setae on the metameron (state 2) (see Mattingly, 1959: fig. 2C, Ge. kabaenensis).

  • 164(125). Alula, marginal scales: (0) absent; (1) narrow; (2) broad. The presence of narrow scales on the margin of the alula (state 1) is the usual condition in Aedini. Mattingly (1957, 1958, 1959) pointed out the value of the varied development of scales on the alula (also see character 165). Broad scales on the margin (state 2) are found in Finlaya, Gymnometopa and Oc. (Fin.) subalbitarsis. Scales are absent (state 0) in Hg. (Hag.) splendens and Ps. (Gra.) jamaicensis. See Harbach & Kitching (1998) for illustrations of state (1) (their fig. 12E) and state (2) (their fig. 12D). See discussion of scale types under character 111.

  • 165(126). Alula, dorsal moderately broad or broad scales: (0) absent; (1) present. See discussion of this character in Reinert et al. (2004). Gymnometopa mediovittata is the only species in the present study that exhibits state (1).

  • 166(127). Remigium, dorsal setae: (0) absent; (1) present, distally; (2) present, proximally. The dorsal remigial setae inserted distally (state 1) (see Tanaka et al., 1979: fig. 219, Oc. (Fin.) hatorii) usually extend noticeably beyond the remigial scales, but are short and hidden beneath the scales in some species. Marks (1962: fig. 1c, Oc. (Cha.) humeralis (Edwards)) illustrated the normally long, stout, blunt-tipped setae (state 2). See Tanaka et al. (1979: fig. 220, Ta. togoi) for an illustration of state (0).

  • 167(128). Remigium, ventral setae: (0) absent; (1) present. Setae are normally absent (state 0) from the ventral surface of the remigium in Aedini. They are present (state 1) on both males and females in the two monobasic subgenera of Opifex (see Belkin, 1968: fig. 2, Op. (Opi.) fuscus), and are also present in the outgroup species Cs. inornata.

  • 168(new). Costa, scales: (0) all dark; (1) one pale-scaled patch at or near base; (2) ≥ 3 pale-scaled patches; (3) pale and dark scales intermixed for all or most of length, not forming defined pattern. Most aedine species have the costa entirely dark-scaled (state 0) (see Tanaka et al., 1979: fig. 217, Oc. (Fin.) japonicus) or have a small to large patch of pale scales at the base of the costa (state 1) (see Tanaka et al., 1979: fig. 222, Oc. (Fin.) seoulensis). A few species, e.g. Oc. (Fin.) banksi and Oc. (Fin.) pseudotaeniatus, have a small dark-scaled patch at the base of the costa and a white-scaled patch near the base on the anterior surface (scored as state 1). Three or more white-scaled patches on the costa (state 2) are typical of genus Finlaya (see Russell, 1996: 61, Fl. kochi). Pale and dark scales are intermixed along the entire length of the costa and do not form a defined pattern (state 3) in some species, e.g. Ps. (Gra.) jamaicensis, Ps. (Pso.) ciliata and the outgroup species Cs. inornata, Ma. titillans and Or. signifera.

  • 169(129). Anal vein: (0) ending approximately at level of intersection of mcu and cubitus; (1) ending noticeably distad of this point. Termination of the anal vein beyond the intersection of the mediocubital crossvein and the cubitus, with the distal portion normally nearly straight or gently curved, is the usual condition in Aedini. Termination near the intersection of mcu and the cubitus, with the distal part sharply curved, occurs in Rhinoskusea, Belkinius, Zeugnomyia and Ae. (Cancraedes)’.

  • 170(131). Anteprocoxal scales: (0) absent; (1) present. Scales are normally absent from the anteprocoxal membrane of Aedini. They are, however, found in subgenus Rusticoidus of Ochlerotatus (present study) and some other species (see Reinert et al., 2004).

  • 171(132). Postprocoxal scales: (0) absent; (1) present. Most taxa examined have the postprocoxal area bare (state 0) (see Zavortink, 1972: fig. 5, Oc. (Pro.) brelandi). Scales are present (state 1) on the postprocoxal membrane in Abraedes, Gymnometopa (see Zavortink, 1972: fig. 38, Gy. mediovittata), Hg. (Haemagogus), Oc. (Rusticoidus), Oc. (Och.) fluviatilis, Oc. (Och.) muelleri, Oc. (Och.) varipalpus and some other Aedini (see Reinert et al., 2004).

  • 172(133). Base of hindcoxa: (0) well below dorsal margin of mesomeron; (1) more or less in line with or slightly above dorsal margin of mesomeron. See the discussion of this character in Reinert et al. (2004). State (1) occurs in Hg. (Haemagogus), Scutomyia and Oc. (Cha.) wattensis in the present study.

  • 173(134). Fore-, mid- and hindfemora, all with narrow, subapical, pale-scaled band: (0) absent; (1) present. In Aedini, the fore-, mid- and hindfemora normally do not have a complete subapical band (state 0). All three femora with a distinct subapical pale-scaled band (state 1) occur in some aedine taxa, e.g. Finlaya, Fredwardsius (see Mattingly, 1965: fig. 5A), Huaedes and Ps. (Grabhamia). Subapical pale-scaled bands are present in Oc. (Fin.) alboannulatus, but these are incomplete on one or more of the femora, and this species is scored (0).

  • 174(new). Midfemur, narrow, median, pale-scaled stripe on anterior surface from base to or near apex: (0) absent; (1) present. Most aedine species do not have a median, pale-scaled stripe on the anterior surface of the midfemur (state 0) (see Tanaka et al., 1979: fig. 223, Oc. (Brh.) alektorovi (=kobayashii)). This feature is present (state 1) in Oc. (Fin.) albilabris, Oc. (Fin.) banksi, Oc. (Fin.) elsiae, Oc. (Fin.) hatorii, Oc. (Fin.) macfarlanei, Oc. (Fin.) notoscriptus and Oc. (Fin.) pseudotaeniatus of the ingroup, and St. aegypti (see Tanaka et al., 1979: fig. 234) and Gy. mediovittata of the aedine outgroup taxa. In the last two species, this stripe is sometimes incomplete in some specimens but is scored (1). Other species have a pale-scaled stripe on the ventral margin of the anterior surface (scored 0).

  • 175(new). Hindfemur, pale scales dorsally and/or anteriorly at apex: (0) absent; (1) present. The apices of the hindfemora in some species of Aedini have dark scales on the dorsal and anterior areas (state 0), e.g. Oc. (Fin.) crossi and Oc. (Fin.) seoulensis (see examples in Tanaka et al., 1979: figs 222, 226, 227, Oc. (Fin.) seoulensis, Do. nipponica and Do. nishikawai). A few species possess a subapical pale-scaled area with a very narrow band or fringe of dark scales at the apex, e.g. Oc. (Fin.) chrysolineatus, Oc. (Fin.) embuensis, Oc. (Fin.) ingrami and Oc. (Fin.) japonicus japonicus (see Tanaka et al., 1979: fig. 217). These species are scored (0). Many species of Aedini have a narrow to broad band of pale scales at the apex of the hindfemur (state 1), e.g. Ta. savoryi and Ta. togoi (see Tanaka et al., 1979: fig. 220).

  • 176(new). Hindtibia: (0) with dark scales only; (1) with pale-scaled areas. The hindtibia may be entirely dark-scaled (state 0), or dark-scaled with pale scales that are intermixed, forming an anterior and/or posterior stripe, developed as a narrow or broad basal, median or apical spot(s) or band(s) (see Banez & Jueco, 1966: pl. 1, Fl. poicilia; Russell, 1996: 61, Fl. kochi), or entirely pale-scaled (state 1).

  • 177(new). Hindtarsomere 1, basal pale scales or band: (0) absent; (1) present. The basal part of hindtarsomere 1 may be dark-scaled (state 0) (see Tanaka et al., 1979: fig. 226, Do. nipponica) or have pale scales forming a narrow to wide band (see Tanaka et al., 1979: fig. 222, Oc. (Fin.) seoulensis) or small spot on the dorsal, ventral or posterior surface (state 1).

  • 178(new). Hindtarsomere 1, one or more median pale-scaled bands: (0) absent; (1) present. The median area of hindtarsomere 1 may be entirely dark-scaled (state 0) or have one or more pale-scaled bands (rarely reduced to dorsal pale-scaled spots) (state 1) as in Finlaya (see Banez & Jueco, 1966: pl. 1, Fl. poicilia; Russell, 1996: 61, Fl. kochi).

  • 179(new). Hindtarsomere 1, apical pale scales or band: (0) absent; (1) present. The apical part of hindtarsomere 1 may be dark-scaled (state 0) (see Zavortink, 1972: fig. 42, Oc. (Och.) muelleri) or have pale scales forming a narrow to wide band or small spot on the dorsal, ventral or posterior surface (state 1) (see Zavortink, 1972: fig. 48, Oc. (Och.) atropalpus).

  • 180(new). Hindtarsomere 2, basal pale scales or band: (0) absent; (1) present. See discussion under character 177.

  • 181(new). Hindtarsomere 2, apical pale scales or band: (0) absent; (1) present. See discussion under character 179.

  • 182(136). Foreungues: (0) both simple; (1) one simple, one toothed; (2) both toothed. Only a few taxa examined have females with both foreungues simple (state 0), e.g. Abraedes, Geoskusea, Howardina (see Berlin, 1969: fig. 16, Hw. walkeri), Levua, Rhinoskusea, Scutomyia and the outgroup species Cs. inornata, Ma. titillans and Or. signifera. State (1) with one unguis simple and the other unguis toothed occurs in the outgroup species Cx. quinquefasciatus. Most aedine taxa have both foreungues toothed (state 2) (see Tanaka et al., 1979: figs 219, 220, Oc. (Fin.) hatorii and Ta. togoi).

  • 183(137). Foreungues (males): (0) both simple; (1) one simple, one toothed; (2) both toothed. The male foreungues are both simple (state 0) and large in Op. (Opi.) fuscus (see Belkin, 1968: figs 2, 18). One unguis is simple and the other toothed (state 1) in specimens examined of Abraedes, Aedes, Hg. (Hag.) splendens, Hw. sexlineata, Levua, Oc. (Cha.) wattensis, Oc. (Och.) muelleri, Rhinoskusea, Scutomyia, Stegomyia and the outgroup species Ma. titillans and Or. signifera. The other species examined have both foreungues toothed (state 2).

  • 184(new). Larger foreunguis, development (males): (0) simple; (1) with 1 tooth; (2) with 2 teeth. The larger foreunguis of males may be simple (state 0) as in Op. (Opi.) fuscus (see Belkin, 1968: figs 2, 18) and St. africana, have a single tooth (state 1) as in Abraedes, Aedes, Gymnometopa (see Zavortink, 1972: fig. 38, Gy. mediovittata), Haemagogus, Isoaedes, Kenknightia, Rhinoskusea, Scutomyia and others, or have two teeth (state 2) (see Zavortink, 1972: fig. 35, Az. ramirezi), one small tooth usually at or near the base and a second larger tooth near midlength as in many of the species examined.

  • 185(138). Midungues (males): (0) both simple; (1) one simple, one toothed; (2) both toothed. Simple midungues (state 0) occur in males of Op. (Opi.) fuscus, Le. geoskusea, Sc. albolineata and St. aegypti. Some species, e.g. Ab. papago, Hg. (Hag.) splendens, Oc. (Cha.) wattensis and Oc. (Mac.) tremulus, exhibit state (1). Most species examined have both midungues toothed (state 2).

  • 186(new). Larger midunguis, development (males): (0) simple; (1) with 1 tooth; (2) with 2 teeth. Males of only a few taxa examined have the larger midunguis simple (state 0), e.g. Hg. (Con.) leucotaeniatus, Le. geoskusea, Op. (Opi.) fuscus, Scutomyia and Stegomyia. The larger midunguis with a tooth (state 1) occurs in several species, e.g. Abraedes, Aedes, Geoskusea, Isoaedes, Kenknightia, Oc. (Mac.) tremulus, Op. (Not.) chathamicus and Zavortinkius. Numerous other taxa have the larger unguis with two teeth (state 2), e.g. Aztecaedes, Downsiomyia, Finlaya, Psorophora and Tanakaius.

  • 187(139). Hindungues (males): (0) both simple; (1) both toothed. Males of most species examined have both hindungues simple (state 0) but some have both toothed (state 1), e.g. Ha. australis, Mu. alternans, Mu. laniger, Op. (Not.) chathamicus, Ps. (Jan.) ferox, Ps. (Pso.) ciliata, Tanakaius and Zavortinkius.

  • 188(140). Hindungues: (0) both simple; (1) both toothed. Females of most taxa examined have both hindungues simple (state 0), but those of a few taxa have both toothed (state 1), e.g. Aedes, Halaedes, Mu. alternans, Mu. laniger, Opifex, Tanakaius, Zavortinkius and subgenus Rusticoidus of Ochlerotatus.

  • 189(141). Laterotergite, scales: (0) absent; (1) present. Scales are absent (state 0) in Geoskusea, Oc. (Fin.) candidoscutellum, Oc. (Och.) atropalpus, Oc. (Och.) epactius, Oc. infirmatus, Oc. scapularis, Opifex, Ps. (Pso.) ciliata and the four outgroup species. Most Aedini have numerous, or at least a few, scales on the laterotergite of abdominal segment I (state 1).

  • 190(142). Abdominal terga, lateral setae (males): (0) few, relatively short; (1) numerous, short to moderately long; (2) numerous, long, with curved apices. The lateral setae on the abdominal terga of males are few in number and relatively short (state 0) in some taxa, e.g. Geoskusea, Hg. (Conopostegus), Hg. (Hag.) splendens, Howardina, Isoaedes, Kenknightia, Oc. (Bruceharrisonius)’, Op. (Opi.) fuscus, Rhinoskusea, Scutomyia, St. aegypti, Tanakaius and Zavortinkius. These setae are numerous and relatively short to moderately long (state 1) in other taxa, e.g. Downsiomyia, Finlaya, Oc. (Mac.) tremulus and Oc. (Mol.) pecuniosus. The setae are numerous and long with apices curved (state 2) in numerous taxa, e.g. Aedes, Mu. alternans, Mu. laniger, Oc. (Protomacleaya) (most species) and Oc. (Ochlerotatus) (many species).

  • 191(143). Abdominal segment VII, shape: (0) laterally compressed; (1) dorsoventrally flattened; (2) cylindrical. Abdominal segment VII (and often segment VIII) of dead, dried females has a distinctive shape when viewed transversely, i.e. laterally compressed, dorsoventrally flattened or cylindrical. The cylindrical shape occurs in Psorophora and the outgroup species Ma. titillans. See Tanaka et al. (1979: fig. 229, Oc. (Fin.) watasei) for state (0) and their fig. 212 (Oc. dorsalis) for state (1).

Female genitalia

The series of papers by Reinert (2000b, c, d, e, f, 2001b, c, d, e, 2002b, c, d, e) on the comparative anatomy of Aedini should be consulted for illustrations and descriptions of female genital structures.

  • 192(144, in part). Intersegmental membrane between segments VII and VIII: (0) short to intermediate; (1) long to very long. Taxa with intersegmental membranes separating segments VII and VIII that allow 50% or less of tergum VIII to be retracted into segment VII are considered short to intermediate whereas those allowing 65% or more retraction are scored long to very long. The very long intersegmental membrane in Psorophora allows for nearly complete retraction of segment VIII into segment VII, whereas taxa with short intersegmental membranes do not allow retraction or only a slight retraction of segment VIII.

  • 193(145, in part). Tergum VIII, development: (0) mostly membranous, with lateral rod-shaped structures; (1) entirely sclerotized (rarely with only small median nonsclerotized area), without lateral rod-shaped structures. Psorophora have tergum and sternum VIII nearly completely membranous except for small scattered, sclerotized islands usually bearing setae and a rod-shaped structure laterally on each side (state 0) (see Reinert, 2000c: figs 1–3). Most Aedini have tergum and sternum VIII entirely sclerotized (state 1) (see figures in Reinert, 2000f, 2001c, 2002e). Tergum VIII of the outgroup species Ma. titillans has distal and lateral sclerotized areas and a large basomesal membranous area (scored 1). Sternum VIII is sclerotized in this species.

  • 194(146). Tergum VIII, setae: (0) on distal 0.60 or less; (1) on distal 0.70 or more. Some aedine taxa have most of the surface (≥ 0.70) of tergum VIII covered with several to numerous setae (state 1), e.g. Aedes, Geoskusea, Halaedes, Kompia, Levua, Oc. (Bruceharrisoniu s)’, Oc. (Ochlerotatus) (most species), Oc. (Pseudoskusea)’, Oc. (Rusticoidus), Op. (Nothoskusea), Ps. (Janthinosoma), St. aegypti and Ta. togoi. Other groups have few to several setae that are restricted to the distal 0.60 or less of the tergum (state 0).

  • 195(147, in part). Tergum VIII, scales: (0) absent, occasionally with 1–3 adventitious scales; (1) numerous. Most aedines examined in the present study have numerous scales on tergum VIII (14 or more scales, some species with scales nearly covering the tergum), especially on the distal part (state 1). Some species have no scales (state 0), e.g. Geoskusea, Levua, Oc. communis, Oc. infirmatus, Oc. scapularis, Oc. (Och.) deserticola, Oc. (Pseudoskusea)’, Psorophora, Rhinoskusea and the outgroup species Ma. titillans. A few species, e.g. Oc. (Fin.) albilabris and Oc. (Fin.) notoscriptus, have no scales or only 1–3 (rarely a specimen with 4 or 5) scales scattered over the tergum, and are scored (0).

  • 196(148). Sternum VIII, position of seta 2-S: (0) insertion noticeably posterior to seta 1-S; (1) insertion laterally at or near same level as seta 1-S. A number of taxa have numerous long setae along the median area that do not occur in a distinct pattern. Since these setae are mainly arranged longitudinally, they are scored as state (0).

  • 197(149, in part). Sternum VIII, scales: (0) absent, occasionally with 1–3 adventitious scales; (1) several to numerous scales, often covering much of surface. Some species of Aedini have no scales on sternum VIII; however, some specimens occasionally have 1–3 scales scattered over the surface (state 0) (see Reinert, 2002e: fig. 3, Oc. (Fin.) geniculatus). Ochlerotatus (Fin.) macfarlanei has no scales or rarely one or two present, but the sternum has numerous short lanceolate setae interspersed with slender setae (scored 0). Most aedine species have several (10 or more scales) (see Reinert, 2002e: fig. 5, Oc. (Fin.) gubernatoris) or numerous scales covering most of the surface of sternum VIII (see Reinert, 2002e: figs 4, 6, Oc. (Brh.) greenii and Fl. kochi) (state 1).

  • 198(150, in part). Tergum IX: (0) wide, width ≥ 2.0 times length; (1) narrower, width ≤ 1.8 times length. Few aedines have tergum IX width greater than or equal to 2.0 the length, e.g. Oc. (Fin.) banksi, Oc. (Fin.) embuensis, Oc. (Fin.) pseudotaeniatus, Oc. (Och.) atropalpus, Oc. (Och.) epactius and Oc. (Och.) fluviatilis. This also applies to the four outgroup species. The three species of Oc. (Ochlerotatus) listed here also have tergum IX divided into two small sclerites that are separated by membrane (see character 199). The outgroup taxa, e.g. Cx. quinquefasciatus, Ma. titillans (see Gerry, 1932: figs 28, 40) and Cs. inornata (see Rees & Onishi, 1951: fig. 35) have a wide and narrow tergum IX.

  • 199(new). Tergum IX, development: (0) single sclerite, usually with small to deep apical emargination; (1) two sclerites connected by membrane. Most aedine species have tergum IX as a single sclerite (state 0). This sclerite may be entirely sclerotized or have a narrow mesal connection between the lateral parts, and it may be darkly pigmented or have the mesal area slightly paler. Two lateral sclerites connected by membrane (state 1) are found in Ke. dissimilis, Ke. harbachi, Oc. (Och.) atropalpus, Oc. (Och.) epactius, Oc. (Och.) fluviatilis, Op. (Not.) chathamicus and Op. (Opi.) fuscus.

  • 200(151). Tergum IX, setae: (0) absent; (1) present distally; (2) present, distally and on much of medial area. Tergum IX with no setae (state 0) (see Reinert, 2002e: fig. 9, Oc. (Fin.) papuensis) occurs in a few species. Most aedine species have one or more setae distally on tergum IX (state 1) (see Reinert, 2002e: figs 2, 5, 9, Oc. (Finlaya) species). Psorophora and Az. ramirezi have setae on the distal area that extend along the mesal area (state 2) (see Reinert, 2000c: figs 1–3, Psorophora).

  • 201(152, in part). Postgenital lobe, position of ventral setae: (0) distal; (1) lateral; (2) on entire surface. After examining a larger series, including new specimens, of the outgroup species, this character was reevaluated; hence, Cs. inornata (see Rees & Onishi, 1951: fig. 35), Cx. quinquefasciatus (see Sirivanakarn, 1976: fig. 14), Ma. titillans (see Ronderos & Bachmann, 1963: fig. 27) and Or. signifera are now scored (1). The presence of numerous (usually 10 or more) setae in elongate, lateral patches on the ventral surface of the postgenital lobe of these species differs from the species of Aedini examined (see Reinert, 2002e: figs 3, 12, 34, 38, Oc. (Fin.) geniculatus, Ke. dissimilis, Oc. (Pro.) triseriatus and Oc. (Rus.) refiki), which have setae located distally on the postgenital lobe, scored 0, except Op. (Opi.) fuscus, see Reinert, 2001c: fig. 1, which has numerous setae covering the entire ventral surface and is scored (2). Some aedine species have a median apical emargination forming two lobes, each bearing a few to several setae distally (see Reinert, 2002e: fig. 14, Le. geoskusea), and these are scored (0).

  • 202(153). Upper vaginal sclerite: (0) absent; (1) present. An upper vaginal sclerite is present in most of the species examined, including most species of Oc. (Finlaya)’. The sclerite may be small and restricted to the inner margin of the upper vaginal lip, or extend as a well-developed plate mesally on the upper vaginal wall. This sclerite is absent, e.g. Aztecaedes, Hg. (Conopostegus), Kompia, Oc. (Finlaya) (few species), Oc. (Ochlerotatus) (most species), Oc. (Rusticoidus), Oc. communis, Oc. infirmatus, Oc. scapularis, Psorophora, Rhinoskusea and Tanakaius. The upper vaginal sclerite is present in the outgroup species Cs. inornata, Cx. quinquefasciatus, Ma. titillans and Or. signifera.

  • 203(154). Lower vaginal sclerite: (0) absent; (1) present. The lower vaginal sclerite is normally absent in Aedini. A poorly developed sclerite on the inner margin of the lower vaginal lip is present in Ge. baisasi. Culex quinquefasciatus has a well-developed, horseshoe-shaped, lower vaginal sclerite located medially on the lower vaginal wall between the inner margins of the lower vaginal lip.

  • 204(155, in part). Insula: (0) tonguelike; (1) liplike. Most taxa examined in the present study have a liplike insula (state 1) that is noticeably wider than long. Taxa examined that have a tonguelike (lingulate) insula (state 0) include Aedes, Haemagogus, Isoaedes, Psorophora, Scutomyia and Stegomyia. See Reinert et al. (2004) for a discussion of this character in tribe Aedini.

  • 205(156). Insular setae: (0) absent; (1) present in lateral patches; (2) present in medial patch. The majority of taxa examined in the present study have lateral patches of setae on the insula (state 1). Aedes, Isoaedes, Scutomyia and Stegomyia have no setae (state 0) but instead may have small tuberculi (each with or without a tiny spicule). Haemagogus and the outgroup taxa Cs. inornata and Cx. quinquefasciatus have setae in a median patch (state 2).

  • 206(new). Cercus, scales: (0) absent; (1) present. Most species have no scales on the cerci (0) (see Appendix 2). Some specimens of a few species have one adventitious scale on one cercus, and are scored (0).

  • 207(157). Spermathecal capsule (s): (0) one; (1) three. Downsiomyia (except two species) and Rhinoskusea have a single large spermathecal capsule (state 0). The other species studied have one large and two smaller spermathecal capsules (state 1). Reinert et al. (2004) provide information on this character in Aedini.

Male genitalia

  • 208(new). Tergum IX, posterior margin: (0) with pair of strongly sclerotized, darkly pigmented, short lobes, often narrow and acute; (1) with pair of poorly sclerotized, lightly to moderately pigmented, broad, gently rounded or flattened lobes. See Zavortink (1972: figures of Oc. (Protomacleaya)’) and Tanaka et al. (1979: fig. 91, Oc. (Och.) hakusanensis) for state (0) and Belkin (1962: figs 207, 233, Op. (Opi.) fuscus and Oc. (Fin.) argyronotum) for state (1). Also, see the discussion under character 209.

  • 209(158). Tergum IX, setae: (0) absent; (1) all slender; (2) some or all stout. Species with strongly sclerotized, short lobes normally bear stout and often flattened setae. Those with weakly sclerotized, broad, gently rounded lobes normally bear longer, slender setae, and if the lobes are flattened and often sloping the setae are often absent (see discussion for character 208).

  • 210(159). Sternum IX, length: (0) short; (1) moderately long; (2) long. Sternum IX is moderately long (state 1) in the majority of aedine species. It is short (state 0) in a few species, e.g. Aedes, Oc. (Fin.) albocinctus, Oc. (Fin.) banksi and Oc. (Fin.) pseudotaeniatus, and relatively long (state 2) in Ge. longiforceps, Rhinoskusea (see Reinert, 1976b: figs 6–8) and Oc. (Och.) sierrensis.

  • 211(160). Sternum IX, setae: (0) absent; (1) present. Most taxa have setae present on sternum IX (state 1). Setae are absent (state 0), e.g. Ia. cavaticus, Op. (Opi.) fuscus, St. aegypti, St. africana and the outgroup species Cx. quinquefasciatus. Setae are present or absent (scored 0,1) in specimens of the outgroup species Ma. titillans and Or. signifera.

  • 212(new). Gonocoxite, dorsomesal apical lobe: (0) absent; (1) present. The dorsomesal apical lobe is absent (state 0) in most taxa examined in the present study. The lobe is present (state 1) in Oc. communis, Oc. infirmatus and Oc. scapularis (see Arnell, 1976: fig. 30, Oc. infirmatus, lobe well developed; fig. 36, Oc. scapularis, lobe moderately developed), Oc. (Rusticoidus) and Rhinoskusea. The outgroup species Cx. quinquefasciatus has a well-developed lobe bearing specialized setae mesally on the dorsal surface of the gonocoxite, but the lobe is subapical. This is scored (0).

  • 213(new). Gonocoxite, dorsomesal basal lobe: (0) absent; (1) present. Geoskusea lack a basomesal lobe on the dorsal surface of the gonocoxite (state 0), but most species (all except Ge. longiforceps) have a densely setose lobe on this surface at approximately midlength of the mesal margin (see Belkin, 1962: figs 214, 215, Ge. becki, Ge. daggyi and Ge. perryi; Tanaka et al., 1979: fig. 127, Ge. baisasi).

  • 214(163). Gonocoxite, scales: (0) absent; (1) present. Most species of Aedini have scales on the gonocoxite (state 1) (see Belkin, 1962: figs 210, 226). A few species have no scales on the gonocoxite (state 0), e.g. Op. (Opi.) fuscus (see Belkin, 1962: fig. 207), Ps. (Pso.) ciliata, as well as the outgroup species Cs. inornata and Cx. quinquefasciatus.

  • 215(162). Gonocoxite, mesal surface: (0) entirely membranous; (1) partly or entirely sclerotized. The mesal surface of the gonocoxite is commonly membranous (state 0) in the species of Aedini examined, however, this area is partially or completely sclerotized (state 1) in Psorophora, Rhinoskusea and the outgroup species Cs. inornata, Cx. quinquefasciatus and Or. signifera.

  • 216(new). Gonocoxite, seta (e) on basomesal area of dorsal surface: (0) all slender; (1) one or more stout or flattened; (2) absent. The basomesal area of the gonocoxite dorsal surface bears only slender setae in state (0) (see illustrations of Oc. (Protomacleaya) in Zavortink, 1972). This area in state (1) has one or more of the setae stout and/or flattened (see Wood et al., 1979: pl. 20, Oc. communis). In Ge. baisasi this area is bare (see Tanaka et al., 1979: fig. 127) and is scored (2).

  • 217(161). Gonocoxite, lateral setae: (0) mostly short; (1) mostly long. Lateral setae are considered short if they are less than 0.80 of the width of the gonocoxite dorsal surface at the point of attachment whereas they are long if they are longer than, and normally noticeably longer than, the width. In most Aedini, the lateral surface of the gonocoxite has numerous long, stout setae (state 1) (see Belkin, 1962: figs 276, 278, Mu. alternans and Mu. painei). Most of the lateral setae of the gonocoxite in Op. (Opi.) fuscus are short (state 0) (see Belkin, 1962: fig. 207).

  • 218(new). Gonocoxite, ventral surface with specialized scales on mesal margin: (0) absent; (1) present. The gonocoxites of Aedini usually lack specialized scales on the mesal margin of the ventral surface (state 0) (see Belkin, 1962: figs 273, 276, 315, Le. geoskusea, Mu. alternans and Sc. albolineata). Well-developed, long, specialized scales (state 1) are found in Downsiomyia, Finlaya (see figures in Belkin, 1962 of Finlaya, as Kochi Group), Haemagogus, Oc. (Fin.) albotaeniatus, Oc. (Fin.) biocellatus, Oc. (Fin.) harperi, Oc. (Fin.) pulchriventer and Tanakaius.

  • 219(164). Gonostylus, attachment to gonocoxite: (0) subapical; (1) apical. The dorsal surface of the gonocoxite extends noticeably beyond the attachment of the gonostylus in Op. (Opi.) fuscus (see Belkin, 1962: fig. 207) whereas the ventral surface extends noticeably beyond the attachment of the gonostylus in the Africanus Group of Stegomyia (see Huang, 1990: fig. 23A, B, St. opok and St. pseudoafricana) and genus Aedes (see Tanaka et al., 1979: fig. 133, Ae. esoensis and Ae. yamadai), and these taxa are scored (0). In Op. (Not.) chathamicus, the dorsal surface of the gonocoxite extends slightly beyond the attachment of the gonostylus, but is scored (1).

  • 220(166, in part). Gonostylus, development: (0) moderately long to long, relatively narrow throughout length but distal part narrower and usually somewhat curved mesally; (1) same as state 0 but distal part somewhat expanded; (2) short to moderately long, basal area broader, bifurcate; (3) proximal and distal parts narrow, middle part noticeably expanded. The gonostylus is normally moderately long to long, relatively narrow throughout its length and the distal part is somewhat narrower and usually slightly curved mesad (state 0) in most aedines examined in the present study (see Belkin, 1962: figs 226, 235, 276, Oc. (Fin.) notoscriptus, Fl. samoana and Mu. alternans). Rhinoskusea (see Reinert, 1976b: fig. 6, Rh. longirostris) and most Stegomyia have the proximal part of the gonostylus relatively narrow and the distal part expanded (state 1). State (2) occurs in a few species, e.g. Aedes, Op. (Opi.) fuscus (see Belkin, 1962: fig. 207) and the outgroup species Ma. titillans. Psorophora have the proximal and distal parts relatively narrow and the middle part noticeably expanded (see Belkin et al., 1970: figs 57, 59, 66, Ps. (Pso.) ciliata, Ps. (Jan.) ferox and Ps. (Gra.) jamaicensis), state (3).

  • 221(new). Gonostylus, subapical seta (e): (0) absent; (1) one or more present. The gonostylus of most aedines studied have one or a few normally short, fine subapical setae (state 1) (see Tanaka et al., 1979: fig. 97, Oc. (Fin.) japonicus), but these setae are stouter (state 1) in some species (see Barraud, 1934: fig. 37d, e, Oc. (Fin.) elsiae and Oc. (Fin.) shortti). Subapical setae are absent (state 0) in some taxa (see Belkin, 1962: fig. 315, Sc. albolineata), but Scutomyia possess a single long, stout spiniform inserted some distance from the apex (see discussion for character 222).

  • 222(167, in part). Gonostylar claw: (0) absent; (1) present, one narrow, short to long spiniform, insertion at or near apex; (2) present, one striated, short, spatulate spiniform, insertion at or near apex; (3) present, one stout, short, clawlike spiniform, insertion at or near apex; (4) present, one stout, long spiniform, insertion some distance from apex; (5) present two or more narrow, short to moderately long spiniforms, insertion distally; (6) present, two stout, short, side by side, clawlike spiniforms, insertion at or near apex. Aedes lack a spiniform (state 0) (see Tanaka et al., 1979: fig. 133, Ae. esoensis and Ae. yamadai). Most aedines examined exhibit state (1) (see Belkin, 1962: figs 210, 254, Ha. australis and Fl. neogeorgiana). A single, short, striated, spatulate claw (state 2) occurs in Rhinoskusea (see Reinert, 1976b: fig. 6, Rh. longirostris) and the outgroup species Or. signifera. A single, dark, stout, clawlike spiniform (state 3) occurs in Op. (Opi.) fuscus (see Belkin, 1962: fig. 207), Psorophora and the outgroup species Ma. titillans. Scutomyia (see figures in Huang, 1979) have a single, stout, long spiniform inserted some distance from the apex (state 4). Two or more narrow, short to moderately long spiniforms inserted on the distal part of the gonostylus (state 5) occur in Oc. (Fin.) elsiae, Oc. (Fin.) macfarlanei and related species (see Barraud, 1934: fig. 37). Two dark, short, clawlike spiniforms inserted side by side near the apex (state 6) occur in Le. geoskusea (see Belkin, 1962: fig. 273) and the outgroup species Cs. inornata. Ochlerotatus (Fin.) biocellatus has a moderately long, slender gonostylar claw with the distal part forked into two attenuate branches (scored 1).

  • 223(new). Gonostylar claw, length: (0) short, ≤ 0.35 gonostylus length; (1) moderately long to long, ≥ 0.39 gonostylus length. Reinert (1990) defined the gonostylar claw length as the ‘absolute length measured from the base to the apex’ and the gonostylus length as the ‘absolute length of gonostylus, minus gonostylar claw, measured from the tip of the basal apodeme to the apex’. Species that do not have a gonostylar claw (see previous character), e.g. Aedes, are scored as missing (–).

  • 224(new). Gonostylus/gonocoxite index: (0) ≤ 0.40; (1) ≥ 0.42. The gonostylus/gonocoxite index was defined by Reinert (1990) as the ‘ratio determined by dividing the gonostylus length by the gonocoxite length’. Length of the gonocoxite (Reinert, 1990) is the ‘length measured along a straight line from the tip of the dorsal attachment of gonocoxite (dga of Knight & Harrison, 1988) to the apex’. See discussion under character 223 for measurement of gonostylus length.

  • 225(168, in part). Claspette, development: (0) basal setose plaque variously developed (setae simple, specialized or both); (1) single short to long, slender, columnar lobe; (2) two lobes, one long and columnar. The claspettes and the basal mesal lobes are homologous structures in Aedini and can be identified as such by their connection basomesally to a spiculate, more-or-less narrow, somewhat troughlike aedeagal guide (see Reinert, 1999b, 2000a). Reinert (2000a) noted that this structure is apparently nearly lost in species of Geoskusea and Oc. (Pseudoskusea) because it appears to be reduced to a spiculate aedeagal guide and a short thin strip extending laterally onto the basosternal area of the mesal surface of the gonocoxite, and bears one or a few short, thin, inconspicuous setae. He also reported that the claspette is without a filament in Rhinoskusea, and is bifurcate in Rh. longirostris and Rh. wardi. In Oc. (Bruceharrisonius) , the claspette comprises a median, ventral, columnar stem with a terminal, large, flattened filament, and a narrow, lateral arm extending from the base of the stem that bears few to several setae, some of which are flattened (scored 2) (see Reinert, 2003: fig. 2A, Oc. (Brh.) greenii). The filament is absent in Op. (Not.) chathamicus and the stem is developed as a bluntly rounded, lobelike process that is densely covered with short, thin setae (see Belkin, 1968: fig. 20). In Halaedes, the claspette appears to be developed as an oblong plaque bearing a number of curved, lanceolate setae, and the dorsal lateral part is fused with the basomesal portion of the tergal surface of the gonocoxite (see Belkin, 1962: fig. 210). Arnell (1973) pointed out that in Hg. (Conopostegus) and two species of subgenus Haemagogus, i.e. Hg. anastasionis and Hg. chrysochlorus, the claspette filament is inserted in a distinct alveolus on the apex of the claspette stem. However, in the other species of Haemagogus the alveolus is either incomplete or entirely absent at the base of the filament. Species of Haemagogus are scored (1). Psorophora have the claspette variously developed from a short plaque, e.g. Ps. (Gra.) jamaicensis, to a long, flattened stem with distal part expanded, e.g. Ps. (Jan.) ferox and Ps. (Pso.) ciliata, that projects dorsocaudally and bears a few to several specialized setae with flattened, fringed or simple apices, or several slender, simple setae, and none or one or more flattened, broadened and strongly contorted filaments (see Belkin et al., 1970: figs 57, 59, 66). These species of Psorophora are scored as (0). In state (2), the slender lobe is normally distinctly columnar and may be short to long. The claspette is absent in the outgroup species Cx. quinquefasciatus, and is scored (–). In this species the lobe with modified setae that arises from the mesal area on the dorsal surface of the gonocoxite is not homologous with the claspette in Aedini, which arises from the ventromedian area near the base of the gonocoxite and is connected mesally with its mate by an aedeagal guide.

  • 226(new). Claspette filament(s), development: (0) ≥ 2 setae; (1) one slender or stout seta; (2) one stout, curved spiniform, elliptical or circular in cross section; (3) one moderately wide to wide, normally elongate, flattened structure. In Oc. (Rusticoidus), the claspette filament is relatively short but transversely annulate (see Gutsevich, Monchadskii & Shtakel'berg, 1974: figs 133, 135, Oc. (Rus.) rusticus and Oc. (Rus.) refiki), and is scored (3). This character is scored as missing (–) for Cx. quinquefasciatus. See discussion under character 225.

  • 227(new). Claspette filament, distinct transverse striations: (0) absent; (1) present. In most species examined in the present study the claspette filament does not have transverse striations (state 0). The claspette filament of Finlaya is long, foliform and with a membranelike sheath or flap that is annulated transversely on approximately the middle third whereas the filament of several species of Oc. (Finlaya) have the sheath absent but with several transverse striations near midlength. These are scored (1). The filament in Oc. (Rusticoidus) has transverse striations but it is shorter, broader and thicker (scored 1) (see discussion under character 226). This character is scored (–) for Cx. quinquefasciatus (see discussion under character 225).

  • 228(170). Aedeagus: (0) single tubelike, scooplike or troughlike structure; (1) comprising two lateral plates (aedeagal sclerites). State (0) occurs in all species examined in the study except in Aedes, Isoaedes, Scutomyia, Stegomyia and the outgroup species Cs. inornata and Cx. quinquefasciatus, which have two lateral plates (state 1). Reinert (2000a) and Reinert et al. (2004) discussed this character and its importance in separating generic-level taxa of Aedini.

  • 229(new). Aedeagus, width: (0) widest in distal third; (1) widest in middle third; (2) widest in proximal third. The various states of the aedeagal width are illustrated in Zavortink (1972), i.e. state (0) fig. 39, Gy. mediovittata, state (1) fig. 43, Oc. (Och.) muelleri and state (2) fig. 15, Oc. (Pro.) burgeri.

  • 230(171). Proctiger, sternal arm: (0) absent; (1) present. The sternal arm of the proctiger is absent (state 0) in all species evaluated except for the following (state 1): Ia. cavaticus, St. aegypti, St. africana and the outgroup species Cx. quinquefasciatus.

  • 231(172). Proctiger, cercal setae: (0) absent; (1) present. Most aedine species examined have cercal setae present on the proctiger (state 1) (see Belkin, 1962: fig. 233, Oc. (Fin.) argyronotum). Opifex (Opi.) fuscus is unusual in that both minute and short cercal setae are present on the proctiger (see Belkin, 1962: fig. 207). Cercal setae are absent (state 0) in Aedes, Isoaedes (see Reinert, 1979: fig. 3, Ia. cavaticus), Scutomyia and Stegomyia.

Biology

  • 232(new). Immature habitat: (0) freshwater ground pools; (1) brackish-water ground and rock pools; (2) freshwater rock pools; (3) freshwater containers (phytotelmata, small artificial containers, rock holes); (4) crab holes. The female selects the habitat in which the immature stages develop when she deposits her eggs. The primary habitat of each study taxon is listed in the data matrix (Appendix 2). Atypical habitats are not included. Belkin (1962) classified mosquitoes into two principal groups based on the habitat requirements of the immature stages, i.e. species that inhabit ground-water habitats and those that inhabit plant-container habitats. He considered the former as undoubtedly ancestral and the later derived. Reinert (2002a) indicated that aedine species with a well-developed ventral brush (setae 4-X) normally inhabit ground-water habitats, and the strong development of the brush seems to be the ancestral condition. By contrast, species with a weakly developed ventral brush inhabit plant containers, and the weaker brush seems to be the more derived condition.

PHYLOGENETIC ANALYSIS

Reinert et al. (2004) conducted phylogenetic analyses using both equal and implied weighting, applied to adult data and immature stage data separately, as well as the data set as a whole. They concluded that down-weighting the more homoplastic characters by the application of implied weights was preferable, and that this procedure should be applied to the entire data set in a ‘total evidence’ approach. We therefore analysed the present data using only implied weights as implemented by PIWE version 3.0 (for Windows) (Goloboff, 1997), with the default value of the concavity constant, K = 3. We performed the analysis by heuristic search, using 5000 replications (mult*5000) and 10 starting cladograms per replicate (hold/10). We then checked that PIWE had found all most parsimonious (‘fittest’) cladograms (MPCs) by searching for successively less fit cladograms using the commands ‘sub n’ (where n is the decrease in fit, in steps of 0.1) and ‘find*’ (to search for all cladograms of best fit–n), up to a maximum of 100 000 cladograms. We then applied the ‘best’ command to this set of 100000 cladograms to confirm that the included set of MPCs was the same as that with which we started. Agreement in grouping within a set of MPCs was summarized using a strict consensus tree (SCT). We assessed clade support using Bremer support (Bremer, 1994) and relative Bremer support (Goloboff & Farris, 2001). Illustrated cladograms were prepared as described by Reinert et al. (2004) except they were first cropped, resized and positioned on pages with CorelDraw 10 before being imported into Microsoft Word 2000 for final editing.

RESULTS AND DISCUSSION

Analysis of the entire data set under implied weights produced four MPCs (Fit = 836.8), the SCT of which (including Bremer and relative Bremer supports for each clade) is shown in Figure 1. The characters that support the branches of this SCT are shown in Figure 2. It should be emphasized that because this is a strict consensus tree, character mappings on branches immediately subtending and arising from polytomies are not necessarily as parsimonious as they are on the MPCs from which the SCT was derived. In particular, apparent parallel gains in clades arising from polytomies may be synapomorphies for subsets of these clades on some (but not all) MPCs. Thus, caution is required not to over-interpret these mappings.

Figure 1

Strict consensus of four MPCs (Fit = 836.8) obtained from the analysis of the data (Appendix 2) under implied weights. Bremer support and relative Bremer support values of each clade are indicated above and below the branches, respectively. Circled numbers adjacent to terminal taxa indicate newly recognized genera: 1, Pseudoskusea stat. nov. ; 2, Himalaius gen. nov. ; 3, Bruceharrisonius stat. nov. ; 4, Vansomerenis gen. nov. ; 5, Macleaya stat. nov. ; 6, Jihlienius gen. nov. ; 7, Molpemyia stat. nov. ; 8, Phagomyia stat. nov. ; 9, Rampamyia gen. nov. ; 10, Dobrotworskyius gen. nov. ; 11, Georgecraigius gen. nov. ; 12, Patmarksia gen. nov. ; 13, Hulecoeteomyia stat. nov. ; 14, Gilesius gen. nov. ; 15, Collessius gen. nov. ; 16, Dahliana gen. nov. ; 17, Jarnellius gen. nov. Species of Oc. (Protomacleaya) are shaded.

Figure 1

Strict consensus of four MPCs (Fit = 836.8) obtained from the analysis of the data (Appendix 2) under implied weights. Bremer support and relative Bremer support values of each clade are indicated above and below the branches, respectively. Circled numbers adjacent to terminal taxa indicate newly recognized genera: 1, Pseudoskusea stat. nov. ; 2, Himalaius gen. nov. ; 3, Bruceharrisonius stat. nov. ; 4, Vansomerenis gen. nov. ; 5, Macleaya stat. nov. ; 6, Jihlienius gen. nov. ; 7, Molpemyia stat. nov. ; 8, Phagomyia stat. nov. ; 9, Rampamyia gen. nov. ; 10, Dobrotworskyius gen. nov. ; 11, Georgecraigius gen. nov. ; 12, Patmarksia gen. nov. ; 13, Hulecoeteomyia stat. nov. ; 14, Gilesius gen. nov. ; 15, Collessius gen. nov. ; 16, Dahliana gen. nov. ; 17, Jarnellius gen. nov. Species of Oc. (Protomacleaya) are shaded.

Figure 2

Strict consensus tree shown in Figure 1 with supporting, unambiguously optimized characters mapped on the branches. Circled numbers adjacent to terminal taxa correspond to the newly recognized genera indicated in Figure 1: 1, Pseudoskusea stat. nov. ; 2, Himalaius gen. nov. ; 3, Bruceharrisonius stat. nov. ; 4, Vansomerenis gen. nov. ; 5, Macleaya stat. nov. ; 6, Jihlienius gen. nov. ; 7, Molpemyia stat. nov. ; 8, Phagomyia stat. nov. ; 9, Rampamyia gen. nov. ; 10, Dobrotworskyius gen. nov. ; 11, Georgecraigius gen. nov. ; 12, Patmarksia gen. nov. ; 13, Hulecoeteomyia stat. nov. ; 14, Gilesius gen. nov. ; 15, Collessius gen. nov. ; 16, Dahliana gen. nov. ; 17, Jarnellius gen. nov.

Figure 2

Strict consensus tree shown in Figure 1 with supporting, unambiguously optimized characters mapped on the branches. Circled numbers adjacent to terminal taxa correspond to the newly recognized genera indicated in Figure 1: 1, Pseudoskusea stat. nov. ; 2, Himalaius gen. nov. ; 3, Bruceharrisonius stat. nov. ; 4, Vansomerenis gen. nov. ; 5, Macleaya stat. nov. ; 6, Jihlienius gen. nov. ; 7, Molpemyia stat. nov. ; 8, Phagomyia stat. nov. ; 9, Rampamyia gen. nov. ; 10, Dobrotworskyius gen. nov. ; 11, Georgecraigius gen. nov. ; 12, Patmarksia gen. nov. ; 13, Hulecoeteomyia stat. nov. ; 14, Gilesius gen. nov. ; 15, Collessius gen. nov. ; 16, Dahliana gen. nov. ; 17, Jarnellius gen. nov.

Aedini is recovered as monophyletic and the relationships among the non-aedine outgroup taxa are the same as reported by Reinert et al. (2004): Culiseta+ (Culex+ (Orthopodomyia+ (Mansonia+ Aedini))). The first clade to branch off the main stem is Psorophora. Next is a group comprising Mucidus in a sister-group relationship with a clade that includes two subclades, one consisting of Geoskusea, Levua and Rhinoskusea (with the last two in a sister-group relationship) and the other comprising Pseudoskusea stat. nov. and Ochlerotatus (which includes the two subgenera Ochlerotatus and Rusticoidus, of which the former is paraphyletic relative to the latter). The next clade to branch off comprises Halaedes+Opifex, followed by Himalaius gen. nov. +Bruceharrisonius stat. nov. (all new taxa are described in Appendix 3). At the next level, a group separates that comprises Oc. (Fin.) candidoscutellum+‘Oc. (Fin.) keefei in a sister-group relationship with Vansomerenis gen. nov. + (Zavortinkius+Kenknightia). This is followed by a clade comprising six species of Oc. (Finlaya)’, and then Oc. (Pro.) zavortinki, which is sister to the remaining taxa. These are divided into two large clades. In the first of these, Oc. (Pro.) knabi is sister to the remaining taxa, which comprise three clades in an unresolved trichotomy (Fig. 1B). There are two small clades, the first comprising Oc. (Fin.) loi+Macleaya stat. nov. (including subgenus Chaetocruiomyia stat. nov.), and the second (Jihlienius gen. nov. +Molpemyia stat. nov.) +Phagomyia stat. nov. In the third, largest clade, Rampamyia gen. nov. is sister to the remaining taxa, which are divided into two almost equal subclades. The first comprises Dobrotworskyius gen. nov. + (Georgecraigius gen. nov. +Patmarksia gen. nov.), of which Georgecraigius includes subgenus Horsfallius subgen. nov. The second subclade comprises Hulecoeteomyia stat. nov. as the sister group of (Gilesius gen. nov. +Tanakaius) +Collessius gen. nov. In the second large clade (Figs 1C, 2C), there are two terminal clades comprising groups of genera: Dahliana gen. nov., Howardina and Finlaya+ (Downsiomyia+Haemagogus); and (Jarnellius gen. nov. + (Abraedes+ (Aztecaedes+ (Gymnometopa+Kompia)))) + ((Isoaedes+Aedes) + (Scutomyia+Stegomyia)). These genera and groups of genera are all interrelated through a set of relationships that also includes ten species of Oc. (Protomacleaya) (shaded in Fig. 1C) and three species of Oc. (Finlaya)’, i.e. oreophilus, harperi and albotaeniatus.

The ambiguity among the four MPCs is relatively minor and concerns just four parts of the cladograms, two of which are linked. The first involves the relationships among the terminal three species of Jarnellius, which are either varipalpus+ (deserticola+sierrensis) or sierrensis+ (deserticola+varipalpus). The second area of ambiguity concerns the relationships of Howardina, which can be placed as the sister group of either Dahliana or the clade comprising Oc. (Fin.) harperi, Oc. (Fin.) albotaeniatus, Finlaya, Downsiomyia and Haemagogus. The third area of ambiguity involves the relationships of Oc. (Fin.) loi+Macleaya and the four species of Phagomyia. When the former clade is placed as the sister group of (Jihlienius+Molpemyia) +Phagomyia, then the relationships among the species of Phagomyia are assamensis+ (lophoventralis+ (gubernatoris+prominens)). Alternatively, Oc. (Fin.) loi+Macleaya can be placed as the sister group of the larger clade comprising Rampamyia, Dobrotworskyius, Georgecraigius, Patmarksia, Hulecoeteomyia, Gilesius, Tanakaius and Collessius, whence the relationships among the species of Phagomyia are (gubernatoris+lophoventralis) + (assamensis+prominens).

COMPARISON WITH REINERT ET AL. (2004)

An exact comparison of the relationships portrayed in Figures 1 and 2 with those reported by Reinert et al. (2004: Fig. 4) is not possible because numerous taxa analysed in the latter study are not included in the present analysis, and vice versa. It is nevertheless revealing to compare the two patterns of relationships. First, and most importantly, all those taxa that Reinert et al. (2004) treated as genera, for which two or more species are included in the current analysis, are recovered here as monophyletic groups. Thus, the monophyly of the following genera is corroborated: Aedes, Downsiomyia, Geoskusea, Haemagogus, Kenknightia, Mucidus, Ochlerotatus, Opifex, Psorophora, Rhinoskusea, Scutomyia, Stegomyia and Zavortinkius. No genus proposed by Reinert et al. failed this test.

Second, a number of more inclusive groups, comprising two or more genera, are also common to both sets of results, often with identical or very similar patterns of interrelationships among their members, and which may eventually form the basis of a higher-level classification of the tribe. The clade Geoskusea+ (Levua+Rhinoskusea) is consistently recovered. A group comprising Downsiomyia+Haemagogus is also found in both studies, although in the present study the interrelationships among the four species of Haemagogus are slightly different in that both subgenera, Haemagogus and Conopostegus, are now recovered as monophyletic groups. Previously the latter subgenus was paraphyletic, although the clade Hg. (Con.) leucocelaenus+Hg. (Haemagogus) was supported by only a single character (71:0) and had very low branch support (Bremer support = 0.1). In the present study, four homoplastic characters (83:2, 107:1, 132:1, 213:1) now support Hg. (Conopostegus), but the Bremer support remains unchanged at 0.1. The more inclusive clade found by Reinert et al. (2004), comprising Oc. (Pro.) triseriatus, Oc. (Fin.) geniculatus, Downsiomyia and Haemagogus, is also recovered in the current study but now also includes Finlaya and Howardina. Previously, Finlaya was placed as the sister group of Molpemyia pecuniosus comb. nov. In the present study, this latter taxon is recovered as the sister group of Jihlienius and included in the other major distal clade from that which contains Finlaya. Reinert et al. (2004) placed Howardina as the sister group to a clade consisting of Abraedes, Aztecaedes, Gymnometopa, Kompia and Oc. (Och.) muelleri. This latter group is also recovered here, although its internal relationships are different. Furthermore, it is placed as the sister group to a clade consisting of representatives of the Aedes genus-group (i.e. Aedes in the previous broad sense; Section B of Belkin, 1962), namely Isoaedes, Aedes, Stegomyia and Scutomyia. These four genera (seven species) represent the much larger clade of 63 species in Reinert et al. (2004), but are, nevertheless, recovered as a monophyletic unit. Their internal relationships are slightly different, however, being (Isoaedes+Aedes) + (Stegomyia+Scutomyia) instead of Isoaedes+ (Aedes+ (Stegomyia+Scutomyia)), but this may be an artefact of the reduction in the number of taxa sampled from this group.

Another clade found by Reinert et al.(2004) comprised (‘Oc. (Fin.) chrysolineatus+‘Oc. (Fin.) macfarlanei) + (‘Oc. (Och.) atropalpus+ (Tanakaius+ (Halaedes+Opifex))). This group has been split up in the present analysis. The species atropalpus, chrysolineatus and macfarlanei are here placed, respectively, in Georgecraigius, Hulecoeteomyia and Collessius. All three, and Tanakaius, are still found together within a single large clade (Figs 1B, 2B), but the addition of numerous species of ‘Finlaya’ to this clade has changed their interrelationships into Georgecraigius+ (Hulecoeteomyia+(Tanakaius+Collessius)). Halaedes and Opifex are still sister groups but are now placed much nearer to the base of the cladogram (Figs 1A, 2A). The two other species included in the study of Reinert et al. (2004), namely Macleaya (Chaetocruiomyia) wattensis comb. nov. and Macleaya (Macleaya) tremulus comb. nov. form a sister-group pair in the same large clade that includes Collessius, Georgecraigius, Hulecoeteomyia and Tanakaius. This pair, together with Finlaya and Molpemyia, was placed in the MPCs of Reinert et al. (2004) as a paraphyletic series between the clade consisting of Oc. (Pro.) triseriatus+ (‘Oc. (Fin.) geniculatus+ (Downsiomyia+Haemagogus)) and that comprising GymnometopaAztecaedes and the Aedes genus-group. However, support for these groupings was low (0.1–0.2) and so changes following the addition of a large number of related taxa are not entirely unexpected.

The two large clades of the present study (Figs 1B, C, 2B, C) were also found in the MPCs of Reinert et al. (2004: fig. 4), although with the differences in internal relationships described above. Between these and the non-aedine outgroup taxa (which also show the same branching pattern as found previously) is a series of clades that Reinert et al. (2004) resolved into two monophyletic groups placed in a trichotomous relationship with the large clade. The first of these clades (with updated nomenclature) was Zavortinkius+ (Kenknightia+ (Bruceharrisonius+Psorophora)), and the second was ((Geoskusea+ (Levua+Rhinoskusea)) + (Mucidus+ (Pseudoskusea+Ochlerotatus)). This latter clade is recovered in the present analysis (Figs 1A, 2A), although Mucidus has moved to a position as the sister group of the two remaining clades. Of the others, Zavortinkius is now sister to Kenknightia in a clade that also includes Oc. (Fin) candidoscutellum, Oc. (Fin.) keefei and Vansomerenis. Bruceharrisonius is placed with a newly included species, Himalaius gilli comb. nov. in another clade, and Psorophora is a separate lineage that is the sister group of the remaining Aedini. Thus, these four clades no longer form two monophyletic groups but are arranged in a paraphyletic series (Figs 1A, 2A). Overall, however, despite the removal and inclusion of a large number of taxa, the results of Reinert et al. (2004) and those of the present study show remarkable congruence.

CLASSIFICATION OF EVALUATED TAXA

When Zavortink (1972) removed Protomacleaya from Aedes (Finlaya), he stated: ‘Although the subgenus is quite diverse, is defined mainly on negative characters, and many of the included species depart from the “normal” in 1 or more characteristics of 1 or more stages, it is, I believe, natural because of the basically similar male genitalia of virtually all species. However, the species of Oc. (Protomacleaya) included in the present analysis (shaded in Fig. 1) are arrayed in a pattern similar to what palaeontologists call a ‘stem group’, in contrast to a terminal, monophyletic ‘crown group’. A crown group is ‘an extant monophyletic taxon that also includes extinct phena with all the diagnostic attributes of that taxon (or can be shown to have secondarily lost such attributes)’, whereas the ‘total group is ‘an extant monophyletic taxon that contains all extinct phena possessing one or more of the diagnostic attributes of the taxon (Smith, 1994: 94–95). The stem group is that portion of the total group that is not included within the crown group and is thus paraphyletic. In the present study, only a paraphyletic group such as Oc. (Pro.) brelandi, Oc. (Pro.) hendersoni and Oc. (Pro.) triseriatus strictly meets the criteria for being a ‘stem group’, in this case the clade comprising Oc. (Pro.) brelandiKompia purpureipes (Fig. 1C); Oc. (Protomacleaya) in its entirety is polyphyletic. It is apparent, however, that many of the groups that consist of a species of Oc. (Protomacleaya) and its larger sister group are poorly diagnosed on short branches, and also that those species of Oc. (Protomacleaya) are poorly characterized in comparison with their sister groups (Fig. 2). Thus, we agree with Zavortink (1972) that the species of Oc. (Protomacleaya) do, in general, resemble each other phenetically, by virtue of what they lack. However, such a paraphyletic/polyphyletic assemblage is in no meaningful evolutionary sense ‘natural’. Ultimately, such ‘groups’ should be split up and classified into their constituent monophyletic parts. As an interim solution, the generic and subgeneric names of such assemblages should be placed in quotation marks to indicate their non-monophyletic status (the single quotation marks are used here merely as a matter of journal style). Therefore, knabi, for example, should be Ochlerotatus (‘Protomacleaya’) knabi or Ochlerotatus (Protomacleaya) knabi (abbreviated as Oc. (Pro.) knabi). This convention is adopted herein for all non-monophyletic taxa that must be retained for the time being.

Pseudoskusea is a small, homogeneous group comprising four species (bancroftianus, culiciformis, multiplex, postspiraculosus) in the Australasian Region. Theobald (1907) established Pseudoskusea as a monobasic genus for Skusea multiplex Theobald. Edwards (1932) classified Pseudoskusea as a subgenus of Aedes, where it remained until Reinert (2000a) transferred it to genus Ochlerotatus when he formally elevated this taxon from subgeneric rank within Aedes. The placement of Pseudoskusea as the sister group to Ochlerotatus s.s. in the preferred cladogram of Reinert et al. (2004: fig. 4) is corroborated by the results of the present analysis (Figs 1A, 2A). Whereas Reinert et al. (2004) were obliged to retain Pseudoskusea in the non-monophyletic Ochlerotatussensu auctorum because its sister-group relationship with Ochlerotatus was not weighting-independent (undoubtedly because it was represented only by a single species), we propose to restore this taxon to its original generic status based on the present results. Support for the sister-group relationship between Pseudoskusea and Ochlerotatus is strong (Bremer support > 0.4; relative Bremer support > 100), but the supporting characters are all homoplastic (Fig. 2A). Likewise, the characters that diagnose Pseudoskusea (10) and Ochlerotatus (9) are also homoplastic. As is usually the case with generic-level groups of Aedini, these genera are polythetic taxa, i.e. they are diagnosed not by unique characters but by unique combinations of characters. Hence, although the individual characters may be independently derived elsewhere, they do not occur in the same combination in any other group.

The sister-group relationship of Himalaius gilli and Bruceharrisonius is strongly supported (Bremer support > 0.4; relative Bremer support > 100) by five homoplastic characters (Figs 1A, 2A). However, we have chosen to recognize gilli and Bruceharrisonius as separate genera because the number of characters that distinguish them is similar to the number that separates other generic pairs, e.g. Zavortinkius and Kenknightia. Because Himalaius exhibits clear distinctions in all life stages, there seems to be no empirical basis for including gilli and Bruceharrisonius as subgenera of a single genus. They are distinct taxa that merit equivalent rank. Consequently, we have chosen to elevate Bruceharrisonius to generic status with the concomitant recognition of gilli as a separate genus. Reinert (2003) provided a full description and illustrations of Bruceharrisonius (as a subgenus of Ochlerotatus). Himalaius, which includes simlensis in addition to gilli, is described in Appendix 3.

The placement of Vansomerenis as the sister group of Zavortinkius+Kenknightia is well supported (Bremer support > 0.4; relative Bremer support > 100), and the monophyly of the former taxon is strongly supported (Bremer support > 0.4; relative Bremer support > 100) (Fig. 1A) by two characters, one of which (117:2, antennal pedicel without scales and/or setae) is unique and not contradicted (Fig. 2A). Additional morphological distinctions (not included in the analysis) of this group of Afrotropical species further corroborate its uniqueness and support its recognition as a distinct lineage within the Ochlerotatus genus-group. We adjudge this lineage to be a genus based on the strength of the character support and the principle of equivalent rank. The diagnostic and differential features that distinguish Vansomerenis are detailed in Appendix 3.

Although support for the clade consisting of Oc. (Fin.) candidoscutellum+‘Oc. (Fin.) keefei (Fig. 1A) is weak (Bremer support = 0.1; relative Bremer support = 3), morphological observations (data not included herein) suggest that this species pair is a natural relationship. Furthermore, morphological data suggest that this taxon may be related to the Oc. (Fin.) biocellatus, but there is no evidence in the four MPCs to unite these taxa (Figs 1A, 2A). The inclusion of Oc. (Fin.) crossi in the biocellatus - seoulensis clade raises additional questions. We carefully examined published descriptions and available material of all known species of ‘Finlaya’ and their apparent allies to determine which species might be closely related to Oc. (Fin.) crossi based on morphological correspondence. Lien (1967) stated that crossi‘is very closely related to suffusus’, the only member of the Suffusus Subgroup (Geniculatus Group) in the classification of Knight & Marks (1952). After examining adult and immature stages of Oc.suffusus, we concur with Lien (1967) and fail to see any unambiguous morphological evidence of a natural relationship between crossi and the other species of the biocellatus - seoulensis clade. Consequently, it will be necessary to include additional related species and/or character data in future studies to resolve the relationships of these species.

The first clade in the unresolved trichotomy (Figs 1B, 2B) is a bit of an enigma. Chaetocruiomyia+Macleaya is placed as sister to Oc. (Fin.) loi. However, support for this sister-group relationship is weak (Bremer support = 0.3; relative Bremer support = 14) and the two clades do not appear to be closely related based on an intuitive interpretation of morphological data that is not included in the analysis. The pairing of Chaetocruiomyia and Macleaya is strongly supported by 15 homoplastic characters (Bremer support > 0.4; relative Bremer support > 100). These two taxa were originally described as genera (Theobald, 1910, 1903, respectively), and later reduced to subgenera within ‘Aedes’ (Edwards, 1932). Despite the large number of morphological features that distinguish them, including a unique character among those that diagnose Chaetocruiomyia (115:3, interocular space with both broad and narrow scales), we propose to recognize these lineages as subgenera of genus Macleaya. Taylor (1929), Marks (1962, 1964), Dobrotworsky (1965) and Lee et al. (1982, 1984) all reported the close similarity and relationship of Chaetocruiomyia and Macleaya in adult and immature stages as well as biology and distribution in the Australasian Region. Additionally, Reinert (2002e) pointed out the close similarity of female genitalia of the two taxa. In addition to the diagnostic combinations of characters arrayed on the branches in Fig. 2B, salient differences between the adults, larvae and pupae of Chaetocruiomyia and Macleaya are listed in Marks (1962). In view of the many morphological differences that distinguish these taxa from Oc. (Fin.) loi, we regard the pairing of Oc. (Fin.) loi with genus Macleaya as an anomaly and retain it in the non-monophyletic assemblage of Oc. (Finlaya) pending further study.

Reinert (1993) resurrected Molpemyia from synonymy with Finlaya as a subgeneric category for three species of ‘Aedes’ (auridorsum, pecuniosus, purpureus), and Reinert (2000a) subsequently transferred the subgenus to Ochlerotatus. In the study of Reinert et al. (2004), this taxon, represented only by pecuniosus, was placed as sister to Fl. kochi + Fl. poicilia, with Macleaya tremula as sister to these three taxa. Whereas a relationship between Finlayas.s. and Macleaya is not evident in the present study, affinities that may exist between Molpemyia and ‘Finlaya’sensu auctorum within the trichotomy (Figs 1B, 2B) are insoluble with the current data set. Although Oc. (Fin.) chungi is strongly supported as sister to Molpemyia (Bremer support > 0.4; relative Bremer support > 100), morphological differences not included in the analysis indicate that chungi is not closely related to Molpemyia. Ochlerotatus (Fin.) chungi and Molpemyia are diagnosed by 16 and 19 homoplastic characters, respectively (Fig. 2B), which exceed the number of such characters that diagnose several traditionally accepted genera, i.e. Armigeres (7), Eretmapodites (15), Haemagogus (13), Heizmannia (8), Udaya (10) and Zeugnomyia (9) (Reinert et al., 2004: figs 5, 6). Therefore, the recognition of chungi and Molpemyia as generic-level taxa is certainly equipollent. For these reasons, in addition to restoring Molpemyia to its original generic status based on shared diagnostic features observed in the three included species (see Reinert, 1993), a new genus, Jihliensius, is proposed for chungi and two apparently related species (Appendix 3).

Phagomyia Theobald is resurrected from synonymy with Finlaya for the clade comprising assamensis, gubernatoris, lophoventralis and prominens (Figs 1B, 2B). This clade is strongly supported (Bremer support > 0.4; relative Bremer support > 100) by a combination of eight homoplastic characters. Of these, six (129:0, 131:1, 133:0, 135:0, 141:2 and 179:1) are common to all four MPCs, and the other two (10:1 and 166:0) appear only after the MPCs have been combined to form the SCT (see caveat on over-interpretation above). Phagomyia seems to be a homogeneous group that currently includes 16 species (see Appendix 4). The genus was proposed originally for three nominal species (Theobald, 1905), including gubernatoris and two species now placed in Ae. (Aedimorphus)’. Culex gubernatoris Giles is the type species of Phagomyia by subsequent designation (Edwards, 1932).

The monophyly of the clade comprising albilabris+notoscriptus is strongly supported by an extensive combination of 16 homoplastic characters (Bremer support > 0.4; relative Bremer support > 100) (Figs 1B, 2B), hence the reason we have chosen to establish genus Rampamyia for this lineage within the Ochlerotatus genus-group. In addition to the two species included in the analysis (Ra. albilabris and Ra. notoscripta), this taxon also includes the allied Ra. quinquelineata (see Appendix 4). These species occur in the Australasian Region. Belkin (1962) placed albilabris and notoscriptus in the Notoscriptus Subgroup of the Mediovittatus Group of ‘Finlaya’ (as a subgenus of ‘Aedes’) and noted that it ‘probably should be recognized as a distinct group’. Rampamyia, as defined here, includes a distinct part of Belkin's Notoscriptus Subgroup. This group is marked by the presence of a patch of broad, overlapping, silvery scales on the antennal pedicels (character 117:1), which only occurs elsewhere as independent derivations in the Stegomyia+Scutomyia and AbraedesKompia clades of the Aedes genus-group and Ochlerotatus genus-group, respectively.

Dobrotworskyius is proposed for the clade represented by alboannulatus, rubrithorax and tubbutiensis (Figs 1B, 2B). Although this genus is not as strongly supported as most other genera of Aedini (Bremer support = 0.3; relative Bremer support = 18), shared morphological characters not included in the analysis indicate that these species form a natural assemblage. The genus, which is confined to Australia, also includes milsoni, occidentalis, rupestris and subbasalis (Appendices 3 and 4).

Georgecraigius is proposed as a genus for three species that occur in the Nearctic (atropalpus, epactius), Neotropical (epactius, fluviatilis) and Palaearctic (atropalpus) regions. These species were classified as species of Aedes (Finlaya) until Zavortink (1972) assigned them to Ae. (Ochlerotatus) . Reinert (2000a) transferred them to Ochlerotatus (Ochlerotatus)’, and they were subsequently left without generic placement when Reinert et al. (2004) restricted the limits of Ochlerotatus. The monophyly of Georgecraigius is strongly supported (Bremer support > 0.4; relative Bremer support > 100) by a combination of 11 homoplastic characters (Fig. 2B). Nine homoplastic characters diagnose fluviatilis and place it in a sister-group relationship with atropalpus+epactius (Figs 1B, 2B). The atropalpus+epactius clade is diagnosed and distinguished from the fluviatilis clade by 20 homoplastic characters. [It is noteworthy that atropalpus and epactius were strongly paired (bootstrap value = 71) in the rDNA RFLP phylogeny of Kumar et al. (1998)]. In view of the number of characters that distinguish the two clades, we propose to recognize them as separate subgenera. The atropalpus+epactius clade is the nominotypical subgenus because atropalpus is the type species of Georgecraigius. The monobasic fluviatilis clade is named and described as Horsfallius subgen. nov. in Appendix 3.

Genus Patmarksia is established for Australasian species that, for the most part, were included in the Papuensis Subgroup of the Alboannulatus Group of Knight & Marks (1952). The sister-group relationship of Patmarksia with Georgecragius is strongly supported by seven homoplastic characters (Bremer support > 0.4; relative Bremer support > 100) (Figs 1B, 2B). Although monophyly of Patmarksia is rather weakly supported by a combination of only four homoplastic characters (Bremer support = 0.3; relative Bremer support = 34) (Figs 1B, 2B), it is further diagnosed and distinguished from Georgecragius by a number of distinctive features that were not included in the present data set (see description in Appendix 3). Relationships of the four species of Patmarksia included in the analysis indicate the possible existence of two species-groups or subgenera within the genus, represented by the pairing of argyronotum+papuensis and argenteitarsis+subalbitarsis. However, because only five homoplastic characters support the latter clade, further analyses that include several or all of the 13 species currently assigned to Patmarksia (see Appendix 4) are required to corroborate the reality of these groups.

Hulecoeteomyia Theobald is resurrected for the clade comprising japonicussherki (Figs 1B, 2B). Although the basal branch of the clade has only weak support (Bremer support = 0.2; relative Bremer support = 10), the group appears to be a natural assemblage of species that share a number of distinctive morphological features (Fig. 2B), especially the broad basal pale bands on hindtarsomeres 1–3 and the development and positions of larval setae 4–7-C. With the exception of japonicus, all species of Hulecoeteomyia included in the analysis exhibit character 95:0 (seta 6-II shorter than seta 7-II). Within Hulecoeteomyia, japonicus is sister to the remaining taxa, which comprise two groups in a pectinate relationship (Figs 1B, 2B). Because the relationships within Hulecoeteomyia are only weakly supported, it is not practical to propose internal subdivisions until additional character data are available.

Gilesius pulchriventer comb. nov. shares a number of characters with species of Hulecoeteomyia but the hindtarsomeres of the adults are dark-scaled and pupal seta 6-II, based on the pupa of Gi. alius, which Lien (1968b) originally described as a subspecies of pulchriventer, is significantly longer than seta 7-II. The sister-group relationship of pulchriventer and Tanakaius (Figs 1B, 2B) is strongly supported (Bremer support > 0.4; relative Bremer support > 100) by a single character of the male genitalia (218:1, gonocoxite with specialized scales on the mesal margin of the ventral surface) that otherwise occurs in the distantly related Oc. (Fin.) biocellatus and the clade comprising Oc. (Fin.) harperiHaemagogus. However, the many morphological differences that distinguish these taxa indicate that they are not so closely related. Gilesius is diagnosed by a combination of 12 homoplastic characters (Fig. 2B), which exceed the number of such characters that define four traditionally accepted genera, i.e. Armigeres (7), Heizmannia (8), Udaya (10) and Zeugnomyia (9) (Reinert et al., 2004: figs 5, 6). Shared morphological characters not included in the analysis also support the monophyly of the group (Appendix 3); thus, this clade is recognized as a genus based on the degree of morphological differentiation and the principle of equivalent rank.

Gilesius+Tanakaius shares a weakly supported sister-group relationship with hatorii+elsiaepseudotaeniatus (Bremer support = 0.2; relative Bremer support = 7), but the monophyly of the latter taxon is strongly supported (Bremer support > 0.4; relative Bremer support > 100) (Fig. 1B) by five homoplastic characters (Fig. 2B). Because, like Gilesius above, the monophyly of this clade is further supported by morphological data that are not included in the analysis, we have chosen to recognize it as a generic-level lineage, genus Collessius, within the Ochlerotatus genus-group. In addition to the five species included in the analysis, this taxon also includes four allied species (Appendix 3). Members of the genus, which occur in the Oriental Region and the Manchurian Subregion of the Palaearctic, exhibit morphological differences, principally in the male and female genitalia, indicative of subgroup affinities that only partially correspond with the pattern of relationships shown in the cladograms, i.e. hatorii+ ((elsiae+macfarlanei) + (banksi+pseudotaeniatus)). Although the characters that diagnose the elsiae+macfarlanei branch include a unique character (222:5, gonostylar claw with two narrow, short to moderately long spiniforms inserted distally) that is not contradicted (Fig. 2B), other anatomical characters suggest that hatorii, and three species not included in the analysis, are more closely allied to this species pair than to banksi and pseudotaeniatus. In view of the morphological contradictions, it is not possible to divide the genus into subgeneric groups; hence, two informal assemblages of species are recognized (Appendix 3) until the relationships of all species of Collessius are resolved.

The monophyly of echinus+geniculatus is strongly supported (Bremer support > 0.4; relative Bremer support > 100) by a combination of ten homoplastic characters (Figs 1C, 2C) and otherwise forms a distinct element within the Eurasian aedine fauna. Edwards (1932) included these species along with species from the Oriental Region and the New World in his Group H of Aedes (Finlaya) . Although similarities have been noted between geniculatus and Oc. (Protomacleaya) (e.g. Becker et al., 2003), the studies by Schick (1970) and Zavortink (1972) did not include or discuss this species in their revisions of species currently placed in Protomacleaya. In a biochemical systematics study that included geniculatus and Oc. (Pro.) triseriatus, Munstermann (1980) reported that ‘When enzyme migration rates of geniculatus and triseriatus were compared, only 7 of the 21 enzymes assayed migrated at similar rates, indicating that the two species are not closely related.’ The results of the present study corroborate this observation. Based on these observations, the strength of character support (Figs 1C, 2C) and other distinguishing features not included in the analysis, generic-level status is proposed for the echinus+geniculatus clade. This genus is formally named and described as Dahliana in Appendix 3.

The pairing of Oc. (Och.) muelleri with deserticola - varipalpus (in an unresolved relationship) is only supported (Bremer support = 0.4; relative Bremer support = 7) by a combination of six homoplastic characters (Fig. 2C), but it is strongly supported as sister to Abraedes+ (Aztecaedes+(Gymnometopa+Kompia)) (Bremer support > 0.4; relative Bremer support > 100) (Figs 1B, 2B). The three species that comprise the latter clade belong to the Varipalpus Group of five treehole-breeding species that occur in western North America (Arnell & Nielsen, 1972). Belkin & McDonald (1957) excluded muelleri from the Varipalpus Group (their varipalpus complex) and considered it ‘an aberrant member of the pulchritarsis-varipalpus group based on differences in the scutal ornamentation of the adults and the absence of teeth on the fore- and midungues of females. Zavortink (1972) pointed out similarities between the adult, larval and pupal stages and distributions of muelleri and members of the Varipalpus Group and placed it in a monobasic group (Muelleri Group) allied ‘with the Pulchritarsis and Varipalpus Groups in a more inclusive Varipalpus Section’. The results of the present study corroborate a close relationship between muelleri and the Varipalpus Group. In as much as the muellerivaripalpis lineage is tantamount to other clades that we regard as genera, we have chosen to identify this monophyletic group as genus Jarnellius. Because a combination of 16 homoplastic characters distinguishes muelleri as an independent lineage within the genus (Fig. 2C), Lewnielsenius subgen. nov. is proposed as a monobasic subgenus for this species. The genus and subgenus are formally recognized and described in Appendix 3.

FINAL COMMENTS

This is the second study of a four-phase project that aims objectively to assess the phylogenetic relationships and classification of Aedini. As with our previous study (Reinert et al., 2004), the results and proposals of the present investigation are likely to be viewed as controversial. In response to this, we unreservedly state that we regard our reclassification of the tribe as a developing empirical hypothesis, subject to change and revision as more information becomes available. As Zavortink (1979) aptly stated, ‘A classification, again like any other hypothesis, has heuristic value, so that its very existence should stimulate further research that may ultimately bring about its modification or demise. With this in mind, it is worth reiterating that the reclassification of species ‘into smaller and more numerous genera that indicate their genetic [recte genealogical] relationships more accurately is a consequence of beta-level taxonomic study; and in comparison with the classifications of other groups of organisms based on beta taxonomy, the total number of mosquito genera should be 225 (Zavortink, 1990). In proportion to the size of the family, we previously estimated that Aedini should include about 87 genera. As a result of our present and previous studies, 63 genera are currently recognized within the tribe. Again, we emphasize that this is a fundamental desideratum for achieving a natural classification of Aedini. Two points support this statement. First, the results of Reinert et al. (2004) and those of the present study show remarkable congruence despite the disjunction between the number of taxa and morphological characters included in the two studies. Second, we would note that the majority of subgenera recognized in the traditional classification of Aedini (Edwards, 1932; Belkin, 1962) are the same taxa that we recognize as polythetically diagnosed genera, the difference being that we choose to recognize explicitly monophyletic groups of generic rank rather than intuitively constructed groups of lower rank within ‘large paraphyletic or polyphyletic (and hence unnatural) genera.

Mitter (1999) appropriately stated: ‘Intelligent selection of the sample of species to be used for molecular study depends on the existence of a sound initial classification, which will almost always be based on morphology. Morphological classifications will also continue to serve as the initial standard for gauging which genes are suitable for solving a given problem.’ Our proposed classification based on a comparative morphological study provides such a framework.

ACKNOWLEDGEMENTS

We gratefully acknowledge Mark Carine (Department of Botany, The Natural History Museum (NHM), London, UK) and M. Anice M. Sallum (Núcleo de Pesquisa Taxonômica e Sistemática em Entomologia Médica, Faculdade de Saúde Pública, Universidade de São Paulo, Brazil), for critically reviewing the manuscript and providing valuable comments. Gratitude is expressed to Richard C. Wilkerson, James E. Pecor and Thomas V. Gaffigan (Walter Reed Biosystematics Unit, Smithsonian Institution, Washington, DC, USA), Theresa M. Howard (NHM), Desmond H. Foley and Greg Daniels (University of Queensland Insect Collection, Department of Zoology and Entomology, University of Queensland, Brisbane, Australia), Jacquie Recsei and Don Colless (Australian National Insect Collection, CSIRO Division of Entomology, Canberra, Australia), Richard F. Darsie, Jr, Florida Medical Entomology Laboratory, Vero Beach, FL, USA), Francis Schaffner (Laboratoire/Cellule Entomologie, EID Méditerranée, Montpellier, France), Gary J. Steck (Florida State Collection of Arthropods, Division of Plant Industry, Gainesville, FL, USA), Vu Duc Huong (National Institute of Malariology, Parasitology and Entomology, Hanoi, Vietnam) and Thomas J. Zavortink (Bohart Museum of Entomology, University of California, Davis, CA, USA) for the loan of specimens. Theresa Howard (NHM) is especially acknowledged for preparing the figures and making contributions to the species inventory. Appreciation is expressed to Kenneth J. Linthicum (CMAVE) for providing research facilities to J.F.R.

REFERENCES

Arnell
JH
1973
.
Mosquito studies (Diptera, Culicidae) XXXII. A revision of the genus Haemagogus
.
Contributions of the American Entomological Institute (Ann Arbor)
10
:
1
174
.

Arnell
JH
1976
.
Mosquito studies (Diptera, Culicidae) XXXIII. A revision of the scapularis group of Aedes (Ochlerotatus) [sic]
.
Contributions of the American Entomological Institute (Ann Arbor)
13
:
1
144
.

Arnell
JH
,
Nielsen
LT
1972
.
Mosquito studies (Diptera, Culicidae) XXVII. The varipalpus [sic] group of Aedes (Ochlerotatus)
.
Contributions of the American Entomological Institute (Ann Arbor)
8
:
1
48
.

Banez
LFL
,
Jueco
NI
1966
.
Adult and larva of Aedes (Finlaya) sorsogonensis [sic] Banez, 1963. Comparison with Aedes (Finlaya) ananae [sic] Knight and Laffoon, 1946 and Aedes (Finlaya) poecilus [sic] (Theobald), 1903
.
Acta Medica Philippina
2
(Series 2, No.
4
):
194
200
.

Barraud
PJ
1923
.
A revision of the culicine mosquitoes of India. Part VI. Some Indian species of the genus Finlaya Theo. adult stage
.
Indian Journal of Medical Research
11
:
475
493
+ plates XXII–XXXIII.

Barraud
PJ
1924
.
Four new mosquitoes from the western Himalayas
.
Indian Journal of Medical Research
11
:
999
1006
.

Barraud
PJ
1934
.
The fauna of British India, including Ceylon and Burma. Diptera. Vol. V. Family Culicidae Tribes Megarhinini and Culicini.
London
:
Taylor & Francis
.

Becker
N
,
Dusan
P
,
Zgomba
M
,
Boase
C
,
Dahl
C
,
Lane
J
,
Kaiser
A
2003
.
Mosquitoes and their control.
New York
:
Kluwer Academic
/
Plenum Publishers
.

Behbahani
A
,
Dutton
TJ
,
Davies
N
,
Townson
H
,
Sinkins
SP
2005
.
Population differentiation and Wolbachia phylogeny in mosquitoes of the Aedes scutellaris group
.
Medical and Veterinary Entomology
19
:
66
71
.

Belkin
JN
1962
.
The mosquitoes of the South Pacific (Diptera, Culicidae) [sic].
Vols
I & II
.
Berkeley
:
University of California Press
.

Belkin
JN
1968
.
Mosquito studies (Diptera, Culicidae) VII. The Culicidae of New Zealand
.
Contributions of the American Entomological Institute (Ann Arbor)
3
:
1
182
.

Belkin
JN
,
Heinemann
SJ
,
Page
WA
1970
.
Mosquito studies (Diptera, Culicidae) XXI. The Culicidae of Jamaica
.
Contributions of the American Entomological Institute (Ann Arbor)
6
:
1
458
.

Belkin
JN
,
McDonald
WA
1957
.
A new species of Aedes (Ochlerotatus) from tree holes in southern Arizona and a discussion of the varipalpus complex (Diptera: Culicidae)
.
Annals of the Entomological Society of America
50
:
179
191
.

Berlin
OGW
1969
.
Mosquito studies (Diptera, Culicidae) XII. A revision of the Neotropical subgenus Howardina [sic] of Aedes [sic]
.
Contributions of the American Entomological Institute (Ann Arbor)
4
:
1
190
.

Besansky
NJ
,
Fahey
GT
1997
.
Utility of the white gene in estimating phylogenetic relationships among mosquitoes (Diptera: Culicidae)
.
Molecular Biology and Evolution
14
:
442
454
.

Bohart
RM
1957
.
Insects of Micronesia, Diptera: Culicidae
.
Bernice P. Bishop Museum, Insects of Micronesia.
(1956)
12
:
1
85
.

Bohart
RM
,
Washino
RK
1978
.
Mosquitoes of California.
Berkeley
:
Division of Agricultural Sciences, University of California.

Bremer
K
1994
.
Branch support and tree stability
.
Cladistics
10
:
295
304
.

Buxton
PA
,
Hopkins
GHE
1925
.
The early stages of Samoan mosquitos [sic]
.
Bulletin of Entomological Research
15
:
295
301
.

Buxton
PA
,
Hopkins
GHE
1927
.
Researches in Polynesia and Melanesia, an account of investigations in Samoa, Tonga, the Ellice Group, and the New Hebrides, in 1924, 1925 Parts I–IV (relating principally to medical entomology).
Memoir Series Number 1.
London
:
The London School of Hygiene and Tropical Medicine.

Carpenter
SJ
,
LaCasse
WJ
1955
.
Mosquitoes of North America (north of Mexico)
.
Berkeley
:
University of California Press
.

Chadee
DD
,
Bennett
H
1990
.
Scanning electron microscopy of the eggs of Haemagogus celeste and Haemagogus equinus (Diptera: Culicidae) from Trinidad, West Indies
.
Mosquito Systematics
22
:
11
14
.

Clements
AN
1999
.
The biology of mosquitoes
, Vol.
2
. Sensory reception and behaviour.
Wallingford, UK
:
CAB International
.

Cook
S
,
Diallo
M
,
Sall
AA
,
Cooper
A
,
Holmes
EC
2005
.
Mitochondrial markers for molecular identification of Aedes mosquitoes (Diptera: Culicidae) involved in transmission of arboviral disease in West Africa
.
Journal of Medical Entomology
42
:
19
28
.

Dahl
C
1988
.
Taxonomic studies on Culex pipiens and C. torrentium
. In:
Service
MW
, ed.
Biosystematics of haematophagous insects.
Systematics Association Special Volume
37
:
149
175
.

Darsie
RF
Jr,
Pradhan
SP
,
Vaidya
RG
1992
.
Notes on the mosquitoes of Nepal. II. New species records from 1991 collections
.
Mosquito Systematics
24
:
23
28
.

Dobrotworsky
NV
1959
.
Notes on Australian mosquitoes (Diptera, Culicidae) [sic]. IV. Aedes alboannulatus [sic] complex in Victoria
.
Proceedings of the Linnean Society of New South Wales
84
:
131
145
.

Dobrotworsky
NV
1962
.
Notes on Australian mosquitoes (Diptera, Culicidae). VI. Five new Victorian species and a description of the larva of Aedes milsoni [sic] (Taylor)
.
Proceedings of the Linnean Society of New South Wales
87
:
291
302
.

Dobrotworsky
NV
1965
.
The mosquitoes of Victoria (Diptera, Culicidae) [sic].
Carlton, Victoria
:
Melbourne University Press
.

Edwards
FW
1922
.
A synopsis of adult Oriental culicine (including megarhinine and sabethine) mosquitoes. Part I
.
Indian Journal of Medical Research
10
:
249
293
.

Edwards
FW
1932
.
Genera insectorum. Diptera, fam. Culicidae. Fascicle 194.
Brussels
:
Desmet-Verteneuil
.

Edwards
FW
1941
.
Mosquitoes of the Ethiopian Region III.–Culicine Adults and Pupae.
London
:
British Museum (Natural History).

Forattini
OP
1965a
.
Entomologia Medica, 2. Culicini: Culex, Aedes e Psorophora.
Sao Paulo
:
Editora da Universidade de Sao Paulo
.

Forattini
OP
1965b
.
Culicidologia Medica, 3. Culicini: Haemagogus, Mansonia, Culiseta [sic]
.
Sabethini. Toxorhynchitini. Arboviroses. Filariose bancroftiana [sic]. Genetica. Taxonomico
.
Sao Paulo
:
Editora da Universidade de Sao Paulo
.

Forattini
OP
1996
.
Culicidologia Medica, 1. Principios gerais morfologia glossario taxonomico.
Sao Paulo
:
Editora da Universidade de Sao Paulo.

Gerry
BI
1932
.
Morphological studies of the female genitalia of Cuban mosquitoes
.
Annals of the Entomological Society of America
25
:
31
75
.

Goloboff
P
1997
.
Pee-Wee: a program for parsimony analysis under implied weights
. Program and documentation, available at ftp.unt.edu.ar/pub/parsimony.

Goloboff
PA
,
Farris
JS
2001
.
Methods for quick consensus estimation
.
Cladistics
17
:
S26
S34
.

Gutsevich
AV
1974a
.
The determination of mosquito females by microscopic preparations of the head I. Taxonomic characters and description of the genera
.
Mosquito Systematics
6
:
243
250
(originally published in Russian in 1972, Parazitologiya, 6: 320–325).

Gutsevich
AV
1974b
.
The determination of mosquito females by microscopic preparations of the head II. Key to genera and subgenera
.
Mosquito Systematics
6: 251–258 (originally published in Russian in 1973, Parazitologiya,
7
:
106
110
).

Gutsevich
AV
1975a
.
The determination of mosquito females (Culicidae) by microscopic preparations of the head. III. A key to species (excluding Aedes)
.
Mosquito Systematics
7
:
153
163
(originally published in Russian in 1973, Parazitologiya, 7:443–449).

Gutsevich
AV
1975b
.
The determination of mosquito females (Culicidae) by microscopic preparations of the head. IV. A key to species of genus Aedes
.
Mosquito Systematics
7: 164–173 (originally published in Russian in 1974, Parazitologiya,
8
:
329
335
).

Gutsevich
AV
,
Monchadskii
AS
,
Shtakel'berg
AA
1974
.
Fauna of the U.S.S.R. Diptera. Mosquitoes Family Culicidae.
Jerusalem
:
Israel Program for Scientific Translations
(translated from Russian).

Harbach
RE
,
Kitching
IJ
1998
.
Phylogeny and classification of the Culicidae (Diptera)
.
Systematic Entomology
23
:
327
370
.

Harbach
RE
,
Knight
KL
1980
.
Taxonomists' glossary of mosquito anatomy
.
Marlton, New Jersey
:
Plexus Publishing
.

Harbach
RE
,
Knight
KL
1982
.
Corrections and additions to Taxonomists' Glossary of Mosquito Anatomy
.
Mosquito Systematics
. (1981)
13
:
201
217
.

Hinton
HE
1981
.
Biology of insect eggs in three volumes
, Vol.
I
.
Oxford
:
Pergamon Press
.

Hochman
RH
,
Reinert
JF
1974
.
Undescribed setae in larvae of Culicidae (Diptera)
.
Mosquito Systematics
6
:
1
10
.

Hopkins
GHE
1952
.
Mosquitoes of the Ethiopian Region I–Larval bionomics of mosquitoes and taxonomy of Culicine larvae
.
London
:
British Museum (Natural History)
.

Howard
LO
,
Dyar
HG
,
Knab
F
1913
.
The mosquitoes of North and Central America and the West Indies. A general consideration of mosquitoes, their habits, and their relations to the human species
.
(1912) Publication Number 159
, Vol.
1
.
Baltimore
:
Carnegie Institution of Washington, The Lord Baltimore Press
.

Howard
LO
,
Dyar
HG
,
Knab
F
1917
.
The mosquitoes of North and Central America and the West Indies. A general consideration of mosquitoes, their habits, and their relations to the human species
. (1912) Publication Number 159, Vol.
4
.
Baltimore
:
Carnegie Institution of Washington, The Lord Baltimore Press
.

Huang
Y-M
1979
.
Medical entomology studies - XI. The subgenus Stegomyia of Aedes in the Oriental Region with keys to the species (Diptera: Culicidae)
.
Contributions of the American Entomological Institute (Ann Arbor)
15
:
1
79
.

Huang
Y-M
1990
.
The subgenus Stegomyia of Aedes in the Afrotropical Region I. The africanus group of species (Diptera: Culicidae)
.
Contributions of the American Entomological Institute (Gainesville)
26
:
1
90
.

Huang
Y-M
1997
.
The subgenus Stegomyia of Aedes in the Afrotropical Region. II. The dendrophilus group of species (Diptera: Culicidae)
.
Contributions of the American Entomological Institute (Gainesville)
29
:
1
104
.

Kalpage
KS
,
Brust
RA
1968
.
Mosquitoes of Manitoba. I. Descriptions and a key to Aedes eggs (Diptera: Culicidae)
.
Canadian Journal of Zoology
46
:
699
718
.

Kaur
R
1992
.
Mosquito types present in the collections of National Institute of Communicable Diseases, Delhi (Diptera: Culicidae)
.
Oriental Insects
26
:
367
382
.

King
WV
,
Hoogstraal
H
1946
.
Species of Aedes (Finlaya) [sic] of the papuensis group in the Australasian Region (Diptera, Culicidae)
.
Proceedings of the Entomological Society of Washington
48
:
135
157
.

Knight
KL
1948
.
The Aedes (Finlaya) albotaeniatus [sic] group of mosquitoes (Diptera, Culicidae)
.
Proceedings of the Entomological Society of Washington
50
:
1
8
.

Knight
KL
1968
.
Contributions to the mosquito fauna of Southeast Asia.–IV. Species of the subgroup chrysolineatus of group D, genus Aedes, subgenus Finlaya Theobald
.
Contributions of the American Entomological Institute (Ann Arbor)
2
:
1
45
.

Knight
KL
,
Harrison
BA
1988
.
A new Aedes (Finlaya) of the niveus-subgroup
.
Mosquito Systematics
. (1987)
19
:
212
236
.

Knight
KL
,
Hull
WB
1951
.
The Aedes mosquitoes of the Philippine Islands I. Keys to species, subgenera Mucidus, Ochlerotatus, and Finlaya (Diptera, Culicidae)
.
Pacific Science
5
:
211
251
.

Knight
KL
,
Laffoon
JL
1971
.
A mosquito taxonomic glossary VIII. The larval chaetotaxy
.
Mosquito Systematics Newsletter
3
:
160
194
.

Knight
KL
,
Marks
EN
1952
.
An annotated checklist of the mosquitoes of the subgenus Finlaya [sic], genus Aedes [sic]
.
Proceedings of the United States National Museum
101
:
513
574
.

Kumar
A
,
Black
WCIV
,
Rai
KS
1998
.
An estimate of phylogenetic relationships among culicine mosquitoes using a restriction map of the rDNA cistron
.
Insect Molecular Biology
7
:
367
373
.

Lee
DJ
,
Hicks
MM
,
Griffiths
M
,
Russell
RC
,
Marks
EN
1982
.
The Culicidae of the Australasian region, Vol. II. Nomenclature, synonymy, literature, distribution, biology and relation to disease. Genera AEDEOMYIA, genus AEDES(subgenera [Aedes], Aedimorphus, Chaetocruiomyia, Christophersiomyia, Edwardsaedes and Finlaya)
Monograph Series. Entomology Monograph Number 2.
Canberra
:
School of Public Health and Tropical Medicine, Australian Government Publishing Service
.

Lee
DJ
,
Hicks
MM
,
Griffiths
M
,
Russell
RC
,
Marks
EN
1984
.
The Culicidae of the Australasian region, Vol. III. Nomenclature, synonymy, literature, distribution, biology and relation to disease. Genus AEDES subgenera Geoskusea, Halaedes, Huaedes, Leptosomatomyia, Levua, Lorrainea, Macleaya, Mucidus, Neomelaniconion, Nothoskusea, Ochlerotatus, Paraedes, Pseudoskusea, Rhinoskusea
Monograph Series. Entomology Monograph Number 2.
Canberra
:
School of Public Health and Tropical Medicine, Australian Government Publishing Service
.

Lien
JC
1967
.
New species of mosquitoes from Taiwan (Diptera: Culicidae) Part I. Two new species of Aedes (Finlaya) [sic]
.
Tropical Medicine
9
:
177
185
.

Lien
JC
1968a
.
New species of mosquitoes from Taiwan (Diptera: Culicidae) Part III. Five new species of Aedes
.
Tropical Medicine
10
:
95
115
.

Lien
JC
1968b
.
New species of mosquitoes from Taiwan (Diptera: Culicidae) Part V. Three new subspecies of Aedes and seven new species of Culex
.
Tropical Medicine
10
:
217
262
.

Linley
JR
1989
.
Egg of Mansonia dyari described and compared with egg of Mansonia titillans (Diptera: Culicidae)
.
Journal of Medical Entomology
26
:
41
45
.

Linley
JR
,
Chadee
DD
1990
.
Fine structure of the eggs of Psorophora columbiae, Ps. cingulata and Ps. ferox (Diptera: Culicidae)
.
Proceedings of the Entomological Society of Washington
92
:
497
511
.

Linley
JR
,
Chadee
DD
1991
.
Fine structure of the eggs of Haemagogus equinus and Hg. janthinomys (Diptera: Culicidae)
.
Journal of Medical Entomology
28
:
434
445
.

Linley
JR
,
Craig
GB
Jr.
1993
.
The egg of Aedes hendersoni and a comparison of its structure with the egg of Aedes triseriatus (Diptera: Culicidae)
.
Mosquito Systematics
25
:
65
76
.

Linley
JR
,
Craig
GB
Jr.
1994
.
Morphology of long- and short-day eggs of Aedes atropalpus and A. epactius (Diptera: Culicidae)
.
Journal of Medical Entomology
31
:
855
867
.

Linley
JR
,
Geary
MJ
,
Russell
RC
1991a
.
The eggs of Aedes (Finlaya) alboannulatus and Aedes (Finlaya) rubrithorax (Diptera: Culicidae)
.
Mosquito Systematics
23
:
132
143
.

Linley
JR
,
Geary
MJ
,
Russell
RC
1991b
.
The eggs of Aedes funereus, Aedes notoscriptus and Aedes alternans (Diptera: Culicidae)
.
Proceedings of the Entomological Society of Washington
93
:
592
612
.

Linley
JR
,
Linley
PA
,
Lounibos
LP
1986
.
Light and scanning electron microscopy of the egg of Mansonia titillans (Diptera: Culicidae)
.
Journal of Medical Entomology
23
:
99
104
.

Lu
B
,
Ji
S
1997
.
Subgenus Finlaya Theobald, 1903
. In:
Lu
B
,
Li
B
,
Ji
S
,
Chen
H
,
Meng
Q
,
Su
L
,
Qu
F
,
Gong
Z
,
Zhang
Z
, eds.
Diptera: Culicidae 1 Fauna Sinica, Insecta
, Vol.
8
.
Beijing
:
Science Press
99
173
, figs 18–60 (in Chinese).

Macdonald
WW
,
Traub
R
1960
.
Malaysian parasites XXXVII. An introduction to the ecology of the mosquitoes of the lowland Dipterocarp forest of Selangor, Malaya
. In:
Macdonald
WW
, ed
Malaysian Parasites XXXV–XLIX.
Studies Number 29.
Kuala Lumpur
:
Institute for Medical Research of the Federation of Malaya
,
79
109
.

Marks
EN
1962
.
A revision of the subgenus Chaetocruiomyia Theobald (Diptera: Culicidae)
.
Department of Entomology, University of Queensland Press
1
:
189
211
.

Marks
EN
1964
.
Notes on the subgenus Chaetocruiomyia Theobald (Diptera: Culicidae)
.
Proceedings of the Linnean Society of New South Wales
89
:
131
147
.

Marks
EN
,
Hodgkin
EP
1958
.
A new species of Aedes (Finlaya) from northern Australia (Diptera, Culicidae)
.
Proceedings of the Linnean Society of New South Wales
83
:
33
39
.

Marshall
JF
1938
.
The British mosquitoes
.
London
:
British Museum (Natural History).

Matsuo
K
,
Lien
JC
,
Yoshida
Y
1974a
.
Scanning electron microscopy of mosquitoes II. The egg surface structure of 13 species of Aedes from Taiwan
.
Journal of Medical Entomology
11
:
179
188
.

Matsuo
K
,
Lien
JC
,
Yoshida
Y
1974b
.
Scanning electron microscopy of mosquitoes Part III. The egg surface structure of 5 species from Taiwan and 2 species from Indonesia
.
Journal of Formosan Medical Association
73
:
437
444
.

Mattingly
PF
1957
.
The Culicine mosquitoes of the Indomalayan area. Part I: Genus Ficalbia Theobald.
London
:
British Museum (Natural History).

Mattingly
PF
1958
.
The culicine mosquitoes of the Indomalayan area. Part III. Genus Aedes Meigen, subgenera Paraedes Edwards, Rhinoskusea Edwards and Cancraedes Edwards.
London
:
British Museum (Natural History)
.

Mattingly
PF
1959
.
The culicine mosquitoes of the Indomalayan area. Part IV. Genus Aedes Meigen, subgenera Skusea Theobald, Diceromyia Theobald, Geoskusea Edwards and Christophersiomyia Barraud.
London
:
British Museum (Natural History)
.

Mattingly
PF
1965
.
The culicine mosquitoes of the Indomalayan area. Part IV. Genus Aedes Meigen, subgenus Stegomyia (Groups A, B and D).
London
:
British Museum (Natural History)
.

Mattingly
PF
1969
.
The biology of mosquito-borne disease
. The Science of Biology Series Number 1.
London
:
George, Allen & Unwin.

Mattingly
PF
1970
.
Mosquito eggs IX. Genus Opifex Hutton
.
Mosquito Systematics Newsletter
2
:
92
97
.

Mattingly
PF
1972
.
Mosquito eggs XXI. Genus Culiseta Felt
.
Mosquito Systematics
4
:
114
127
.

Mitter
C
1999
.
Sketches of U.S. systematic entomology, circa 1850–2000: return of a golden age?
Annals of the Entomological Society of America
92
:
798
811
.

Moriya
K
,
Yabe
T
,
Harada
F
1973
.
Chorionic markings of some aedine mosquitoes in Japan I. Preliminary observations by a scanning electron microscope and a reflected lighting microscope
.
Japanese Journal of Sanitary Zoology
24
:
47
55
.

Munstermann
LE
1980
.
Speciation in northern Aedes tree hole mosquitoes
. In:
Proceedings of XVI International Congress of Entomology.
Kyoto
,
Japan
11-S-2
,
13
,
321
.

Myers
CM
1967
.
Identification and descriptions of Aedes eggs from California and Nevada (Diptera: Culicidae)
.
The Canadian Entomologist
99
:
975
806
.

Penn
GH
1949
.
The pupae of the mosquitoes of New Guinea
.
Pacific Science
3
:
3
85
.

Pratt
HD
,
Kidwell
AS
1969
.
Eggs of mosquitoes found in Aedes aegypti oviposition traps
.
Mosquito News
29
:
545
548
.

Rees
DM
,
Onishi
K
1951
.
Morphology of the terminalia and internal reproductive organs, and copulation in the mosquito, Culiseta inornata [sic] (Williston)
.
Proceedings of the Entomological Society of Washington
53
:
233
246
.

Reinert
JF
1976a
.
A ventromedian cervical sclerite of mosquito larvae (Diptera: Culicidae)
.
Mosquito Systematics
8
:
205
208
.

Reinert
JF
1976b
.
Medical entomology studies - V. The subgenus Rhinoskusea of the genus Aedes (Diptera: Culicidae)
.
Contributions of the American Entomological Institute (Ann Arbor)
13
:
1
60
.

Reinert
JF
1979
.
A description of Isoaedes, a new subgenus of Aedes Meigen, and its type-species, Ae. (Isa.) cavaticus new species (Diptera: Culicidae)
.
Mosquito Systematics
11
:
144
162
.

Reinert
JF
1980
.
A literature review of pupal seta 13-CT in the family Culicidae (Diptera) and a report of its occurrence in Aedes aurotaeniatus Edwards
.
Mosquito Systematics
12
:
441
444
.

Reinert
JF
1990
.
Medical entomology studies -XVII. Biosystematics of Kenknightia, a new subgenus of the mosquito genus Aedes Meigen from the Oriental Region (Diptera: Culicidae)
.
Contributions of the American Entomological Institute (Gainesville)
26
:
1
119
.

Reinert
JF
1993
.
Redescription of Molpemyia, and its revalidation as a subgenus of Aedes (Diptera: Culicidae)
.
Mosquito Systematics
25
:
41
63
.

Reinert
JF
1999a
.
The subgenus Rusticoidus of genus Aedes (Diptera: Culicidae) in Europe and Asia
.
European Mosquito Bulletin
4
:
1
7
.

Reinert
JF
1999b
.
Descriptions of Zavortinkius, a new subgenus of Aedes, and the eleven included species from the Afrotropical Region (Diptera: Culicidae)
.
Contributions of the American Entomological Institute (Gainesville)
31
:
1
105
.

Reinert
JF
2000a
.
New classification for the composite genus Aedes (Diptera: Culicidae: Aedini), elevation of subgenus Ochlerotatus to generic rank, reclassification of the other subgenera, and notes on certain subgenera and species
.
Journal of the American Mosquito Control Association
16
:
175
188
.

Reinert
JF
2000b
.
Comparative anatomy of the female genitalia of genera and subgenera in tribe Aedini (Diptera: Culicidae). Part I. Introduction, preparation techniques, and anatomical terminology
.
Contributions of the American Entomological Institute (Gainesville)
32
:
1
18
.

Reinert
JF
2000c
.
Comparative anatomy of the female genitalia of genera and subgenera in tribe Aedini (Diptera: Culicidae). Part II. Genus Psorophora Robineau-Desvoidy
.
Contributions of the American Entomological Institute (Gainesville)
32
:
19
35
.

Reinert
JF
2000d
.
Comparative anatomy of the female genitalia of genera and subgenera in tribe Aedini (Diptera: Culicidae). Part III. Genus Udaya Thurman
.
Contributions of the American Entomological Institute (Gainesville)
32
:
37
44
.

Reinert
JF
2000e
.
Comparative anatomy of the female genitalia of genera and subgenera in tribe Aedini (Diptera: Culicidae). Part IV. Genus Zeugnomyia Leicester
.
Contributions of the American Entomological Institute (Gainesville)
32
:
45
52
.

Reinert
JF
2000f
.
Comparative anatomy of the female genitalia of genera and subgenera in tribe Aedini (Diptera: Culicidae). Part V. Genus Aedes Meigen
.
Contributions of the American Entomological Institute (Gainesville)
32
:
1
102
.

Reinert
JF
2001a
.
Revised list of abbreviations for genera and subgenera of Culicidae (Diptera) and notes on generic and subgeneric changes
.
Journal of the American Mosquito Control Association
17
:
51
55
.

Reinert
JF
2001b
.
Comparative anatomy of the female genitalia of genera and subgenera in tribe Aedini (Diptera: Culicidae). Part VI. Genus Ayurakitia Thurman
.
Contributions of the American Entomological Institute (Gainesville)
32
:
1
7
.

Reinert
JF
2001c
.
Comparative anatomy of the female genitalia of genera and subgenera in tribe Aedini (Diptera: Culicidae). Part VII. Genus Opifex Hutton
.
Contributions of the American Entomological Institute (Gainesville)
32
:
9
15
.

Reinert
JF
2001d
.
Comparative anatomy of the female genitalia of genera and subgenera in tribe Aedini (Diptera: Culicidae). Part VIII. Genus Verrallina Theobald
.
Contributions of the American Entomological Institute (Gainesville)
32
:
17
39
.

Reinert
JF
2001e
.
Comparative anatomy of the female genitalia of genera and subgenera in tribe Aedini (Diptera: Culicidae). Part IX. Genus Eretmapodites Theobald
.
Contributions of the American Entomological Institute (Gainesville)
32
:
41
50
.

Reinert
JF
2002a
.
A comparative analysis of the ventral brush and its attachment to segment X of fourth-stage larvae in tribe Aedini (Diptera: Culicidae)
.
Journal of the American Mosquito Control Association
18
:
16
25
.

Reinert
JF
2002b
.
Comparative anatomy of the female genitalia of genera and subgenera in tribe Aedini (Diptera: Culicidae). Part X. Genus Heizmannia Ludlow
.
Contributions of the American Entomological Institute (Gainesville)
32
:
1
15
.

Reinert
JF
2002c
.
Comparative anatomy of the female genitalia of genera and subgenera in tribe Aedini (Diptera: Culicidae). Part XI. Genus Haemagogus Williston
.
Contributions of the American Entomological Institute (Gainesville)
32
:
17
30
.

Reinert
JF
2002d
.
Comparative anatomy of the female genitalia of genera and subgenera in tribe Aedini (Diptera: Culicidae). Part XII. Genus Armigeres Theobald
.
Contributions of the American Entomological Institute (Gainesville)
32
:
31
46
.

Reinert
JF
2002e
.
Comparative anatomy of the female genitalia of genera and subgenera in tribe Aedini (Diptera: Culicidae). Part XIII. Genus Ochlerotatus Lynch Arribalzaga
.
Contributions of the American Entomological Institute (Gainesville)
33
:
1
111
.

Reinert
JF
2003
.
Description of Bruceharrisonius, a new subgenus of Ochlerotatus, and a redescription of its type species Oc. (Brh) greenii
.
Journal of the American Mosquito Control Association
19
:
309
322
.

Reinert
JF
2005
.
List of species described in the egg stage of tribe Aedini (Diptera: Culicidae) with their literature citations
.
Journal of the American Mosquito Control Association
21
:
252
262
.

Reinert
JF
,
Harbach
RE
,
Kitching
IJ
2004
.
Phylogeny and classification of Aedini (Diptera: Culicidae) based on morphological characters of all life stages
.
Zoological Journal of the Linnean Society
142
:
289
368
.

Ronderos
RA
,
Bachmann
AO
1963
.
Mansoniini [sic] neotropicales. I. (Diptera-Culicidae) [sic]
.
Revista de la Sociedad Entomologia Argentina
26
:
57
65
.

Ross
HH
1947
.
The mosquitoes of Illinois (Diptera, Culicidae)
.
Bulletin of the Illinois Natural History Survey (Urbana, Illinois)
24
:
1
96
.

Ross
HH
,
Horsfall
WR
1965
.
A synopsis of the mosquitoes of Illinois (Diptera, Culicidae)
.
Illinois Natural History Survey Biological Notes, Number
52
:
1
50
.

Russell
RC
1996
.
A colour photo atlas of mosquitoes of southeastern Australia.
Sydney
:
Published by the author.

Russell
PF
,
West
LS
,
Manwell
RD
,
Macdonald
G
1963
.
Practical malariology
, 2nd edn.
London
:
Oxford University Press
.

Schick
RX
1970
.
Mosquito studies (Diptera, Culicidae) XX. The Terrens Group of Aedes (Finlaya)
.
Contributions of the American Entomological Institute (Ann Arbor)
5
:
1
158
.

Sirivanakarn
S
1976
.
Medical entomology studies - III. A revision of the subgenus Culex in the Oriental Region (Diptera: Culicidae)
.
Contributions of the American Entomological Institute (Ann Arbor)
12
:
1
272
.

Smith
AB
1994
.
Systematics and the fossil record: documenting evolutionary patterns.
Oxford
:
Blackwell Scientific Publications
.

Tanaka
K
2002
.
Studies on the pupal mosquitoes of Japan (6) Aedes (Finlaya) (Diptera, Culicidae)
.
Japanese Journal of Systematic Entomology
8
:
137
177
.

Tanaka
K
,
Mizusawa
K
,
Saugstad
ES
1979
.
A revision of the adult and larval mosquitoes of Japan (including the Ryukyu Archipelago and the Ogasawara Islands) and Korea (Diptera: Culicidae)
.
Contributions of the American Entomological Institute (Ann Arbor)
16
:
1
987
.

Taylor
FH
1929
.
Notes on Australian Culicidae (Dipt.)
.
Bulletin of Entomological Research
20
:
271
277
.

Theobald
FV
1903
.
Two new Australian culicids
.
The Entomologist
36
:
154
157
.

Theobald
FV
1905
.
Genera insectorum. Diptera, fam. Culicidae
. Fascicle 26,
Belgium
.

Theobald
FV
1907
.
A monograph of the Culicidae or mosquitoes. Mainly compiled from collections received at the British Museum
, Vol.
IV
.
London
:
British Museum (Natural History)
.

Theobald
FV
1910
.
A monograph of the Culicidae or mosquitoes. Mainly compiled from collections received at the British Museum
, Vol.
V
.
London
:
British Museum (Natural History)
.

Tyson
WH
1970
.
Contributions to the mosquito fauna of Southeast Asia VIII. Genus Aedes, subgenus Mucidus Theobald in Southeast Asia
.
Contributions of the American Entomological Institute (Ann Arbor)
6
:
28
80
.

Villa
AS-C
,
Rodriguez
JAR
,
Llera
JLG
1985
.
Estudio de la larva de cuarto estado de las especies del subgenero Finlaya (Diptera: Culicidae) [sic] de Espana peninsular
.
Sociedade Portuguesa de Entomologia Suplemento
1
:
2
12
.

Wesson
DW
,
Porter
CH
,
Collins
FH
1992
.
Sequence and secondary structure comparisons of ITS rDNA in mosquitoes (Diptera: Culicidae)
.
Molecular Phylogenetics and Evolution
1
:
253
269
.

Wood
DM
,
Dang
PT
,
Ellis
RA
1979
.
The insects and arachnids of Canada Part 6. The mosquitoes of Canada, Diptera: Culicidae.
Publication 1686.
Quebec
:
Canadian Government Printing Centre.

Yamaguti
S
,
LaCasse
WJ
1951
.
Mosquito fauna of North America Part V - Genus Aedes [sic](with 116 original plates).
Office of the Surgeon, HQ. Japan Logistical Command, APO 343.

Zavortink
TJ
1972
.
Mosquito studies (Diptera, Culicidae) XXVIII. The New World species formerly placed in Aedes (Finlaya)
.
Contributions of the American Entomological Institute (Ann Arbor)
8
:
1
206
.

Zavortink
TJ
1979
.
Mosquito studies (Diptera, Culicidae) XXXV. The sabethine genus Johnbelkinia and a preliminary reclassification of the composite genus Trichoprosopon
.
Contributions of the American Entomological Institute (Ann Arbor)
17
:
1
61
.

Zavortink
TJ
1985
.
Observations on the ecology of treeholes and treehole mosquitoes in the southwestern United States
. In:
Lounibos
LP
,
Rey
JR
,
Frank
JH
, eds.
Ecology of mosquitoes
. Proceedings of a Workshop Florida Medical Entomology Laboratory,
Vero Beach, Florida
,
473
487
.

Zavortink
TJ
1990
.
Classical taxonomy of mosquitoes - a memorial to John N. Belkin
.
Journal of the American Mosquito Control Association
6
:
593
599
.

Appendices

APPENDIX 1

List of species examined in detail during the comparative morphological analysis. An asterisk (*) following a species–author combination denotes a type species of a generic-level taxon. Geographical distribution of species is indicated by region: 1, Nearctic; 2, Palaearctic; 3, Afrotropical; 4, Oriental; 5, Australasian; 6, Neotropical. An asterisk following a geographical region indicates a relatively recent introduction.

GenusSubgenusSpeciesDistribution
Aedine ingroup taxa 
Downsiomyia Vargas  leonis (Colless) 
  nivea (Ludlow)* 
Finlaya Theobald  kochi (Doenitz)* 
  poicilia (Theobald) 4,5 
Haemagogus Conopostegus Dyar leucocelaenus (Dyar & Shannon)* 
Williston  leucotaeniatus (Komp) 
 Haemagogus equinus Theobald 
  splendens Williston* 
Halaedes Belkin  australis (Erichson)* 
Ochlerotatus Lynch    
Arribalizaga ‘Finlaya’ Theobald albilabris (Edwards) 
  alboannulatus (Macquart) 
  albocinctus (Barraud) 
  albotaeniatus (Leicester) 
  argenteitarsis (Brug) 
  argyronotum (Belkin) 
  assamensis (Theobald) 
  banksi (Edwards) 
  biocellatus (Taylor) 
  candidoscutellum (Marks) 
  chrysolineatus (Theobald) 
  chungi (Lien) 
  crossi (Lien) 
  echinus Edwards 
  elsiae (Barraud) 
  embuensis (Edwards) 
  geniculatus (Olivier) 
  gilli (Barraud) 
  gubernatoris (Giles) 
  harperi (Knight) 
  hatorii (Yamada) 2,4 
  ingrami (Edwards) 
  j. japonicus (Theobald) 1*,2*,4 
  jugraensis (Leicester) 
  keefei (King & Hoogstraal) 
  loi (Lien) 
  lophoventralis (Theobald) 
  luteostriatus (Robinson) 
  macfarlanei Edwards 
  notoscriptus (Skuse) 
  oreophilus Edwards 2,4 
  papuensis (Taylor) 
  prominens (Barraud) 
  pseudotaeniatus (Giles) 
  pulchritorax (Edwards) 
  pulchriventer (Giles) 2,4 
  rubrithorax (Macquart) 
  seoulensis (Yamada) 
  sherki (Knight) 
  subalbitarsis (King & Hoogstraal) 
  tubbutiensis (Dobrotworsky) 
 Macleaya Theobald tremulus (Theobald)* 
 Molpemyia Theobald pecuniosus (Edwards) 
 Ochlerotatus’ atropalpus (Coquillett) 1,2* 
  deserticola (Zavortink) 
  epactius (Dyar & Knab) 1,6 
  fluviatilis (Lutz) 
  muelleri (Dyar) 
  sierrensis (Theobald) 
  varipalpus (Coquillett) 
 Protomacleaya Theobald berlini (Zavortink) 
  brelandi (Zavortink) 
  burgeri (Zavortink) 
  galindoi (Schick) 
  hendersoni (Cockerell) 
  homoeopus (Dyar) 
  knabi (Coquillett) 
  kompi (Vargas & Downs) 
  terrens (Walker) 
  triseriatus (Say)* 
  zavortinki (Schick) 
  zoosophus (Dyar & Knab) 
Opifex Hutton Nothoskusea Dumbleton chathamicus (Dumbleton)* 
 Opifex fuscus Hutton* 
Tanakaius Reinert, Harbach & Kitching  savoryi (Bohart) 
  togoi (Theobald)* 1*,2,4 
Aedine outgroup taxa    
Abraedes Zavortink  papago (Zavortink)* 
Aedes Meigen  cinereus Meigen* 1,2 
  esoensis Yamada 
Aztecaedes Zavortink  ramirezi (Vargas & Downs)* 
Geoskusea Edwards  baisasi (Knight & Hull) 
  longiforceps (Edwards) 
Gymnometopa Coquillett  mediovittata (Coquillett)* 
Howardina Theobald  sexlineata (Theobald) 
  walkeri (Theobald)* 
Isoaedes Reinert  cavaticus (Reinert)* 
Kenknightia Reinert  dissimilis (Leicester)* 
  harbachi (Reinert) 
Kompia Aitkin  purpureipes (Aitken)* 
Levua Stone & Bohart  geoskusea (Amos)* 
Mucidus Theobald  alternans (Westwood)* 
  laniger (Wiedemann) 
Ochlerotatus Rusticoidus Shevchenko & refiki (Medschid)* 
 Prudkina rusticus (Rossi) 
 Unassigned to subgenus communis (de Geer) 1,2 
  infirmatus Dyar & Knab 
  scapularis (Rondani)* 1,6 
Ochlerotatus’ Bruceharrisonius Reinert alektorovi (Stackelberg) 
  greenii (Theobald)* 
 Chaetocruiomyia Theobald wattensis (Taylor) 
 Pseudoskusea Theobald bancroftianus (Edwards) 
  postspiraculosus (Dobrotworsky) 
Psorophora Robineau-Desvoidy Grabhamia Theobald jamaicensis (Theobald)* 
Janthinosoma Lynch ferox (von Humboldt) 1,6 
 Arribalzaga   
 Psorophora ciliata (Fabricius)* 1,6 
Rhinoskusea Edwards  longirostris (Leicester)* 4,5 
  wardi (Reinert) 
Scutomyia Theobald  albolineata (Theobald)* 4,5 
  arboricola Knight & Rozeboom 
Stegomyia Theobald  aegypti (Linnaeus)* 1*,2*,3,4*,5*,6* 
  africana (Theobald) 
Zavortinkius Reinert  fulgens (Edwards) 
  longipalpis (Grunberg)* 
Non-aedine outgroup taxa    
Culex Linneaus Culex quinquefasciatus Say 1*,2*,3,4*,5*,6* 
Culiseta Felt Culiseta inornata (Williston) 
Mansonia Blanchard Mansonia titillans (Walker)* 1,6 
Orthopodomyia Theobald  signifera (Coquillett) 1,6 
GenusSubgenusSpeciesDistribution
Aedine ingroup taxa 
Downsiomyia Vargas  leonis (Colless) 
  nivea (Ludlow)* 
Finlaya Theobald  kochi (Doenitz)* 
  poicilia (Theobald) 4,5 
Haemagogus Conopostegus Dyar leucocelaenus (Dyar & Shannon)* 
Williston  leucotaeniatus (Komp) 
 Haemagogus equinus Theobald 
  splendens Williston* 
Halaedes Belkin  australis (Erichson)* 
Ochlerotatus Lynch    
Arribalizaga ‘Finlaya’ Theobald albilabris (Edwards) 
  alboannulatus (Macquart) 
  albocinctus (Barraud) 
  albotaeniatus (Leicester) 
  argenteitarsis (Brug) 
  argyronotum (Belkin) 
  assamensis (Theobald) 
  banksi (Edwards) 
  biocellatus (Taylor) 
  candidoscutellum (Marks) 
  chrysolineatus (Theobald) 
  chungi (Lien) 
  crossi (Lien) 
  echinus Edwards 
  elsiae (Barraud) 
  embuensis (Edwards) 
  geniculatus (Olivier) 
  gilli (Barraud) 
  gubernatoris (Giles) 
  harperi (Knight) 
  hatorii (Yamada) 2,4 
  ingrami (Edwards) 
  j. japonicus (Theobald) 1*,2*,4 
  jugraensis (Leicester) 
  keefei (King & Hoogstraal) 
  loi (Lien) 
  lophoventralis (Theobald) 
  luteostriatus (Robinson) 
  macfarlanei Edwards 
  notoscriptus (Skuse) 
  oreophilus Edwards 2,4 
  papuensis (Taylor) 
  prominens (Barraud) 
  pseudotaeniatus (Giles) 
  pulchritorax (Edwards) 
  pulchriventer (Giles) 2,4 
  rubrithorax (Macquart) 
  seoulensis (Yamada) 
  sherki (Knight) 
  subalbitarsis (King & Hoogstraal) 
  tubbutiensis (Dobrotworsky) 
 Macleaya Theobald tremulus (Theobald)* 
 Molpemyia Theobald pecuniosus (Edwards) 
 Ochlerotatus’ atropalpus (Coquillett) 1,2* 
  deserticola (Zavortink) 
  epactius (Dyar & Knab) 1,6 
  fluviatilis (Lutz) 
  muelleri (Dyar) 
  sierrensis (Theobald) 
  varipalpus (Coquillett) 
 Protomacleaya Theobald berlini (Zavortink) 
  brelandi (Zavortink) 
  burgeri (Zavortink) 
  galindoi (Schick) 
  hendersoni (Cockerell) 
  homoeopus (Dyar) 
  knabi (Coquillett) 
  kompi (Vargas & Downs) 
  terrens (Walker) 
  triseriatus (Say)* 
  zavortinki (Schick) 
  zoosophus (Dyar & Knab) 
Opifex Hutton Nothoskusea Dumbleton chathamicus (Dumbleton)* 
 Opifex fuscus Hutton* 
Tanakaius Reinert, Harbach & Kitching  savoryi (Bohart) 
  togoi (Theobald)* 1*,2,4 
Aedine outgroup taxa    
Abraedes Zavortink  papago (Zavortink)* 
Aedes Meigen  cinereus Meigen* 1,2 
  esoensis Yamada 
Aztecaedes Zavortink  ramirezi (Vargas & Downs)* 
Geoskusea Edwards  baisasi (Knight & Hull) 
  longiforceps (Edwards) 
Gymnometopa Coquillett  mediovittata (Coquillett)* 
Howardina Theobald  sexlineata (Theobald) 
  walkeri (Theobald)* 
Isoaedes Reinert  cavaticus (Reinert)* 
Kenknightia Reinert  dissimilis (Leicester)* 
  harbachi (Reinert) 
Kompia Aitkin  purpureipes (Aitken)* 
Levua Stone & Bohart  geoskusea (Amos)* 
Mucidus Theobald  alternans (Westwood)* 
  laniger (Wiedemann) 
Ochlerotatus Rusticoidus Shevchenko & refiki (Medschid)* 
 Prudkina rusticus (Rossi) 
 Unassigned to subgenus communis (de Geer) 1,2 
  infirmatus Dyar & Knab 
  scapularis (Rondani)* 1,6 
Ochlerotatus’ Bruceharrisonius Reinert alektorovi (Stackelberg) 
  greenii (Theobald)* 
 Chaetocruiomyia Theobald wattensis (Taylor) 
 Pseudoskusea Theobald bancroftianus (Edwards) 
  postspiraculosus (Dobrotworsky) 
Psorophora Robineau-Desvoidy Grabhamia Theobald jamaicensis (Theobald)* 
Janthinosoma Lynch ferox (von Humboldt) 1,6 
 Arribalzaga   
 Psorophora ciliata (Fabricius)* 1,6 
Rhinoskusea Edwards  longirostris (Leicester)* 4,5 
  wardi (Reinert) 
Scutomyia Theobald  albolineata (Theobald)* 4,5 
  arboricola Knight & Rozeboom 
Stegomyia Theobald  aegypti (Linnaeus)* 1*,2*,3,4*,5*,6* 
  africana (Theobald) 
Zavortinkius Reinert  fulgens (Edwards) 
  longipalpis (Grunberg)* 
Non-aedine outgroup taxa    
Culex Linneaus Culex quinquefasciatus Say 1*,2*,3,4*,5*,6* 
Culiseta Felt Culiseta inornata (Williston) 
Mansonia Blanchard Mansonia titillans (Walker)* 1,6 
Orthopodomyia Theobald  signifera (Coquillett) 1,6 
APPENDIX 1

List of species examined in detail during the comparative morphological analysis. An asterisk (*) following a species–author combination denotes a type species of a generic-level taxon. Geographical distribution of species is indicated by region: 1, Nearctic; 2, Palaearctic; 3, Afrotropical; 4, Oriental; 5, Australasian; 6, Neotropical. An asterisk following a geographical region indicates a relatively recent introduction.

GenusSubgenusSpeciesDistribution
Aedine ingroup taxa 
Downsiomyia Vargas  leonis (Colless) 
  nivea (Ludlow)* 
Finlaya Theobald  kochi (Doenitz)* 
  poicilia (Theobald) 4,5 
Haemagogus Conopostegus Dyar leucocelaenus (Dyar & Shannon)* 
Williston  leucotaeniatus (Komp) 
 Haemagogus equinus Theobald 
  splendens Williston* 
Halaedes Belkin  australis (Erichson)* 
Ochlerotatus Lynch    
Arribalizaga ‘Finlaya’ Theobald albilabris (Edwards) 
  alboannulatus (Macquart) 
  albocinctus (Barraud) 
  albotaeniatus (Leicester) 
  argenteitarsis (Brug) 
  argyronotum (Belkin) 
  assamensis (Theobald) 
  banksi (Edwards) 
  biocellatus (Taylor) 
  candidoscutellum (Marks) 
  chrysolineatus (Theobald) 
  chungi (Lien) 
  crossi (Lien) 
  echinus Edwards 
  elsiae (Barraud) 
  embuensis (Edwards) 
  geniculatus (Olivier) 
  gilli (Barraud) 
  gubernatoris (Giles) 
  harperi (Knight) 
  hatorii (Yamada) 2,4 
  ingrami (Edwards) 
  j. japonicus (Theobald) 1*,2*,4 
  jugraensis (Leicester) 
  keefei (King & Hoogstraal) 
  loi (Lien) 
  lophoventralis (Theobald) 
  luteostriatus (Robinson) 
  macfarlanei Edwards 
  notoscriptus (Skuse) 
  oreophilus Edwards 2,4 
  papuensis (Taylor) 
  prominens (Barraud) 
  pseudotaeniatus (Giles) 
  pulchritorax (Edwards) 
  pulchriventer (Giles) 2,4 
  rubrithorax (Macquart) 
  seoulensis (Yamada) 
  sherki (Knight) 
  subalbitarsis (King & Hoogstraal) 
  tubbutiensis (Dobrotworsky) 
 Macleaya Theobald tremulus (Theobald)* 
 Molpemyia Theobald pecuniosus (Edwards) 
 Ochlerotatus’ atropalpus (Coquillett) 1,2* 
  deserticola (Zavortink) 
  epactius (Dyar & Knab) 1,6 
  fluviatilis (Lutz) 
  muelleri (Dyar) 
  sierrensis (Theobald) 
  varipalpus (Coquillett) 
 Protomacleaya Theobald berlini (Zavortink) 
  brelandi (Zavortink) 
  burgeri (Zavortink) 
  galindoi (Schick) 
  hendersoni (Cockerell) 
  homoeopus (Dyar) 
  knabi (Coquillett) 
  kompi (Vargas & Downs) 
  terrens (Walker) 
  triseriatus (Say)* 
  zavortinki (Schick) 
  zoosophus (Dyar & Knab) 
Opifex Hutton Nothoskusea Dumbleton chathamicus (Dumbleton)* 
 Opifex fuscus Hutton* 
Tanakaius Reinert, Harbach & Kitching  savoryi (Bohart) 
  togoi (Theobald)* 1*,2,4 
Aedine outgroup taxa    
Abraedes Zavortink  papago (Zavortink)* 
Aedes Meigen  cinereus Meigen* 1,2 
  esoensis Yamada 
Aztecaedes Zavortink  ramirezi (Vargas & Downs)* 
Geoskusea Edwards  baisasi (Knight & Hull) 
  longiforceps (Edwards) 
Gymnometopa Coquillett  mediovittata (Coquillett)* 
Howardina Theobald  sexlineata (Theobald) 
  walkeri (Theobald)* 
Isoaedes Reinert  cavaticus (Reinert)* 
Kenknightia Reinert  dissimilis (Leicester)* 
  harbachi (Reinert) 
Kompia Aitkin  purpureipes (Aitken)* 
Levua Stone & Bohart  geoskusea (Amos)* 
Mucidus Theobald  alternans (Westwood)* 
  laniger (Wiedemann) 
Ochlerotatus Rusticoidus Shevchenko & refiki (Medschid)* 
 Prudkina rusticus (Rossi) 
 Unassigned to subgenus communis (de Geer) 1,2 
  infirmatus Dyar & Knab 
  scapularis (Rondani)* 1,6 
Ochlerotatus’ Bruceharrisonius Reinert alektorovi (Stackelberg) 
  greenii (Theobald)* 
 Chaetocruiomyia Theobald wattensis (Taylor) 
 Pseudoskusea Theobald bancroftianus (Edwards) 
  postspiraculosus (Dobrotworsky) 
Psorophora Robineau-Desvoidy Grabhamia Theobald jamaicensis (Theobald)* 
Janthinosoma Lynch ferox (von Humboldt) 1,6 
 Arribalzaga   
 Psorophora ciliata (Fabricius)* 1,6 
Rhinoskusea Edwards  longirostris (Leicester)* 4,5 
  wardi (Reinert) 
Scutomyia Theobald  albolineata (Theobald)* 4,5 
  arboricola Knight & Rozeboom 
Stegomyia Theobald  aegypti (Linnaeus)* 1*,2*,3,4*,5*,6* 
  africana (Theobald) 
Zavortinkius Reinert  fulgens (Edwards) 
  longipalpis (Grunberg)* 
Non-aedine outgroup taxa    
Culex Linneaus Culex quinquefasciatus Say 1*,2*,3,4*,5*,6* 
Culiseta Felt Culiseta inornata (Williston) 
Mansonia Blanchard Mansonia titillans (Walker)* 1,6 
Orthopodomyia Theobald  signifera (Coquillett) 1,6 
GenusSubgenusSpeciesDistribution
Aedine ingroup taxa 
Downsiomyia Vargas  leonis (Colless) 
  nivea (Ludlow)* 
Finlaya Theobald  kochi (Doenitz)* 
  poicilia (Theobald) 4,5 
Haemagogus Conopostegus Dyar leucocelaenus (Dyar & Shannon)* 
Williston  leucotaeniatus (Komp) 
 Haemagogus equinus Theobald 
  splendens Williston* 
Halaedes Belkin  australis (Erichson)* 
Ochlerotatus Lynch    
Arribalizaga ‘Finlaya’ Theobald albilabris (Edwards) 
  alboannulatus (Macquart) 
  albocinctus (Barraud) 
  albotaeniatus (Leicester) 
  argenteitarsis (Brug) 
  argyronotum (Belkin) 
  assamensis (Theobald) 
  banksi (Edwards) 
  biocellatus (Taylor) 
  candidoscutellum (Marks) 
  chrysolineatus (Theobald) 
  chungi (Lien) 
  crossi (Lien) 
  echinus Edwards 
  elsiae (Barraud) 
  embuensis (Edwards) 
  geniculatus (Olivier) 
  gilli (Barraud) 
  gubernatoris (Giles) 
  harperi (Knight) 
  hatorii (Yamada) 2,4 
  ingrami (Edwards) 
  j. japonicus (Theobald) 1*,2*,4 
  jugraensis (Leicester) 
  keefei (King & Hoogstraal) 
  loi (Lien) 
  lophoventralis (Theobald) 
  luteostriatus (Robinson) 
  macfarlanei Edwards 
  notoscriptus (Skuse) 
  oreophilus Edwards 2,4 
  papuensis (Taylor) 
  prominens (Barraud) 
  pseudotaeniatus (Giles) 
  pulchritorax (Edwards) 
  pulchriventer (Giles) 2,4 
  rubrithorax (Macquart) 
  seoulensis (Yamada) 
  sherki (Knight) 
  subalbitarsis (King & Hoogstraal) 
  tubbutiensis (Dobrotworsky) 
 Macleaya Theobald tremulus (Theobald)* 
 Molpemyia Theobald pecuniosus (Edwards) 
 Ochlerotatus’ atropalpus (Coquillett) 1,2* 
  deserticola (Zavortink) 
  epactius (Dyar & Knab) 1,6 
  fluviatilis (Lutz) 
  muelleri (Dyar) 
  sierrensis (Theobald) 
  varipalpus (Coquillett) 
 Protomacleaya Theobald berlini (Zavortink) 
  brelandi (Zavortink) 
  burgeri (Zavortink) 
  galindoi (Schick) 
  hendersoni (Cockerell) 
  homoeopus (Dyar) 
  knabi (Coquillett) 
  kompi (Vargas & Downs) 
  terrens (Walker) 
  triseriatus (Say)* 
  zavortinki (Schick) 
  zoosophus (Dyar & Knab) 
Opifex Hutton Nothoskusea Dumbleton chathamicus (Dumbleton)* 
 Opifex fuscus Hutton* 
Tanakaius Reinert, Harbach & Kitching  savoryi (Bohart) 
  togoi (Theobald)* 1*,2,4 
Aedine outgroup taxa    
Abraedes Zavortink  papago (Zavortink)* 
Aedes Meigen  cinereus Meigen* 1,2 
  esoensis Yamada 
Aztecaedes Zavortink  ramirezi (Vargas & Downs)* 
Geoskusea Edwards  baisasi (Knight & Hull) 
  longiforceps (Edwards) 
Gymnometopa Coquillett  mediovittata (Coquillett)* 
Howardina Theobald  sexlineata (Theobald) 
  walkeri (Theobald)* 
Isoaedes Reinert  cavaticus (Reinert)* 
Kenknightia Reinert  dissimilis (Leicester)* 
  harbachi (Reinert) 
Kompia Aitkin  purpureipes (Aitken)* 
Levua Stone & Bohart  geoskusea (Amos)* 
Mucidus Theobald  alternans (Westwood)* 
  laniger (Wiedemann) 
Ochlerotatus Rusticoidus Shevchenko & refiki (Medschid)* 
 Prudkina rusticus (Rossi) 
 Unassigned to subgenus communis (de Geer) 1,2 
  infirmatus Dyar & Knab 
  scapularis (Rondani)* 1,6 
Ochlerotatus’ Bruceharrisonius Reinert alektorovi (Stackelberg) 
  greenii (Theobald)* 
 Chaetocruiomyia Theobald wattensis (Taylor) 
 Pseudoskusea Theobald bancroftianus (Edwards) 
  postspiraculosus (Dobrotworsky) 
Psorophora Robineau-Desvoidy Grabhamia Theobald jamaicensis (Theobald)* 
Janthinosoma Lynch ferox (von Humboldt) 1,6 
 Arribalzaga   
 Psorophora ciliata (Fabricius)* 1,6 
Rhinoskusea Edwards  longirostris (Leicester)* 4,5 
  wardi (Reinert) 
Scutomyia Theobald  albolineata (Theobald)* 4,5 
  arboricola Knight & Rozeboom 
Stegomyia Theobald  aegypti (Linnaeus)* 1*,2*,3,4*,5*,6* 
  africana (Theobald) 
Zavortinkius Reinert  fulgens (Edwards) 
  longipalpis (Grunberg)* 
Non-aedine outgroup taxa    
Culex Linneaus Culex quinquefasciatus Say 1*,2*,3,4*,5*,6* 
Culiseta Felt Culiseta inornata (Williston) 
Mansonia Blanchard Mansonia titillans (Walker)* 1,6 
Orthopodomyia Theobald  signifera (Coquillett) 1,6 

APPENDIX 2

Data matrix for 116 taxa and 232 characters used in the analysis. See text for explanation of coding.

1111111111222222222233333333334
1234567890123456789012345678901234567890
Cs.  (Cus.) inornata 
Cx. (Cux.) quinquefasciatus 
Ma. (Man.) titillans 
Or. signifera 
Ab. papago 
Ae. cinereus 
Ae. esoensis 
Az. ramirezi 01 01 
Ge. baisasi 
Ge. longiforceps 
Gy. mediovittata 
Hw. sexlineata 
Hw. walkeri 
Ia. cavaticus 01 01 01 
Ke. dissimilis 01 01 
Ke. harbachi 01 
Ko. purpureipes 
Le. geoskusea 
Mu. alternans 
Mu. laniger 
‘Oc. (Brh.) alektorovi 
‘Oc. (Brh.) greenii 01 
‘Oc. (Cha.) wattensis 
Oc. (Och.) communis 01 01 
Oc. (Och.) infirmatus 
Oc. (Och.) scapularis 
‘Oc. (Psk.) bancroftianus 01 
‘Oc. (Psk.) postspiraculosus 
Oc. (Rus.) refiki 
Oc. (Rus.) rusticus 01 
Ps. (Gra.) jamaicensis 
Ps. (Jan.) ferox 01 
Ps. (Pso.) ciliata 
Rh. longirostris 01 01 
Rh. wardi 
Sc. albolineata 
Sc. arboricola 
St. aegypti 01 12 
St. africana 01 
Za. fulgens 01 
Za. longipalpis 
Do. leonis 
Do. nivea 
Fl. kochi 
Fl. poicilia 
Ha. australis 
Hg. (Con.) leucocelaenus 
Hg. (Con.) leucotaeniatus 01 
Hg. (Hag.) equinus 01 
Hg. (Hag.) splendens 
‘Oc. (Fin.) albilabris 01 01 
‘Oc. (Fin.) alboannulatus 
‘Oc. (Fin.) albocinctus 
‘Oc. (Fin.) albotaeniatus 01 
‘Oc. (Fin.) argenteitarsis 01 
‘Oc. (Fin.) argyronotum 
‘Oc. (Fin.) assamensis 
‘Oc. (Fin.) banksi 01 
‘Oc. (Fin.) biocellatus 01 
‘Oc. (Fin.) candidoscutellum 
‘Oc. (Fin.) chrysolineatus 01 01 
‘Oc. (Fin.) chungi 
‘Oc. (Fin.) crossi 
‘Oc. (Fin.) echinus 
‘Oc. (Fin.) elsiae 
‘Oc. (Fin.) embuensis 01 01 
‘Oc. (Fin.) geniculatus 
‘Oc. (Fin.) gilli 
‘Oc. (Fin.) gubernatoris 
‘Oc. (Fin.) harperi 
‘Oc. (Fin.) hatorii 01 01 
‘Oc. (Fin.) ingrami 01 
‘Oc. (Fin.) j. japonicus 01 01 01 01 
‘Oc. (Fin.) jugaraensis 
‘Oc. (Fin.) keefei 01 
‘Oc. (Fin.) loi 01 
‘Oc. (Fin.) lophoventralis 
‘Oc. (Fin.) luteostriatus 01 01 
‘Oc. (Fin.) macfarlanei 01 
‘Oc. (Fin.) notoscriptus 01 
‘Oc. (Fin.) oreophilus 12 01 
‘Oc. (Fin.) papuensis 01 01 
‘Oc. (Fin.) prominens 
‘Oc. (Fin.) pseudotaeniatus 
‘Oc. (Fin.) pulchrithorax 01 01 
‘Oc. (Fin.) pulchriventer 01 
‘Oc. (Fin.) rubrithorax 
‘Oc. (Fin.) seoulensis 01 01 
‘Oc. (Fin.) sherki 
‘Oc. (Fin.) subalbitarsis 01 
‘Oc. (Fin.) tubbutiensis 
‘Oc. (Mac.) tremulus 
‘Oc. (Mol.) pecuniosus 
‘Oc. (Och.) atropalpus 01 01 01 01 
‘Oc. (Och.) deserticola 
‘Oc. (Och.) epactius 01 
‘Oc. (Och.) fluviatilis 01 
‘Oc. (Och.) muelleri 
‘Oc. (Och.) sierrensis 01 
‘Oc. (Och.) varipalpus 01 
‘Oc. (Pro.) berlini 
‘Oc. (Pro.) brelandi 
‘Oc. (Pro.) burgeri 
‘Oc. (Pro.) galindoi 
‘Oc. (Pro.) hendersoni 
‘Oc. (Pro.) homoeopus 01 
‘Oc. (Pro.) knabi 01 01 
‘Oc. (Pro.) kompi 
‘Oc. (Pro.) terrens 
‘Oc. (Pro.) triseriatus 01 
‘Oc. (Pro.) zavortinki 
‘Oc. (Pro.) zoosophus 
Op. (Not.) chathamicus 
Op. (Opi.) fuscus 01 
Ta. savoryi 01 01 
Ta. togoi 
1111111111222222222233333333334
1234567890123456789012345678901234567890
Cs.  (Cus.) inornata 
Cx. (Cux.) quinquefasciatus 
Ma. (Man.) titillans 
Or. signifera 
Ab. papago 
Ae. cinereus 
Ae. esoensis 
Az. ramirezi 01 01 
Ge. baisasi 
Ge. longiforceps 
Gy. mediovittata 
Hw. sexlineata 
Hw. walkeri 
Ia. cavaticus 01 01 01 
Ke. dissimilis 01 01 
Ke. harbachi 01 
Ko. purpureipes 
Le. geoskusea 
Mu. alternans 
Mu. laniger 
‘Oc. (Brh.) alektorovi 
‘Oc. (Brh.) greenii 01 
‘Oc. (Cha.) wattensis 
Oc. (Och.) communis 01 01 
Oc. (Och.) infirmatus 
Oc. (Och.) scapularis 
‘Oc. (Psk.) bancroftianus 01 
‘Oc. (Psk.) postspiraculosus 
Oc. (Rus.) refiki 
Oc. (Rus.) rusticus 01 
Ps. (Gra.) jamaicensis 
Ps. (Jan.) ferox 01 
Ps. (Pso.) ciliata 
Rh. longirostris 01 01 
Rh. wardi 
Sc. albolineata 
Sc. arboricola 
St. aegypti 01 12 
St. africana 01 
Za. fulgens 01 
Za. longipalpis 
Do. leonis 
Do. nivea 
Fl. kochi 
Fl. poicilia 
Ha. australis 
Hg. (Con.) leucocelaenus 
Hg. (Con.) leucotaeniatus 01 
Hg. (Hag.) equinus 01 
Hg. (Hag.) splendens 
‘Oc. (Fin.) albilabris 01 01 
‘Oc. (Fin.) alboannulatus 
‘Oc. (Fin.) albocinctus 
‘Oc. (Fin.) albotaeniatus 01 
‘Oc. (Fin.) argenteitarsis 01 
‘Oc. (Fin.) argyronotum 
‘Oc. (Fin.) assamensis 
‘Oc. (Fin.) banksi 01 
‘Oc. (Fin.) biocellatus 01 
‘Oc. (Fin.) candidoscutellum 
‘Oc. (Fin.) chrysolineatus 01 01 
‘Oc. (Fin.) chungi 
‘Oc. (Fin.) crossi 
‘Oc. (Fin.) echinus 
‘Oc. (Fin.) elsiae 
‘Oc. (Fin.) embuensis 01 01 
‘Oc. (Fin.) geniculatus 
‘Oc. (Fin.) gilli 
‘Oc. (Fin.) gubernatoris 
‘Oc. (Fin.) harperi 
‘Oc. (Fin.) hatorii 01 01 
‘Oc. (Fin.) ingrami 01 
‘Oc. (Fin.) j. japonicus 01 01 01 01 
‘Oc. (Fin.) jugaraensis 
‘Oc. (Fin.) keefei 01 
‘Oc. (Fin.) loi 01 
‘Oc. (Fin.) lophoventralis 
‘Oc. (Fin.) luteostriatus 01 01 
‘Oc. (Fin.) macfarlanei 01 
‘Oc. (Fin.) notoscriptus 01 
‘Oc. (Fin.) oreophilus 12 01 
‘Oc. (Fin.) papuensis 01 01 
‘Oc. (Fin.) prominens 
‘Oc. (Fin.) pseudotaeniatus 
‘Oc. (Fin.) pulchrithorax 01 01 
‘Oc. (Fin.) pulchriventer 01 
‘Oc. (Fin.) rubrithorax 
‘Oc. (Fin.) seoulensis 01 01 
‘Oc. (Fin.) sherki 
‘Oc. (Fin.) subalbitarsis 01 
‘Oc. (Fin.) tubbutiensis 
‘Oc. (Mac.) tremulus 
‘Oc. (Mol.) pecuniosus 
‘Oc. (Och.) atropalpus 01 01 01 01 
‘Oc. (Och.) deserticola 
‘Oc. (Och.) epactius 01 
‘Oc. (Och.) fluviatilis 01 
‘Oc. (Och.) muelleri 
‘Oc. (Och.) sierrensis 01 
‘Oc. (Och.) varipalpus 01 
‘Oc. (Pro.) berlini 
‘Oc. (Pro.) brelandi 
‘Oc. (Pro.) burgeri 
‘Oc. (Pro.) galindoi 
‘Oc. (Pro.) hendersoni 
‘Oc. (Pro.) homoeopus 01 
‘Oc. (Pro.) knabi 01 01 
‘Oc. (Pro.) kompi 
‘Oc. (Pro.) terrens 
‘Oc. (Pro.) triseriatus 01 
‘Oc. (Pro.) zavortinki 
‘Oc. (Pro.) zoosophus 
Op. (Not.) chathamicus 
Op. (Opi.) fuscus 01 
Ta. savoryi 01 01 
Ta. togoi 
APPENDIX 2

Data matrix for 116 taxa and 232 characters used in the analysis. See text for explanation of coding.

1111111111222222222233333333334
1234567890123456789012345678901234567890
Cs.  (Cus.) inornata 
Cx. (Cux.) quinquefasciatus 
Ma. (Man.) titillans 
Or. signifera 
Ab. papago 
Ae. cinereus 
Ae. esoensis 
Az. ramirezi 01 01 
Ge. baisasi 
Ge. longiforceps 
Gy. mediovittata 
Hw. sexlineata 
Hw. walkeri 
Ia. cavaticus 01 01 01 
Ke. dissimilis 01 01 
Ke. harbachi 01 
Ko. purpureipes 
Le. geoskusea 
Mu. alternans 
Mu. laniger 
‘Oc. (Brh.) alektorovi 
‘Oc. (Brh.) greenii 01 
‘Oc. (Cha.) wattensis 
Oc. (Och.) communis 01 01 
Oc. (Och.) infirmatus 
Oc. (Och.) scapularis 
‘Oc. (Psk.) bancroftianus 01 
‘Oc. (Psk.) postspiraculosus 
Oc. (Rus.) refiki 
Oc. (Rus.) rusticus 01 
Ps. (Gra.) jamaicensis 
Ps. (Jan.) ferox 01 
Ps. (Pso.) ciliata 
Rh. longirostris 01 01 
Rh. wardi 
Sc. albolineata 
Sc. arboricola 
St. aegypti 01 12 
St. africana 01 
Za. fulgens 01 
Za. longipalpis 
Do. leonis 
Do. nivea 
Fl. kochi 
Fl. poicilia 
Ha. australis 
Hg. (Con.) leucocelaenus 
Hg. (Con.) leucotaeniatus 01 
Hg. (Hag.) equinus 01 
Hg. (Hag.) splendens 
‘Oc. (Fin.) albilabris 01 01 
‘Oc. (Fin.) alboannulatus 
‘Oc. (Fin.) albocinctus 
‘Oc. (Fin.) albotaeniatus 01 
‘Oc. (Fin.) argenteitarsis 01 
‘Oc. (Fin.) argyronotum 
‘Oc. (Fin.) assamensis 
‘Oc. (Fin.) banksi 01 
‘Oc. (Fin.) biocellatus 01 
‘Oc. (Fin.) candidoscutellum 
‘Oc. (Fin.) chrysolineatus 01 01 
‘Oc. (Fin.) chungi 
‘Oc. (Fin.) crossi 
‘Oc. (Fin.) echinus 
‘Oc. (Fin.) elsiae 
‘Oc. (Fin.) embuensis 01 01 
‘Oc. (Fin.) geniculatus 
‘Oc. (Fin.) gilli 
‘Oc. (Fin.) gubernatoris 
‘Oc. (Fin.) harperi 
‘Oc. (Fin.) hatorii 01 01 
‘Oc. (Fin.) ingrami 01 
‘Oc. (Fin.) j. japonicus 01 01 01 01 
‘Oc. (Fin.) jugaraensis 
‘Oc. (Fin.) keefei 01 
‘Oc. (Fin.) loi 01 
‘Oc. (Fin.) lophoventralis 
‘Oc. (Fin.) luteostriatus 01 01 
‘Oc. (Fin.) macfarlanei 01 
‘Oc. (Fin.) notoscriptus 01 
‘Oc. (Fin.) oreophilus 12 01 
‘Oc. (Fin.) papuensis 01 01 
‘Oc. (Fin.) prominens 
‘Oc. (Fin.) pseudotaeniatus 
‘Oc. (Fin.) pulchrithorax 01 01 
‘Oc. (Fin.) pulchriventer 01 
‘Oc. (Fin.) rubrithorax 
‘Oc. (Fin.) seoulensis 01 01 
‘Oc. (Fin.) sherki 
‘Oc. (Fin.) subalbitarsis 01 
‘Oc. (Fin.) tubbutiensis 
‘Oc. (Mac.) tremulus 
‘Oc. (Mol.) pecuniosus 
‘Oc. (Och.) atropalpus 01 01 01 01 
‘Oc. (Och.) deserticola 
‘Oc. (Och.) epactius 01 
‘Oc. (Och.) fluviatilis 01 
‘Oc. (Och.) muelleri 
‘Oc. (Och.) sierrensis 01 
‘Oc. (Och.) varipalpus 01 
‘Oc. (Pro.) berlini 
‘Oc. (Pro.) brelandi 
‘Oc. (Pro.) burgeri 
‘Oc. (Pro.) galindoi 
‘Oc. (Pro.) hendersoni 
‘Oc. (Pro.) homoeopus 01 
‘Oc. (Pro.) knabi 01 01 
‘Oc. (Pro.) kompi 
‘Oc. (Pro.) terrens 
‘Oc. (Pro.) triseriatus 01 
‘Oc. (Pro.) zavortinki 
‘Oc. (Pro.) zoosophus 
Op. (Not.) chathamicus 
Op. (Opi.) fuscus 01 
Ta. savoryi 01 01 
Ta. togoi 
1111111111222222222233333333334
1234567890123456789012345678901234567890
Cs.  (Cus.) inornata 
Cx. (Cux.) quinquefasciatus 
Ma. (Man.) titillans 
Or. signifera 
Ab. papago 
Ae. cinereus 
Ae. esoensis 
Az. ramirezi 01 01 
Ge. baisasi 
Ge. longiforceps 
Gy. mediovittata 
Hw. sexlineata 
Hw. walkeri 
Ia. cavaticus 01 01 01 
Ke. dissimilis 01 01 
Ke. harbachi 01 
Ko. purpureipes 
Le. geoskusea 
Mu. alternans 
Mu. laniger 
‘Oc. (Brh.) alektorovi 
‘Oc. (Brh.) greenii 01 
‘Oc. (Cha.) wattensis 
Oc. (Och.) communis 01 01 
Oc. (Och.) infirmatus 
Oc. (Och.) scapularis 
‘Oc. (Psk.) bancroftianus 01 
‘Oc. (Psk.) postspiraculosus 
Oc. (Rus.) refiki 
Oc. (Rus.) rusticus 01 
Ps. (Gra.) jamaicensis 
Ps. (Jan.) ferox 01 
Ps. (Pso.) ciliata 
Rh. longirostris 01 01 
Rh. wardi 
Sc. albolineata 
Sc. arboricola 
St. aegypti 01 12 
St. africana 01 
Za. fulgens 01 
Za. longipalpis 
Do. leonis 
Do. nivea 
Fl. kochi 
Fl. poicilia 
Ha. australis 
Hg. (Con.) leucocelaenus 
Hg. (Con.) leucotaeniatus 01 
Hg. (Hag.) equinus 01 
Hg. (Hag.) splendens 
‘Oc. (Fin.) albilabris 01 01 
‘Oc. (Fin.) alboannulatus 
‘Oc. (Fin.) albocinctus 
‘Oc. (Fin.) albotaeniatus 01 
‘Oc. (Fin.) argenteitarsis 01 
‘Oc. (Fin.) argyronotum 
‘Oc. (Fin.) assamensis 
‘Oc. (Fin.) banksi 01 
‘Oc. (Fin.) biocellatus 01 
‘Oc. (Fin.) candidoscutellum 
‘Oc. (Fin.) chrysolineatus 01 01 
‘Oc. (Fin.) chungi 
‘Oc. (Fin.) crossi 
‘Oc. (Fin.) echinus 
‘Oc. (Fin.) elsiae 
‘Oc. (Fin.) embuensis 01 01 
‘Oc. (Fin.) geniculatus 
‘Oc. (Fin.) gilli 
‘Oc. (Fin.) gubernatoris 
‘Oc. (Fin.) harperi 
‘Oc. (Fin.) hatorii 01 01 
‘Oc. (Fin.) ingrami 01 
‘Oc. (Fin.) j. japonicus 01 01 01 01 
‘Oc. (Fin.) jugaraensis 
‘Oc. (Fin.) keefei 01 
‘Oc. (Fin.) loi 01 
‘Oc. (Fin.) lophoventralis 
‘Oc. (Fin.) luteostriatus 01 01 
‘Oc. (Fin.) macfarlanei 01 
‘Oc. (Fin.) notoscriptus 01 
‘Oc. (Fin.) oreophilus 12 01 
‘Oc. (Fin.) papuensis 01 01 
‘Oc. (Fin.) prominens 
‘Oc. (Fin.) pseudotaeniatus 
‘Oc. (Fin.) pulchrithorax 01 01 
‘Oc. (Fin.) pulchriventer 01 
‘Oc. (Fin.) rubrithorax 
‘Oc. (Fin.) seoulensis 01 01 
‘Oc. (Fin.) sherki 
‘Oc. (Fin.) subalbitarsis 01 
‘Oc. (Fin.) tubbutiensis 
‘Oc. (Mac.) tremulus 
‘Oc. (Mol.) pecuniosus 
‘Oc. (Och.) atropalpus 01 01 01 01 
‘Oc. (Och.) deserticola 
‘Oc. (Och.) epactius 01 
‘Oc. (Och.) fluviatilis 01 
‘Oc. (Och.) muelleri 
‘Oc. (Och.) sierrensis 01 
‘Oc. (Och.) varipalpus 01 
‘Oc. (Pro.) berlini 
‘Oc. (Pro.) brelandi 
‘Oc. (Pro.) burgeri 
‘Oc. (Pro.) galindoi 
‘Oc. (Pro.) hendersoni 
‘Oc. (Pro.) homoeopus 01 
‘Oc. (Pro.) knabi 01 01 
‘Oc. (Pro.) kompi 
‘Oc. (Pro.) terrens 
‘Oc. (Pro.) triseriatus 01 
‘Oc. (Pro.) zavortinki 
‘Oc. (Pro.) zoosophus 
Op. (Not.) chathamicus 
Op. (Opi.) fuscus 01 
Ta. savoryi 01 01 
Ta. togoi 

APPENDIX 2 Continued
4444444445555555555666666666677777777778
1234567890123456789012345678901234567890
Cs.  (Cus.) inornata 01 
Cx. (Cux.) quinquefasciatus 
Ma. (Man.) titillans 
Or. signifera 
Ab. papago 01 
Ae. cinereus 12 01 
Ae. esoensis 
Az. ramirezi 01 
Ge. baisasi 01 
Ge. longiforceps 01 
Gy. mediovittata 
Hw. sexlineata 
Hw. walkeri 
Ia. cavaticus 01 12 
Ke. dissimilis 
Ke. harbachi 
Ko. purpureipes 
Le. geoskusea 
Mu. alternans 01 01 12 
Mu. laniger 
‘Oc. (Brh.) alektorovi 01 
‘Oc. (Brh.) greenii 
‘Oc. (Cha.) wattensis 
Oc. (Och.) communis 01 
Oc. (Och.) infirmatus 01 01 
Oc. (Och.) scapularis 01 
‘Oc. (Psk.) bancroftianus 
‘Oc. (Psk.) postspiraculosus 01 
Oc. (Rus.) refiki 
Oc. (Rus.) rusticus 
Ps. (Gra.) jamaicensis 01 01 
Ps. (Jan.) ferox 01 
Ps. (Pso.) ciliata 01 
Rh. longirostris 01 01 
Rh. wardi 01 
Sc. albolineata 
Sc. arboricola 01 
St. aegypti 01 12 
St. africana 
Za. fulgens 
Za. longipalpis 
Do. leonis 12 
Do. nivea 12 
Fl. kochi 
Fl. poicilia 
Ha. australis 
Hg. (Con.) leucocelaenus 12 
Hg. (Con.) leucotaeniatus 
Hg. (Hag.) equinus 
Hg. (Hag.) splendens 
‘Oc. (Fin.) albilabris 01 01 
‘Oc. (Fin.) alboannulatus 
‘Oc. (Fin.) albocinctus 
‘Oc. (Fin.) albotaeniatus 
‘Oc. (Fin.) argenteitarsis 
‘Oc. (Fin.) argyronotum 
‘Oc. (Fin.) assamensis 
‘Oc. (Fin.) banksi 
‘Oc. (Fin.) biocellatus 
‘Oc. (Fin.) candidoscutellum 
‘Oc. (Fin.) chrysolineatus 
‘Oc. (Fin.) chungi 
‘Oc. (Fin.) crossi 
‘Oc. (Fin.) echinus 
‘Oc. (Fin.) elsiae 
‘Oc. (Fin.) embuensis 
‘Oc. (Fin.) geniculatus 
‘Oc. (Fin.) gilli 
‘Oc. (Fin.) gubernatoris 
‘Oc. (Fin.) harperi 
‘Oc. (Fin.) hatorii 01 01 01 
‘Oc. (Fin.) ingrami 
‘Oc. (Fin.) j. japonicus 01 
‘Oc. (Fin.) jugaraensis 01 
‘Oc. (Fin.) keefei 
‘Oc. (Fin.) loi 01 
‘Oc. (Fin.) lophoventralis 
‘Oc. (Fin.) luteostriatus 
‘Oc. (Fin.) macfarlanei 
‘Oc. (Fin.) notoscriptus 01 01 01 
‘Oc. (Fin.) oreophilus 
‘Oc. (Fin.) papuensis 
‘Oc. (Fin.) prominens 
‘Oc. (Fin.) pseudotaeniatus 
‘Oc. (Fin.) pulchrithorax 01 12 
‘Oc. (Fin.) pulchriventer 01 
‘Oc. (Fin.) rubrithorax 
‘Oc. (Fin.) seoulensis 
‘Oc. (Fin.) sherki 01 
‘Oc. (Fin.) subalbitarsis 
‘Oc. (Fin.) tubbutiensis 
‘Oc. (Mac.) tremulus 
‘Oc. (Mol.) pecuniosus 01 
‘Oc. (Och.) atropalpus 01 01 
‘Oc. (Och.) deserticola 
‘Oc. (Och.) epactius 
‘Oc. (Och.) fluviatilis 01 
‘Oc. (Och.) muelleri 01 
‘Oc. (Och.) sierrensis 01 
‘Oc. (Och.) varipalpus 01 01 
‘Oc. (Pro.) berlini 
‘Oc. (Pro.) brelandi 
‘Oc. (Pro.) burgeri 
‘Oc. (Pro.) galindoi 
‘Oc. (Pro.) hendersoni 
‘Oc. (Pro.) homoeopus 
‘Oc. (Pro.) knabi 
‘Oc. (Pro.) kompi 
‘Oc. (Pro.) terrens 
‘Oc. (Pro.) triseriatus 12 
‘Oc. (Pro.) zavortinki 01 
‘Oc. (Pro.) zoosophus 
Op. (Not.) chathamicus 
Op. (Opi.) fuscus 01 
Ta. savoryi 
Ta. togoi 
4444444445555555555666666666677777777778
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Cs.  (Cus.) inornata 01 
Cx. (Cux.) quinquefasciatus 
Ma. (Man.) titillans 
Or. signifera 
Ab. papago 01 
Ae. cinereus 12 01 
Ae. esoensis 
Az. ramirezi 01 
Ge. baisasi 01 
Ge. longiforceps 01 
Gy. mediovittata 
Hw. sexlineata 
Hw. walkeri 
Ia. cavaticus 01 12 
Ke. dissimilis 
Ke. harbachi 
Ko. purpureipes 
Le. geoskusea 
Mu. alternans 01 01 12 
Mu. laniger 
‘Oc. (Brh.) alektorovi 01 
‘Oc. (Brh.) greenii 
‘Oc. (Cha.) wattensis 
Oc. (Och.) communis 01 
Oc. (Och.) infirmatus 01 01 
Oc. (Och.) scapularis 01 
‘Oc. (Psk.) bancroftianus 
‘Oc. (Psk.) postspiraculosus 01 
Oc. (Rus.) refiki 
Oc. (Rus.) rusticus 
Ps. (Gra.) jamaicensis 01 01 
Ps. (Jan.) ferox 01 
Ps. (Pso.) ciliata 01 
Rh. longirostris 01 01 
Rh. wardi 01 
Sc. albolineata 
Sc. arboricola 01 
St. aegypti 01 12 
St. africana 
Za. fulgens 
Za. longipalpis 
Do. leonis 12 
Do. nivea 12 
Fl. kochi 
Fl. poicilia 
Ha. australis 
Hg. (Con.) leucocelaenus 12 
Hg. (Con.) leucotaeniatus 
Hg. (Hag.) equinus 
Hg. (Hag.) splendens 
‘Oc. (Fin.) albilabris 01 01 
‘Oc. (Fin.) alboannulatus 
‘Oc. (Fin.) albocinctus 
‘Oc. (Fin.) albotaeniatus 
‘Oc. (Fin.) argenteitarsis 
‘Oc. (Fin.) argyronotum 
‘Oc. (Fin.) assamensis 
‘Oc. (Fin.) banksi 
‘Oc. (Fin.) biocellatus 
‘Oc. (Fin.) candidoscutellum 
‘Oc. (Fin.) chrysolineatus 
‘Oc. (Fin.) chungi 
‘Oc. (Fin.) crossi 
‘Oc. (Fin.) echinus 
‘Oc. (Fin.) elsiae 
‘Oc. (Fin.) embuensis 
‘Oc. (Fin.) geniculatus 
‘Oc. (Fin.) gilli 
‘Oc. (Fin.) gubernatoris 
‘Oc. (Fin.) harperi 
‘Oc. (Fin.) hatorii 01 01 01 
‘Oc. (Fin.) ingrami 
‘Oc. (Fin.) j. japonicus 01 
‘Oc. (Fin.) jugaraensis 01 
‘Oc. (Fin.) keefei 
‘Oc. (Fin.) loi 01 
‘Oc. (Fin.) lophoventralis 
‘Oc. (Fin.) luteostriatus 
‘Oc. (Fin.) macfarlanei 
‘Oc. (Fin.) notoscriptus 01 01 01 
‘Oc. (Fin.) oreophilus 
‘Oc. (Fin.) papuensis 
‘Oc. (Fin.) prominens 
‘Oc. (Fin.) pseudotaeniatus 
‘Oc. (Fin.) pulchrithorax 01 12 
‘Oc. (Fin.) pulchriventer 01 
‘Oc. (Fin.) rubrithorax 
‘Oc. (Fin.) seoulensis 
‘Oc. (Fin.) sherki 01 
‘Oc. (Fin.) subalbitarsis 
‘Oc. (Fin.) tubbutiensis 
‘Oc. (Mac.) tremulus 
‘Oc. (Mol.) pecuniosus 01 
‘Oc. (Och.) atropalpus 01 01 
‘Oc. (Och.) deserticola 
‘Oc. (Och.) epactius 
‘Oc. (Och.) fluviatilis 01 
‘Oc. (Och.) muelleri 01 
‘Oc. (Och.) sierrensis 01 
‘Oc. (Och.) varipalpus 01 01 
‘Oc. (Pro.) berlini 
‘Oc. (Pro.) brelandi 
‘Oc. (Pro.) burgeri 
‘Oc. (Pro.) galindoi 
‘Oc. (Pro.) hendersoni 
‘Oc. (Pro.) homoeopus 
‘Oc. (Pro.) knabi 
‘Oc. (Pro.) kompi 
‘Oc. (Pro.) terrens 
‘Oc. (Pro.) triseriatus 12 
‘Oc. (Pro.) zavortinki 01 
‘Oc. (Pro.) zoosophus 
Op. (Not.) chathamicus 
Op. (Opi.) fuscus 01 
Ta. savoryi 
Ta. togoi 
APPENDIX 2 Continued
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Cs.  (Cus.) inornata 01 
Cx. (Cux.) quinquefasciatus 
Ma. (Man.) titillans 
Or. signifera 
Ab. papago 01 
Ae. cinereus 12 01 
Ae. esoensis 
Az. ramirezi 01 
Ge. baisasi 01 
Ge. longiforceps 01 
Gy. mediovittata 
Hw. sexlineata 
Hw. walkeri 
Ia. cavaticus 01 12 
Ke. dissimilis 
Ke. harbachi 
Ko. purpureipes 
Le. geoskusea 
Mu. alternans 01 01 12 
Mu. laniger 
‘Oc. (Brh.) alektorovi 01 
‘Oc. (Brh.) greenii 
‘Oc. (Cha.) wattensis 
Oc. (Och.) communis 01 
Oc. (Och.) infirmatus 01 01 
Oc. (Och.) scapularis 01 
‘Oc. (Psk.) bancroftianus 
‘Oc. (Psk.) postspiraculosus 01 
Oc. (Rus.) refiki 
Oc. (Rus.) rusticus 
Ps. (Gra.) jamaicensis 01 01 
Ps. (Jan.) ferox 01 
Ps. (Pso.) ciliata 01 
Rh. longirostris 01 01 
Rh. wardi 01 
Sc. albolineata 
Sc. arboricola 01 
St. aegypti 01 12 
St. africana 
Za. fulgens 
Za. longipalpis 
Do. leonis 12 
Do. nivea 12 
Fl. kochi 
Fl. poicilia 
Ha. australis 
Hg. (Con.) leucocelaenus 12 
Hg. (Con.) leucotaeniatus 
Hg. (Hag.) equinus 
Hg. (Hag.) splendens 
‘Oc. (Fin.) albilabris 01 01 
‘Oc. (Fin.) alboannulatus 
‘Oc. (Fin.) albocinctus 
‘Oc. (Fin.) albotaeniatus 
‘Oc. (Fin.) argenteitarsis 
‘Oc. (Fin.) argyronotum 
‘Oc. (Fin.) assamensis 
‘Oc. (Fin.) banksi 
‘Oc. (Fin.) biocellatus 
‘Oc. (Fin.) candidoscutellum 
‘Oc. (Fin.) chrysolineatus 
‘Oc. (Fin.) chungi 
‘Oc. (Fin.) crossi 
‘Oc. (Fin.) echinus 
‘Oc. (Fin.) elsiae 
‘Oc. (Fin.) embuensis 
‘Oc. (Fin.) geniculatus 
‘Oc. (Fin.) gilli 
‘Oc. (Fin.) gubernatoris 
‘Oc. (Fin.) harperi 
‘Oc. (Fin.) hatorii 01 01 01 
‘Oc. (Fin.) ingrami 
‘Oc. (Fin.) j. japonicus 01 
‘Oc. (Fin.) jugaraensis 01 
‘Oc. (Fin.) keefei 
‘Oc. (Fin.) loi 01 
‘Oc. (Fin.) lophoventralis 
‘Oc. (Fin.) luteostriatus 
‘Oc. (Fin.) macfarlanei 
‘Oc. (Fin.) notoscriptus 01 01 01 
‘Oc. (Fin.) oreophilus 
‘Oc. (Fin.) papuensis 
‘Oc. (Fin.) prominens 
‘Oc. (Fin.) pseudotaeniatus 
‘Oc. (Fin.) pulchrithorax 01 12 
‘Oc. (Fin.) pulchriventer 01 
‘Oc. (Fin.) rubrithorax 
‘Oc. (Fin.) seoulensis 
‘Oc. (Fin.) sherki 01 
‘Oc. (Fin.) subalbitarsis 
‘Oc. (Fin.) tubbutiensis 
‘Oc. (Mac.) tremulus 
‘Oc. (Mol.) pecuniosus 01 
‘Oc. (Och.) atropalpus 01 01 
‘Oc. (Och.) deserticola 
‘Oc. (Och.) epactius 
‘Oc. (Och.) fluviatilis 01 
‘Oc. (Och.) muelleri 01 
‘Oc. (Och.) sierrensis 01 
‘Oc. (Och.) varipalpus 01 01 
‘Oc. (Pro.) berlini 
‘Oc. (Pro.) brelandi 
‘Oc. (Pro.) burgeri 
‘Oc. (Pro.) galindoi 
‘Oc. (Pro.) hendersoni 
‘Oc. (Pro.) homoeopus 
‘Oc. (Pro.) knabi 
‘Oc. (Pro.) kompi 
‘Oc. (Pro.) terrens 
‘Oc. (Pro.) triseriatus 12 
‘Oc. (Pro.) zavortinki 01 
‘Oc. (Pro.) zoosophus 
Op. (Not.) chathamicus 
Op. (Opi.) fuscus 01 
Ta. savoryi 
Ta. togoi 
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Cs.  (Cus.) inornata 01 
Cx. (Cux.) quinquefasciatus 
Ma. (Man.) titillans 
Or. signifera 
Ab. papago 01 
Ae. cinereus 12 01 
Ae. esoensis 
Az. ramirezi 01 
Ge. baisasi 01 
Ge. longiforceps 01 
Gy. mediovittata 
Hw. sexlineata 
Hw. walkeri 
Ia. cavaticus 01 12 
Ke. dissimilis 
Ke. harbachi 
Ko. purpureipes 
Le. geoskusea 
Mu. alternans 01 01 12 
Mu. laniger 
‘Oc. (Brh.) alektorovi 01 
‘Oc. (Brh.) greenii 
‘Oc. (Cha.) wattensis 
Oc. (Och.) communis 01 
Oc. (Och.) infirmatus 01 01 
Oc. (Och.) scapularis 01 
‘Oc. (Psk.) bancroftianus 
‘Oc. (Psk.) postspiraculosus 01 
Oc. (Rus.) refiki 
Oc. (Rus.) rusticus 
Ps. (Gra.) jamaicensis 01 01 
Ps. (Jan.) ferox 01 
Ps. (Pso.) ciliata 01 
Rh. longirostris 01 01 
Rh. wardi 01 
Sc. albolineata 
Sc. arboricola 01 
St. aegypti 01 12 
St. africana 
Za. fulgens 
Za. longipalpis 
Do. leonis 12 
Do. nivea 12 
Fl. kochi 
Fl. poicilia 
Ha. australis 
Hg. (Con.) leucocelaenus 12 
Hg. (Con.) leucotaeniatus 
Hg. (Hag.) equinus 
Hg. (Hag.) splendens 
‘Oc. (Fin.) albilabris 01 01 
‘Oc. (Fin.) alboannulatus 
‘Oc. (Fin.) albocinctus 
‘Oc. (Fin.) albotaeniatus 
‘Oc. (Fin.) argenteitarsis 
‘Oc. (Fin.) argyronotum 
‘Oc. (Fin.) assamensis 
‘Oc. (Fin.) banksi 
‘Oc. (Fin.) biocellatus 
‘Oc. (Fin.) candidoscutellum 
‘Oc. (Fin.) chrysolineatus 
‘Oc. (Fin.) chungi 
‘Oc. (Fin.) crossi 
‘Oc. (Fin.) echinus 
‘Oc. (Fin.) elsiae 
‘Oc. (Fin.) embuensis 
‘Oc. (Fin.) geniculatus 
‘Oc. (Fin.) gilli 
‘Oc. (Fin.) gubernatoris 
‘Oc. (Fin.) harperi 
‘Oc. (Fin.) hatorii 01 01 01 
‘Oc. (Fin.) ingrami 
‘Oc. (Fin.) j. japonicus 01 
‘Oc. (Fin.) jugaraensis 01 
‘Oc. (Fin.) keefei 
‘Oc. (Fin.) loi 01 
‘Oc. (Fin.) lophoventralis 
‘Oc. (Fin.) luteostriatus 
‘Oc. (Fin.) macfarlanei 
‘Oc. (Fin.) notoscriptus 01 01 01 
‘Oc. (Fin.) oreophilus 
‘Oc. (Fin.) papuensis 
‘Oc. (Fin.) prominens 
‘Oc. (Fin.) pseudotaeniatus 
‘Oc. (Fin.) pulchrithorax 01 12 
‘Oc. (Fin.) pulchriventer 01 
‘Oc. (Fin.) rubrithorax 
‘Oc. (Fin.) seoulensis 
‘Oc. (Fin.) sherki 01 
‘Oc. (Fin.) subalbitarsis 
‘Oc. (Fin.) tubbutiensis 
‘Oc. (Mac.) tremulus 
‘Oc. (Mol.) pecuniosus 01 
‘Oc. (Och.) atropalpus 01 01 
‘Oc. (Och.) deserticola 
‘Oc. (Och.) epactius 
‘Oc. (Och.) fluviatilis 01 
‘Oc. (Och.) muelleri 01 
‘Oc. (Och.) sierrensis 01 
‘Oc. (Och.) varipalpus 01 01 
‘Oc. (Pro.) berlini 
‘Oc. (Pro.) brelandi 
‘Oc. (Pro.) burgeri 
‘Oc. (Pro.) galindoi 
‘Oc. (Pro.) hendersoni 
‘Oc. (Pro.) homoeopus 
‘Oc. (Pro.) knabi 
‘Oc. (Pro.) kompi 
‘Oc. (Pro.) terrens 
‘Oc. (Pro.) triseriatus 12 
‘Oc. (Pro.) zavortinki 01 
‘Oc. (Pro.) zoosophus 
Op. (Not.) chathamicus 
Op. (Opi.) fuscus 01 
Ta. savoryi 
Ta. togoi 

APPENDIX 2 Continued
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Cs.  (Cus.) inornata 01 12 
Cx. (Cux.) quinquefasciatus 
Ma. (Man.) titillans – 
Or. signifera 
Ab. papago 
Ae. cinereus 
Ae. esoensis 
Az. ramirezi 
Ge. baisasi 01 
Ge. longiforceps 
Gy. mediovittata 
Hw. sexlineata 01 
Hw. walkeri 01 01 01 
Ia. cavaticus 01 01 01 12 
Ke. dissimilis 
Ke. harbachi 01 
Ko. purpureipes 
Le. geoskusea 
Mu. alternans 01 
Mu. laniger 
‘Oc. (Brh.) alektorovi 01 01 01 
‘Oc. (Brh.) greenii 
‘Oc. (Cha.) wattensis 
Oc. (Och.) communis 01 
Oc. (Och.) infirmatus 
Oc. (Och.) scapularis 
‘Oc. (Psk.) bancroftianus 
‘Oc. (Psk.) postspiraculosus 01 
Oc. (Rus.) refiki 
Oc. (Rus.) rusticus 
Ps. (Gra.) jamaicensis 01 
Ps. (Jan.) ferox 12 
Ps. (Pso.) ciliata 01 01 
Rh. longirostris 
Rh. wardi 01 
Sc. albolineata 
Sc. arboricola 01 
St. aegypti 
St. africana 01 
Za. fulgens 01 
Za. longipalpis 
Do. leonis 12 
Do. nivea 12 
Fl. kochi 
Fl. poicilia 01 
Ha. australis 
Hg. (Con.) leucocelaenus 
Hg. (Con.) leucotaeniatus 
Hg. (Hag.) equinus 
Hg. (Hag.) splendens 
‘Oc. (Fin.) albilabris 
‘Oc. (Fin.) alboannulatus 
‘Oc. (Fin.) albocinctus 
‘Oc. (Fin.) albotaeniatus 
‘Oc. (Fin.) argenteitarsis 
‘Oc. (Fin.) argyronotum 01 01 01 
‘Oc. (Fin.) assamensis 
‘Oc. (Fin.) banksi 01 01 
‘Oc. (Fin.) biocellatus 
‘Oc. (Fin.) candidoscutellum 
‘Oc. (Fin.) chrysolineatus 
‘Oc. (Fin.) chungi 
‘Oc. (Fin.) crossi 
‘Oc. (Fin.) echinus 
‘Oc. (Fin.) elsiae 
‘Oc. (Fin.) embuensis 
‘Oc. (Fin.) geniculatus 01 
‘Oc. (Fin.) gilli 
‘Oc. (Fin.) gubernatoris 
‘Oc. (Fin.) harperi 
‘Oc. (Fin.) hatorii 
‘Oc. (Fin.) ingrami 
‘Oc. (Fin.) j. japonicus 
‘Oc. (Fin.) jugaraensis 
‘Oc. (Fin.) keefei 
‘Oc. (Fin.) loi 
‘Oc. (Fin.) lophoventralis 01 
‘Oc. (Fin.) luteostriatus 
‘Oc. (Fin.) macfarlanei 01 
‘Oc. (Fin.) notoscriptus 01 
‘Oc. (Fin.) oreophilus 
‘Oc. (Fin.) papuensis 
‘Oc. (Fin.) prominens 
‘Oc. (Fin.) pseudotaeniatus 01 
‘Oc. (Fin.) pulchrithorax 
‘Oc. (Fin.) pulchriventer 
‘Oc. (Fin.) rubrithorax 
‘Oc. (Fin.) seoulensis 
‘Oc. (Fin.) sherki 
‘Oc. (Fin.) subalbitarsis 
‘Oc. (Fin.) tubbutiensis 
‘Oc. (Mac.) tremulus 
‘Oc. (Mol.) pecuniosus 01 
‘Oc. (Och.) atropalpus 01 
‘Oc. (Och.) deserticola 
‘Oc. (Och.) epactius 
‘Oc. (Och.) fluviatilis 
‘Oc. (Och.) muelleri 
‘Oc. (Och.) sierrensis 12 
‘Oc. (Och.) varipalpus 12 
‘Oc. (Pro.) berlini 01 
‘Oc. (Pro.) brelandi 
‘Oc. (Pro.) burgeri 01 
‘Oc. (Pro.) galindoi 01 
‘Oc. (Pro.) hendersoni 01 01 
‘Oc. (Pro.) homoeopus 
‘Oc. (Pro.) knabi 
‘Oc. (Pro.) kompi 
‘Oc. (Pro.) terrens 01 
‘Oc. (Pro.) triseriatus 01 
‘Oc. (Pro.) zavortinki 01 
‘Oc. (Pro.) zoosophus 
Op. (Not.) chathamicus 
Op. (Opi.) fuscus 
Ta. savoryi 01 01 
Ta. togoi 01 
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Cs.  (Cus.) inornata 01 12 
Cx. (Cux.) quinquefasciatus 
Ma. (Man.) titillans – 
Or. signifera 
Ab. papago 
Ae. cinereus 
Ae. esoensis 
Az. ramirezi 
Ge. baisasi 01 
Ge. longiforceps 
Gy. mediovittata 
Hw. sexlineata 01 
Hw. walkeri 01 01 01 
Ia. cavaticus 01 01 01 12 
Ke. dissimilis 
Ke. harbachi 01 
Ko. purpureipes 
Le. geoskusea 
Mu. alternans 01 
Mu. laniger 
‘Oc. (Brh.) alektorovi 01 01 01 
‘Oc. (Brh.) greenii 
‘Oc. (Cha.) wattensis 
Oc. (Och.) communis 01 
Oc. (Och.) infirmatus 
Oc. (Och.) scapularis 
‘Oc. (Psk.) bancroftianus 
‘Oc. (Psk.) postspiraculosus 01 
Oc. (Rus.) refiki 
Oc. (Rus.) rusticus 
Ps. (Gra.) jamaicensis 01 
Ps. (Jan.) ferox 12 
Ps. (Pso.) ciliata 01 01 
Rh. longirostris 
Rh. wardi 01 
Sc. albolineata 
Sc. arboricola 01 
St. aegypti 
St. africana 01 
Za. fulgens 01 
Za. longipalpis 
Do. leonis 12 
Do. nivea 12 
Fl. kochi 
Fl. poicilia 01 
Ha. australis 
Hg. (Con.) leucocelaenus 
Hg. (Con.) leucotaeniatus 
Hg. (Hag.) equinus 
Hg. (Hag.) splendens 
‘Oc. (Fin.) albilabris 
‘Oc. (Fin.) alboannulatus 
‘Oc. (Fin.) albocinctus 
‘Oc. (Fin.) albotaeniatus 
‘Oc. (Fin.) argenteitarsis 
‘Oc. (Fin.) argyronotum 01 01 01 
‘Oc. (Fin.) assamensis 
‘Oc. (Fin.) banksi 01 01 
‘Oc. (Fin.) biocellatus 
‘Oc. (Fin.) candidoscutellum 
‘Oc. (Fin.) chrysolineatus 
‘Oc. (Fin.) chungi 
‘Oc. (Fin.) crossi 
‘Oc. (Fin.) echinus 
‘Oc. (Fin.) elsiae 
‘Oc. (Fin.) embuensis 
‘Oc. (Fin.) geniculatus 01 
‘Oc. (Fin.) gilli 
‘Oc. (Fin.) gubernatoris 
‘Oc. (Fin.) harperi 
‘Oc. (Fin.) hatorii 
‘Oc. (Fin.) ingrami 
‘Oc. (Fin.) j. japonicus 
‘Oc. (Fin.) jugaraensis 
‘Oc. (Fin.) keefei 
‘Oc. (Fin.) loi 
‘Oc. (Fin.) lophoventralis 01 
‘Oc. (Fin.) luteostriatus 
‘Oc. (Fin.) macfarlanei 01 
‘Oc. (Fin.) notoscriptus 01 
‘Oc. (Fin.) oreophilus 
‘Oc. (Fin.) papuensis 
‘Oc. (Fin.) prominens 
‘Oc. (Fin.) pseudotaeniatus 01 
‘Oc. (Fin.) pulchrithorax 
‘Oc. (Fin.) pulchriventer 
‘Oc. (Fin.) rubrithorax 
‘Oc. (Fin.) seoulensis 
‘Oc. (Fin.) sherki 
‘Oc. (Fin.) subalbitarsis 
‘Oc. (Fin.) tubbutiensis 
‘Oc. (Mac.) tremulus 
‘Oc. (Mol.) pecuniosus 01 
‘Oc. (Och.) atropalpus 01 
‘Oc. (Och.) deserticola 
‘Oc. (Och.) epactius 
‘Oc. (Och.) fluviatilis 
‘Oc. (Och.) muelleri 
‘Oc. (Och.) sierrensis 12 
‘Oc. (Och.) varipalpus 12 
‘Oc. (Pro.) berlini 01 
‘Oc. (Pro.) brelandi 
‘Oc. (Pro.) burgeri 01 
‘Oc. (Pro.) galindoi 01 
‘Oc. (Pro.) hendersoni 01 01 
‘Oc. (Pro.) homoeopus 
‘Oc. (Pro.) knabi 
‘Oc. (Pro.) kompi 
‘Oc. (Pro.) terrens 01 
‘Oc. (Pro.) triseriatus 01 
‘Oc. (Pro.) zavortinki 01 
‘Oc. (Pro.) zoosophus 
Op. (Not.) chathamicus 
Op. (Opi.) fuscus 
Ta. savoryi 01 01 
Ta. togoi 01 
APPENDIX 2 Continued
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Cs.  (Cus.) inornata 01 12 
Cx. (Cux.) quinquefasciatus 
Ma. (Man.) titillans – 
Or. signifera 
Ab. papago 
Ae. cinereus 
Ae. esoensis 
Az. ramirezi 
Ge. baisasi 01 
Ge. longiforceps 
Gy. mediovittata 
Hw. sexlineata 01 
Hw. walkeri 01 01 01 
Ia. cavaticus 01 01 01 12 
Ke. dissimilis 
Ke. harbachi 01 
Ko. purpureipes 
Le. geoskusea 
Mu. alternans 01 
Mu. laniger 
‘Oc. (Brh.) alektorovi 01 01 01 
‘Oc. (Brh.) greenii 
‘Oc. (Cha.) wattensis 
Oc. (Och.) communis 01 
Oc. (Och.) infirmatus 
Oc. (Och.) scapularis 
‘Oc. (Psk.) bancroftianus 
‘Oc. (Psk.) postspiraculosus 01 
Oc. (Rus.) refiki 
Oc. (Rus.) rusticus 
Ps. (Gra.) jamaicensis 01 
Ps. (Jan.) ferox 12 
Ps. (Pso.) ciliata 01 01 
Rh. longirostris 
Rh. wardi 01 
Sc. albolineata 
Sc. arboricola 01 
St. aegypti 
St. africana 01 
Za. fulgens 01 
Za. longipalpis 
Do. leonis 12 
Do. nivea 12 
Fl. kochi 
Fl. poicilia 01 
Ha. australis 
Hg. (Con.) leucocelaenus 
Hg. (Con.) leucotaeniatus 
Hg. (Hag.) equinus 
Hg. (Hag.) splendens 
‘Oc. (Fin.) albilabris 
‘Oc. (Fin.) alboannulatus 
‘Oc. (Fin.) albocinctus 
‘Oc. (Fin.) albotaeniatus 
‘Oc. (Fin.) argenteitarsis 
‘Oc. (Fin.) argyronotum 01 01 01 
‘Oc. (Fin.) assamensis 
‘Oc. (Fin.) banksi 01 01 
‘Oc. (Fin.) biocellatus 
‘Oc. (Fin.) candidoscutellum 
‘Oc. (Fin.) chrysolineatus 
‘Oc. (Fin.) chungi 
‘Oc. (Fin.) crossi 
‘Oc. (Fin.) echinus 
‘Oc. (Fin.) elsiae 
‘Oc. (Fin.) embuensis 
‘Oc. (Fin.) geniculatus 01 
‘Oc. (Fin.) gilli 
‘Oc. (Fin.) gubernatoris 
‘Oc. (Fin.) harperi 
‘Oc. (Fin.) hatorii 
‘Oc. (Fin.) ingrami 
‘Oc. (Fin.) j. japonicus 
‘Oc. (Fin.) jugaraensis 
‘Oc. (Fin.) keefei 
‘Oc. (Fin.) loi 
‘Oc. (Fin.) lophoventralis 01 
‘Oc. (Fin.) luteostriatus 
‘Oc. (Fin.) macfarlanei 01 
‘Oc. (Fin.) notoscriptus 01 
‘Oc. (Fin.) oreophilus 
‘Oc. (Fin.) papuensis 
‘Oc. (Fin.) prominens 
‘Oc. (Fin.) pseudotaeniatus 01 
‘Oc. (Fin.) pulchrithorax 
‘Oc. (Fin.) pulchriventer 
‘Oc. (Fin.) rubrithorax 
‘Oc. (Fin.) seoulensis 
‘Oc. (Fin.) sherki 
‘Oc. (Fin.) subalbitarsis 
‘Oc. (Fin.) tubbutiensis 
‘Oc. (Mac.) tremulus 
‘Oc. (Mol.) pecuniosus 01 
‘Oc. (Och.) atropalpus 01 
‘Oc. (Och.) deserticola 
‘Oc. (Och.) epactius 
‘Oc. (Och.) fluviatilis 
‘Oc. (Och.) muelleri 
‘Oc. (Och.) sierrensis 12 
‘Oc. (Och.) varipalpus 12 
‘Oc. (Pro.) berlini 01 
‘Oc. (Pro.) brelandi 
‘Oc. (Pro.) burgeri 01 
‘Oc. (Pro.) galindoi 01 
‘Oc. (Pro.) hendersoni 01 01 
‘Oc. (Pro.) homoeopus 
‘Oc. (Pro.) knabi 
‘Oc. (Pro.) kompi 
‘Oc. (Pro.) terrens 01 
‘Oc. (Pro.) triseriatus 01 
‘Oc. (Pro.) zavortinki 01 
‘Oc. (Pro.) zoosophus 
Op. (Not.) chathamicus 
Op. (Opi.) fuscus 
Ta. savoryi 01 01 
Ta. togoi 01 
111111111111111111111
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Cs.  (Cus.) inornata 01 12 
Cx. (Cux.) quinquefasciatus 
Ma. (Man.) titillans – 
Or. signifera 
Ab. papago 
Ae. cinereus 
Ae. esoensis 
Az. ramirezi 
Ge. baisasi 01 
Ge. longiforceps 
Gy. mediovittata 
Hw. sexlineata 01 
Hw. walkeri 01 01 01 
Ia. cavaticus 01 01 01 12 
Ke. dissimilis 
Ke. harbachi 01 
Ko. purpureipes 
Le. geoskusea 
Mu. alternans 01 
Mu. laniger 
‘Oc. (Brh.) alektorovi 01 01 01 
‘Oc. (Brh.) greenii 
‘Oc. (Cha.) wattensis 
Oc. (Och.) communis 01 
Oc. (Och.) infirmatus 
Oc. (Och.) scapularis 
‘Oc. (Psk.) bancroftianus 
‘Oc. (Psk.) postspiraculosus 01 
Oc. (Rus.) refiki 
Oc. (Rus.) rusticus 
Ps. (Gra.) jamaicensis 01 
Ps. (Jan.) ferox 12 
Ps. (Pso.) ciliata 01 01 
Rh. longirostris 
Rh. wardi 01 
Sc. albolineata 
Sc. arboricola 01 
St. aegypti 
St. africana 01 
Za. fulgens 01 
Za. longipalpis 
Do. leonis 12 
Do. nivea 12 
Fl. kochi 
Fl. poicilia 01 
Ha. australis 
Hg. (Con.) leucocelaenus 
Hg. (Con.) leucotaeniatus 
Hg. (Hag.) equinus 
Hg. (Hag.) splendens 
‘Oc. (Fin.) albilabris 
‘Oc. (Fin.) alboannulatus 
‘Oc. (Fin.) albocinctus 
‘Oc. (Fin.) albotaeniatus 
‘Oc. (Fin.) argenteitarsis 
‘Oc. (Fin.) argyronotum 01 01 01 
‘Oc. (Fin.) assamensis 
‘Oc. (Fin.) banksi 01 01 
‘Oc. (Fin.) biocellatus 
‘Oc. (Fin.) candidoscutellum 
‘Oc. (Fin.) chrysolineatus 
‘Oc. (Fin.) chungi 
‘Oc. (Fin.) crossi 
‘Oc. (Fin.) echinus 
‘Oc. (Fin.) elsiae 
‘Oc. (Fin.) embuensis 
‘Oc. (Fin.) geniculatus 01 
‘Oc. (Fin.) gilli 
‘Oc. (Fin.) gubernatoris 
‘Oc. (Fin.) harperi 
‘Oc. (Fin.) hatorii 
‘Oc. (Fin.) ingrami 
‘Oc. (Fin.) j. japonicus 
‘Oc. (Fin.) jugaraensis 
‘Oc. (Fin.) keefei 
‘Oc. (Fin.) loi 
‘Oc. (Fin.) lophoventralis 01 
‘Oc. (Fin.) luteostriatus 
‘Oc. (Fin.) macfarlanei 01 
‘Oc. (Fin.) notoscriptus 01 
‘Oc. (Fin.) oreophilus 
‘Oc. (Fin.) papuensis 
‘Oc. (Fin.) prominens 
‘Oc. (Fin.) pseudotaeniatus 01 
‘Oc. (Fin.) pulchrithorax 
‘Oc. (Fin.) pulchriventer 
‘Oc. (Fin.) rubrithorax 
‘Oc. (Fin.) seoulensis 
‘Oc. (Fin.) sherki 
‘Oc. (Fin.) subalbitarsis 
‘Oc. (Fin.) tubbutiensis 
‘Oc. (Mac.) tremulus 
‘Oc. (Mol.) pecuniosus 01 
‘Oc. (Och.) atropalpus 01 
‘Oc. (Och.) deserticola 
‘Oc. (Och.) epactius 
‘Oc. (Och.) fluviatilis 
‘Oc. (Och.) muelleri 
‘Oc. (Och.) sierrensis 12 
‘Oc. (Och.) varipalpus 12 
‘Oc. (Pro.) berlini 01 
‘Oc. (Pro.) brelandi 
‘Oc. (Pro.) burgeri 01 
‘Oc. (Pro.) galindoi 01 
‘Oc. (Pro.) hendersoni 01 01 
‘Oc. (Pro.) homoeopus 
‘Oc. (Pro.) knabi 
‘Oc. (Pro.) kompi 
‘Oc. (Pro.) terrens 01 
‘Oc. (Pro.) triseriatus 01 
‘Oc. (Pro.) zavortinki 01 
‘Oc. (Pro.) zoosophus 
Op. (Not.) chathamicus 
Op. (Opi.) fuscus 
Ta. savoryi 01 01 
Ta. togoi 01 

APPENDIX 2 Continued
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Cs.  (Cus.) inornata 12 
Cx. (Cux.) quinquefasciatus 
Ma. (Man.) titillans 
Or. signifera 
Ab. papago 01 
Ae. cinereus 
Ae. esoensis 
Az. ramirezi 
Ge. baisasi 
Ge. longiforceps 12 
Gy. mediovittata 
Hw. sexlineata 
Hw. walkeri 12 
Ia. cavaticus 01 
Ke. dissimilis 01 
Ke. harbachi 
Ko. purpureipes 01 
Le. geoskusea 
Mu. alternans 
Mu. laniger 01 
‘Oc. (Brh.) alektorovi 01 01 01 
‘Oc. (Brh.) greenii 
‘Oc. (Cha.) wattensis 01 
Oc. (Och.) communis 01 01 12 12 01 
Oc. (Och.) infirmatus 01 
Oc. (Och.) scapularis 01 
‘Oc. (Psk.) bancroftianus 
‘Oc. (Psk.) postspiraculosus 
Oc. (Rus.) refiki 01 
Oc. (Rus.) rusticus 
Ps. (Gra.) jamaicensis 12 
Ps. (Jan.) ferox 
Ps. (Pso.) ciliata 01 
Rh. longirostris 01 
Rh. wardi 
Sc. albolineata 
Sc. arboricola 
St. aegypti 01 
St. africana 01 
Za. fulgens 
Za. longipalpis 
Do. leonis 
Do. nivea 
Fl. kochi 
Fl. poicilia 
Ha. australis 
Hg. (Con.) leucocelaenus 
Hg. (Con.) leucotaeniatus 01 
Hg. (Hag.) equinus 
Hg. (Hag.) splendens 
‘Oc. (Fin.) albilabris 01 
‘Oc. (Fin.) alboannulatus 
‘Oc. (Fin.) albocinctus 
‘Oc. (Fin.) albotaeniatus 
‘Oc. (Fin.) argenteitarsis 
‘Oc. (Fin.) argyronotum 
‘Oc. (Fin.) assamensis 01 
‘Oc. (Fin.) banksi 
‘Oc. (Fin.) biocellatus 
‘Oc. (Fin.) candidoscutellum 
‘Oc. (Fin.) chrysolineatus 01 
‘Oc. (Fin.) chungi 
‘Oc. (Fin.) crossi 
‘Oc. (Fin.) echinus 
‘Oc. (Fin.) elsiae 
‘Oc. (Fin.) embuensis 
‘Oc. (Fin.) geniculatus 
‘Oc. (Fin.) gilli 
‘Oc. (Fin.) gubernatoris 
‘Oc. (Fin.) harperi 01 
‘Oc. (Fin.) hatorii 
‘Oc. (Fin.) ingrami 
‘Oc. (Fin.) j. japonicus 
‘Oc. (Fin.) jugaraensis 
‘Oc. (Fin.) keefei 01 01 
‘Oc. (Fin.) loi 
‘Oc. (Fin.) lophoventralis 
‘Oc. (Fin.) luteostriatus 
‘Oc. (Fin.) macfarlanei 
‘Oc. (Fin.) notoscriptus 
‘Oc. (Fin.) oreophilus 
‘Oc. (Fin.) papuensis 
‘Oc. (Fin.) prominens 
‘Oc. (Fin.) pseudotaeniatus 
‘Oc. (Fin.) pulchrithorax 
‘Oc. (Fin.) pulchriventer 
‘Oc. (Fin.) rubrithorax 
‘Oc. (Fin.) seoulensis 
‘Oc. (Fin.) sherki 
‘Oc. (Fin.) subalbitarsis 
‘Oc. (Fin.) tubbutiensis 
‘Oc. (Mac.) tremulus 
‘Oc. (Mol.) pecuniosus 
‘Oc. (Och.) atropalpus 
‘Oc. (Och.) deserticola 
‘Oc. (Och.) epactius 
‘Oc. (Och.) fluviatilis 01 12 01 
‘Oc. (Och.) muelleri 
‘Oc. (Och.) sierrensis 
‘Oc. (Och.) varipalpus 02 12 
‘Oc. (Pro.) berlini 
‘Oc. (Pro.) brelandi 
‘Oc. (Pro.) burgeri 
‘Oc. (Pro.) galindoi 
‘Oc. (Pro.) hendersoni 
‘Oc. (Pro.) homoeopus 
‘Oc. (Pro.) knabi 01 
‘Oc. (Pro.) kompi 
‘Oc. (Pro.) terrens 
‘Oc. (Pro.) triseriatus 12 01 
‘Oc. (Pro.) zavortinki 
‘Oc. (Pro.) zoosophus 
Op. (Not.) chathamicus 
Op. (Opi.) fuscus 
Ta. savoryi 01 
Ta. togoi 01 
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Cs.  (Cus.) inornata 12 
Cx. (Cux.) quinquefasciatus 
Ma. (Man.) titillans 
Or. signifera 
Ab. papago 01 
Ae. cinereus 
Ae. esoensis 
Az. ramirezi 
Ge. baisasi 
Ge. longiforceps 12 
Gy. mediovittata 
Hw. sexlineata 
Hw. walkeri 12 
Ia. cavaticus 01 
Ke. dissimilis 01 
Ke. harbachi 
Ko. purpureipes 01 
Le. geoskusea 
Mu. alternans 
Mu. laniger 01 
‘Oc. (Brh.) alektorovi 01 01 01 
‘Oc. (Brh.) greenii 
‘Oc. (Cha.) wattensis 01 
Oc. (Och.) communis 01 01 12 12 01 
Oc. (Och.) infirmatus 01 
Oc. (Och.) scapularis 01 
‘Oc. (Psk.) bancroftianus 
‘Oc. (Psk.) postspiraculosus 
Oc. (Rus.) refiki 01 
Oc. (Rus.) rusticus 
Ps. (Gra.) jamaicensis 12 
Ps. (Jan.) ferox 
Ps. (Pso.) ciliata 01 
Rh. longirostris 01 
Rh. wardi 
Sc. albolineata 
Sc. arboricola 
St. aegypti 01 
St. africana 01 
Za. fulgens 
Za. longipalpis 
Do. leonis 
Do. nivea 
Fl. kochi 
Fl. poicilia 
Ha. australis 
Hg. (Con.) leucocelaenus 
Hg. (Con.) leucotaeniatus 01 
Hg. (Hag.) equinus 
Hg. (Hag.) splendens 
‘Oc. (Fin.) albilabris 01 
‘Oc. (Fin.) alboannulatus 
‘Oc. (Fin.) albocinctus 
‘Oc. (Fin.) albotaeniatus 
‘Oc. (Fin.) argenteitarsis 
‘Oc. (Fin.) argyronotum 
‘Oc. (Fin.) assamensis 01 
‘Oc. (Fin.) banksi 
‘Oc. (Fin.) biocellatus 
‘Oc. (Fin.) candidoscutellum 
‘Oc. (Fin.) chrysolineatus 01 
‘Oc. (Fin.) chungi 
‘Oc. (Fin.) crossi 
‘Oc. (Fin.) echinus 
‘Oc. (Fin.) elsiae 
‘Oc. (Fin.) embuensis 
‘Oc. (Fin.) geniculatus 
‘Oc. (Fin.) gilli 
‘Oc. (Fin.) gubernatoris 
‘Oc. (Fin.) harperi 01 
‘Oc. (Fin.) hatorii 
‘Oc. (Fin.) ingrami 
‘Oc. (Fin.) j. japonicus 
‘Oc. (Fin.) jugaraensis 
‘Oc. (Fin.) keefei 01 01 
‘Oc. (Fin.) loi 
‘Oc. (Fin.) lophoventralis 
‘Oc. (Fin.) luteostriatus 
‘Oc. (Fin.) macfarlanei 
‘Oc. (Fin.) notoscriptus 
‘Oc. (Fin.) oreophilus 
‘Oc. (Fin.) papuensis 
‘Oc. (Fin.) prominens 
‘Oc. (Fin.) pseudotaeniatus 
‘Oc. (Fin.) pulchrithorax 
‘Oc. (Fin.) pulchriventer 
‘Oc. (Fin.) rubrithorax 
‘Oc. (Fin.) seoulensis 
‘Oc. (Fin.) sherki 
‘Oc. (Fin.) subalbitarsis 
‘Oc. (Fin.) tubbutiensis 
‘Oc. (Mac.) tremulus 
‘Oc. (Mol.) pecuniosus 
‘Oc. (Och.) atropalpus 
‘Oc. (Och.) deserticola 
‘Oc. (Och.) epactius 
‘Oc. (Och.) fluviatilis 01 12 01 
‘Oc. (Och.) muelleri 
‘Oc. (Och.) sierrensis 
‘Oc. (Och.) varipalpus 02 12 
‘Oc. (Pro.) berlini 
‘Oc. (Pro.) brelandi 
‘Oc. (Pro.) burgeri 
‘Oc. (Pro.) galindoi 
‘Oc. (Pro.) hendersoni 
‘Oc. (Pro.) homoeopus 
‘Oc. (Pro.) knabi 01 
‘Oc. (Pro.) kompi 
‘Oc. (Pro.) terrens 
‘Oc. (Pro.) triseriatus 12 01 
‘Oc. (Pro.) zavortinki 
‘Oc. (Pro.) zoosophus 
Op. (Not.) chathamicus 
Op. (Opi.) fuscus 
Ta. savoryi 01 
Ta. togoi 01 
APPENDIX 2 Continued
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Cs.  (Cus.) inornata 12 
Cx. (Cux.) quinquefasciatus 
Ma. (Man.) titillans 
Or. signifera 
Ab. papago 01 
Ae. cinereus 
Ae. esoensis 
Az. ramirezi 
Ge. baisasi 
Ge. longiforceps 12 
Gy. mediovittata 
Hw. sexlineata 
Hw. walkeri 12 
Ia. cavaticus 01 
Ke. dissimilis 01 
Ke. harbachi 
Ko. purpureipes 01 
Le. geoskusea 
Mu. alternans 
Mu. laniger 01 
‘Oc. (Brh.) alektorovi 01 01 01 
‘Oc. (Brh.) greenii 
‘Oc. (Cha.) wattensis 01 
Oc. (Och.) communis 01 01 12 12 01 
Oc. (Och.) infirmatus 01 
Oc. (Och.) scapularis 01 
‘Oc. (Psk.) bancroftianus 
‘Oc. (Psk.) postspiraculosus 
Oc. (Rus.) refiki 01 
Oc. (Rus.) rusticus 
Ps. (Gra.) jamaicensis 12 
Ps. (Jan.) ferox 
Ps. (Pso.) ciliata 01 
Rh. longirostris 01 
Rh. wardi 
Sc. albolineata 
Sc. arboricola 
St. aegypti 01 
St. africana 01 
Za. fulgens 
Za. longipalpis 
Do. leonis 
Do. nivea 
Fl. kochi 
Fl. poicilia 
Ha. australis 
Hg. (Con.) leucocelaenus 
Hg. (Con.) leucotaeniatus 01 
Hg. (Hag.) equinus 
Hg. (Hag.) splendens 
‘Oc. (Fin.) albilabris 01 
‘Oc. (Fin.) alboannulatus 
‘Oc. (Fin.) albocinctus 
‘Oc. (Fin.) albotaeniatus 
‘Oc. (Fin.) argenteitarsis 
‘Oc. (Fin.) argyronotum 
‘Oc. (Fin.) assamensis 01 
‘Oc. (Fin.) banksi 
‘Oc. (Fin.) biocellatus 
‘Oc. (Fin.) candidoscutellum 
‘Oc. (Fin.) chrysolineatus 01 
‘Oc. (Fin.) chungi 
‘Oc. (Fin.) crossi 
‘Oc. (Fin.) echinus 
‘Oc. (Fin.) elsiae 
‘Oc. (Fin.) embuensis 
‘Oc. (Fin.) geniculatus 
‘Oc. (Fin.) gilli 
‘Oc. (Fin.) gubernatoris 
‘Oc. (Fin.) harperi 01 
‘Oc. (Fin.) hatorii 
‘Oc. (Fin.) ingrami 
‘Oc. (Fin.) j. japonicus 
‘Oc. (Fin.) jugaraensis 
‘Oc. (Fin.) keefei 01 01 
‘Oc. (Fin.) loi 
‘Oc. (Fin.) lophoventralis 
‘Oc. (Fin.) luteostriatus 
‘Oc. (Fin.) macfarlanei 
‘Oc. (Fin.) notoscriptus 
‘Oc. (Fin.) oreophilus 
‘Oc. (Fin.) papuensis 
‘Oc. (Fin.) prominens 
‘Oc. (Fin.) pseudotaeniatus 
‘Oc. (Fin.) pulchrithorax 
‘Oc. (Fin.) pulchriventer 
‘Oc. (Fin.) rubrithorax 
‘Oc. (Fin.) seoulensis 
‘Oc. (Fin.) sherki 
‘Oc. (Fin.) subalbitarsis 
‘Oc. (Fin.) tubbutiensis 
‘Oc. (Mac.) tremulus 
‘Oc. (Mol.) pecuniosus 
‘Oc. (Och.) atropalpus 
‘Oc. (Och.) deserticola 
‘Oc. (Och.) epactius 
‘Oc. (Och.) fluviatilis 01 12 01 
‘Oc. (Och.) muelleri 
‘Oc. (Och.) sierrensis 
‘Oc. (Och.) varipalpus 02 12 
‘Oc. (Pro.) berlini 
‘Oc. (Pro.) brelandi 
‘Oc. (Pro.) burgeri 
‘Oc. (Pro.) galindoi 
‘Oc. (Pro.) hendersoni 
‘Oc. (Pro.) homoeopus 
‘Oc. (Pro.) knabi 01 
‘Oc. (Pro.) kompi 
‘Oc. (Pro.) terrens 
‘Oc. (Pro.) triseriatus 12 01 
‘Oc. (Pro.) zavortinki 
‘Oc. (Pro.) zoosophus 
Op. (Not.) chathamicus 
Op. (Opi.) fuscus 
Ta. savoryi 01 
Ta. togoi 01 
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Cs.  (Cus.) inornata 12 
Cx. (Cux.) quinquefasciatus 
Ma. (Man.) titillans 
Or. signifera 
Ab. papago 01 
Ae. cinereus 
Ae. esoensis 
Az. ramirezi 
Ge. baisasi 
Ge. longiforceps 12 
Gy. mediovittata 
Hw. sexlineata 
Hw. walkeri 12 
Ia. cavaticus 01 
Ke. dissimilis 01 
Ke. harbachi 
Ko. purpureipes 01 
Le. geoskusea 
Mu. alternans 
Mu. laniger 01 
‘Oc. (Brh.) alektorovi 01 01 01 
‘Oc. (Brh.) greenii 
‘Oc. (Cha.) wattensis 01 
Oc. (Och.) communis 01 01 12 12 01 
Oc. (Och.) infirmatus 01 
Oc. (Och.) scapularis 01 
‘Oc. (Psk.) bancroftianus 
‘Oc. (Psk.) postspiraculosus 
Oc. (Rus.) refiki 01 
Oc. (Rus.) rusticus 
Ps. (Gra.) jamaicensis 12 
Ps. (Jan.) ferox 
Ps. (Pso.) ciliata 01 
Rh. longirostris 01 
Rh. wardi 
Sc. albolineata 
Sc. arboricola 
St. aegypti 01 
St. africana 01 
Za. fulgens 
Za. longipalpis 
Do. leonis 
Do. nivea 
Fl. kochi 
Fl. poicilia 
Ha. australis 
Hg. (Con.) leucocelaenus 
Hg. (Con.) leucotaeniatus 01 
Hg. (Hag.) equinus 
Hg. (Hag.) splendens 
‘Oc. (Fin.) albilabris 01 
‘Oc. (Fin.) alboannulatus 
‘Oc. (Fin.) albocinctus 
‘Oc. (Fin.) albotaeniatus 
‘Oc. (Fin.) argenteitarsis 
‘Oc. (Fin.) argyronotum 
‘Oc. (Fin.) assamensis 01 
‘Oc. (Fin.) banksi 
‘Oc. (Fin.) biocellatus 
‘Oc. (Fin.) candidoscutellum 
‘Oc. (Fin.) chrysolineatus 01 
‘Oc. (Fin.) chungi 
‘Oc. (Fin.) crossi 
‘Oc. (Fin.) echinus 
‘Oc. (Fin.) elsiae 
‘Oc. (Fin.) embuensis 
‘Oc. (Fin.) geniculatus 
‘Oc. (Fin.) gilli 
‘Oc. (Fin.) gubernatoris 
‘Oc. (Fin.) harperi 01 
‘Oc. (Fin.) hatorii 
‘Oc. (Fin.) ingrami 
‘Oc. (Fin.) j. japonicus 
‘Oc. (Fin.) jugaraensis 
‘Oc. (Fin.) keefei 01 01 
‘Oc. (Fin.) loi 
‘Oc. (Fin.) lophoventralis 
‘Oc. (Fin.) luteostriatus 
‘Oc. (Fin.) macfarlanei 
‘Oc. (Fin.) notoscriptus 
‘Oc. (Fin.) oreophilus 
‘Oc. (Fin.) papuensis 
‘Oc. (Fin.) prominens 
‘Oc. (Fin.) pseudotaeniatus 
‘Oc. (Fin.) pulchrithorax 
‘Oc. (Fin.) pulchriventer 
‘Oc. (Fin.) rubrithorax 
‘Oc. (Fin.) seoulensis 
‘Oc. (Fin.) sherki 
‘Oc. (Fin.) subalbitarsis 
‘Oc. (Fin.) tubbutiensis 
‘Oc. (Mac.) tremulus 
‘Oc. (Mol.) pecuniosus 
‘Oc. (Och.) atropalpus 
‘Oc. (Och.) deserticola 
‘Oc. (Och.) epactius 
‘Oc. (Och.) fluviatilis 01 12 01 
‘Oc. (Och.) muelleri 
‘Oc. (Och.) sierrensis 
‘Oc. (Och.) varipalpus 02 12 
‘Oc. (Pro.) berlini 
‘Oc. (Pro.) brelandi 
‘Oc. (Pro.) burgeri 
‘Oc. (Pro.) galindoi 
‘Oc. (Pro.) hendersoni 
‘Oc. (Pro.) homoeopus 
‘Oc. (Pro.) knabi 01 
‘Oc. (Pro.) kompi 
‘Oc. (Pro.) terrens 
‘Oc. (Pro.) triseriatus 12 01 
‘Oc. (Pro.) zavortinki 
‘Oc. (Pro.) zoosophus 
Op. (Not.) chathamicus 
Op. (Opi.) fuscus 
Ta. savoryi 01 
Ta. togoi 01 

APPENDIX 2 Continued
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Cs.  (Cus.) inornata 
Cx. (Cux.) quinquefasciatus 
Ma. (Man.) titillans 
Or. signifera 
Ab. papago 
Ae. cinereus 
Ae. esoensis 
Az. ramirezi 01 
Ge. baisasi 
Ge. longiforceps 
Gy. mediovittata 01 
Hw. sexlineata 
Hw. walkeri 01 
Ia. cavaticus 
Ke. dissimilis 01 
Ke. harbachi 01 
Ko. purpureipes 
Le. geoskusea 
Mu. alternans 
Mu. laniger 01 
‘Oc. (Brh.) alektorovi 
‘Oc. (Brh.) greenii 01 
‘Oc. (Cha.) wattensis 
Oc. (Och.) communis 
Oc. (Och.) infirmatus 
Oc. (Och.) scapularis 
‘Oc. (Psk.) bancroftianus 01 
‘Oc. (Psk.) postspiraculosus 
Oc. (Rus.) refiki 
Oc. (Rus.) rusticus 
Ps. (Gra.) jamaicensis 
Ps. (Jan.) ferox 01 
Ps. (Pso.) ciliata 
Rh. longirostris 01 01 
Rh. wardi 01 
Sc. albolineata 
Sc. arboricola 
St. aegypti 
St. africana 
Za. fulgens 
Za. longipalpis 
Do. leonis 
Do. nivea 
Fl. kochi 
Fl. poicilia 
Ha. australis 
Hg. (Con.) leucocelaenus 
Hg. (Con.) leucotaeniatus 12 
Hg. (Hag.) equinus 01 
Hg. (Hag.) splendens 
‘Oc. (Fin.) albilabris 
‘Oc. (Fin.) alboannulatus 
‘Oc. (Fin.) albocinctus 
‘Oc. (Fin.) albotaeniatus 01 
‘Oc. (Fin.) argenteitarsis 
‘Oc. (Fin.) argyronotum 
‘Oc. (Fin.) assamensis 
‘Oc. (Fin.) banksi 
‘Oc. (Fin.) biocellatus 
‘Oc. (Fin.) candidoscutellum 
‘Oc. (Fin.) chrysolineatus 
‘Oc. (Fin.) chungi 
‘Oc. (Fin.) crossi 
‘Oc. (Fin.) echinus 
‘Oc. (Fin.) elsiae 
‘Oc. (Fin.) embuensis 
‘Oc. (Fin.) geniculatus 
‘Oc. (Fin.) gilli 
‘Oc. (Fin.) gubernatoris 
‘Oc. (Fin.) harperi 01 
‘Oc. (Fin.) hatorii 
‘Oc. (Fin.) ingrami 01 
‘Oc. (Fin.) j. japonicus 
‘Oc. (Fin.) jugaraensis 
‘Oc. (Fin.) keefei 
‘Oc. (Fin.) loi 
‘Oc. (Fin.) lophoventralis 
‘Oc. (Fin.) luteostriatus 
‘Oc. (Fin.) macfarlanei 
‘Oc. (Fin.) notoscriptus 
‘Oc. (Fin.) oreophilus 
‘Oc. (Fin.) papuensis 
‘Oc. (Fin.) prominens 
‘Oc. (Fin.) pseudotaeniatus 
‘Oc. (Fin.) pulchrithorax 
‘Oc. (Fin.) pulchriventer 
‘Oc. (Fin.) rubrithorax 
‘Oc. (Fin.) seoulensis 
‘Oc. (Fin.) sherki 
‘Oc. (Fin.) subalbitarsis 
‘Oc. (Fin.) tubbutiensis 01 
‘Oc. (Mac.) tremulus 
‘Oc. (Mol.) pecuniosus 01 
‘Oc. (Och.) atropalpus 
‘Oc. (Och.) deserticola 
‘Oc. (Och.) epactius 
‘Oc. (Och.) fluviatilis 
‘Oc. (Och.) muelleri 01 
‘Oc. (Och.) sierrensis 
‘Oc. (Och.) varipalpus 01 
‘Oc. (Pro.) berlini 
‘Oc. (Pro.) brelandi 
‘Oc. (Pro.) burgeri 
‘Oc. (Pro.) galindoi 
‘Oc. (Pro.) hendersoni 
‘Oc. (Pro.) homoeopus 
‘Oc. (Pro.) knabi 01 
‘Oc. (Pro.) kompi 01 
‘Oc. (Pro.) terrens 
‘Oc. (Pro.) triseriatus 
‘Oc. (Pro.) zavortinki 
‘Oc. (Pro.) zoosophus 
Op. (Not.) chathamicus 01 
Op. (Opi.) fuscus 01 01 
Ta. savoryi 
Ta. togoi 
1111111111111111111111111111111111111112
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Cs.  (Cus.) inornata 
Cx. (Cux.) quinquefasciatus 
Ma. (Man.) titillans 
Or. signifera 
Ab. papago 
Ae. cinereus 
Ae. esoensis 
Az. ramirezi 01 
Ge. baisasi 
Ge. longiforceps 
Gy. mediovittata 01 
Hw. sexlineata 
Hw. walkeri 01 
Ia. cavaticus 
Ke. dissimilis 01 
Ke. harbachi 01 
Ko. purpureipes 
Le. geoskusea 
Mu. alternans 
Mu. laniger 01 
‘Oc. (Brh.) alektorovi 
‘Oc. (Brh.) greenii 01 
‘Oc. (Cha.) wattensis 
Oc. (Och.) communis 
Oc. (Och.) infirmatus 
Oc. (Och.) scapularis 
‘Oc. (Psk.) bancroftianus 01 
‘Oc. (Psk.) postspiraculosus 
Oc. (Rus.) refiki 
Oc. (Rus.) rusticus 
Ps. (Gra.) jamaicensis 
Ps. (Jan.) ferox 01 
Ps. (Pso.) ciliata 
Rh. longirostris 01 01 
Rh. wardi 01 
Sc. albolineata 
Sc. arboricola 
St. aegypti 
St. africana 
Za. fulgens 
Za. longipalpis 
Do. leonis 
Do. nivea 
Fl. kochi 
Fl. poicilia 
Ha. australis 
Hg. (Con.) leucocelaenus 
Hg. (Con.) leucotaeniatus 12 
Hg. (Hag.) equinus 01 
Hg. (Hag.) splendens 
‘Oc. (Fin.) albilabris 
‘Oc. (Fin.) alboannulatus 
‘Oc. (Fin.) albocinctus 
‘Oc. (Fin.) albotaeniatus 01 
‘Oc. (Fin.) argenteitarsis 
‘Oc. (Fin.) argyronotum 
‘Oc. (Fin.) assamensis 
‘Oc. (Fin.) banksi 
‘Oc. (Fin.) biocellatus 
‘Oc. (Fin.) candidoscutellum 
‘Oc. (Fin.) chrysolineatus 
‘Oc. (Fin.) chungi 
‘Oc. (Fin.) crossi 
‘Oc. (Fin.) echinus 
‘Oc. (Fin.) elsiae 
‘Oc. (Fin.) embuensis 
‘Oc. (Fin.) geniculatus 
‘Oc. (Fin.) gilli 
‘Oc. (Fin.) gubernatoris 
‘Oc. (Fin.) harperi 01 
‘Oc. (Fin.) hatorii 
‘Oc. (Fin.) ingrami 01 
‘Oc. (Fin.) j. japonicus 
‘Oc. (Fin.) jugaraensis 
‘Oc. (Fin.) keefei 
‘Oc. (Fin.) loi 
‘Oc. (Fin.) lophoventralis 
‘Oc. (Fin.) luteostriatus 
‘Oc. (Fin.) macfarlanei 
‘Oc. (Fin.) notoscriptus 
‘Oc. (Fin.) oreophilus 
‘Oc. (Fin.) papuensis 
‘Oc. (Fin.) prominens 
‘Oc. (Fin.) pseudotaeniatus 
‘Oc. (Fin.) pulchrithorax 
‘Oc. (Fin.) pulchriventer 
‘Oc. (Fin.) rubrithorax 
‘Oc. (Fin.) seoulensis 
‘Oc. (Fin.) sherki 
‘Oc. (Fin.) subalbitarsis 
‘Oc. (Fin.) tubbutiensis 01 
‘Oc. (Mac.) tremulus 
‘Oc. (Mol.) pecuniosus 01 
‘Oc. (Och.) atropalpus 
‘Oc. (Och.) deserticola 
‘Oc. (Och.) epactius 
‘Oc. (Och.) fluviatilis 
‘Oc. (Och.) muelleri 01 
‘Oc. (Och.) sierrensis 
‘Oc. (Och.) varipalpus 01 
‘Oc. (Pro.) berlini 
‘Oc. (Pro.) brelandi 
‘Oc. (Pro.) burgeri 
‘Oc. (Pro.) galindoi 
‘Oc. (Pro.) hendersoni 
‘Oc. (Pro.) homoeopus 
‘Oc. (Pro.) knabi 01 
‘Oc. (Pro.) kompi 01 
‘Oc. (Pro.) terrens 
‘Oc. (Pro.) triseriatus 
‘Oc. (Pro.) zavortinki 
‘Oc. (Pro.) zoosophus 
Op. (Not.) chathamicus 01 
Op. (Opi.) fuscus 01 01 
Ta. savoryi 
Ta. togoi 
APPENDIX 2 Continued
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Cs.  (Cus.) inornata 
Cx. (Cux.) quinquefasciatus 
Ma. (Man.) titillans 
Or. signifera 
Ab. papago 
Ae. cinereus 
Ae. esoensis 
Az. ramirezi 01 
Ge. baisasi 
Ge. longiforceps 
Gy. mediovittata 01 
Hw. sexlineata 
Hw. walkeri 01 
Ia. cavaticus 
Ke. dissimilis 01 
Ke. harbachi 01 
Ko. purpureipes 
Le. geoskusea 
Mu. alternans 
Mu. laniger 01 
‘Oc. (Brh.) alektorovi 
‘Oc. (Brh.) greenii 01 
‘Oc. (Cha.) wattensis 
Oc. (Och.) communis 
Oc. (Och.) infirmatus 
Oc. (Och.) scapularis 
‘Oc. (Psk.) bancroftianus 01 
‘Oc. (Psk.) postspiraculosus 
Oc. (Rus.) refiki 
Oc. (Rus.) rusticus 
Ps. (Gra.) jamaicensis 
Ps. (Jan.) ferox 01 
Ps. (Pso.) ciliata 
Rh. longirostris 01 01 
Rh. wardi 01 
Sc. albolineata 
Sc. arboricola 
St. aegypti 
St. africana 
Za. fulgens 
Za. longipalpis 
Do. leonis 
Do. nivea 
Fl. kochi 
Fl. poicilia 
Ha. australis 
Hg. (Con.) leucocelaenus 
Hg. (Con.) leucotaeniatus 12 
Hg. (Hag.) equinus 01 
Hg. (Hag.) splendens 
‘Oc. (Fin.) albilabris 
‘Oc. (Fin.) alboannulatus 
‘Oc. (Fin.) albocinctus 
‘Oc. (Fin.) albotaeniatus 01 
‘Oc. (Fin.) argenteitarsis 
‘Oc. (Fin.) argyronotum 
‘Oc. (Fin.) assamensis 
‘Oc. (Fin.) banksi 
‘Oc. (Fin.) biocellatus 
‘Oc. (Fin.) candidoscutellum 
‘Oc. (Fin.) chrysolineatus 
‘Oc. (Fin.) chungi 
‘Oc. (Fin.) crossi 
‘Oc. (Fin.) echinus 
‘Oc. (Fin.) elsiae 
‘Oc. (Fin.) embuensis 
‘Oc. (Fin.) geniculatus 
‘Oc. (Fin.) gilli 
‘Oc. (Fin.) gubernatoris 
‘Oc. (Fin.) harperi 01 
‘Oc. (Fin.) hatorii 
‘Oc. (Fin.) ingrami 01 
‘Oc. (Fin.) j. japonicus 
‘Oc. (Fin.) jugaraensis 
‘Oc. (Fin.) keefei 
‘Oc. (Fin.) loi 
‘Oc. (Fin.) lophoventralis 
‘Oc. (Fin.) luteostriatus 
‘Oc. (Fin.) macfarlanei 
‘Oc. (Fin.) notoscriptus 
‘Oc. (Fin.) oreophilus 
‘Oc. (Fin.) papuensis 
‘Oc. (Fin.) prominens 
‘Oc. (Fin.) pseudotaeniatus 
‘Oc. (Fin.) pulchrithorax 
‘Oc. (Fin.) pulchriventer 
‘Oc. (Fin.) rubrithorax 
‘Oc. (Fin.) seoulensis 
‘Oc. (Fin.) sherki 
‘Oc. (Fin.) subalbitarsis 
‘Oc. (Fin.) tubbutiensis 01 
‘Oc. (Mac.) tremulus 
‘Oc. (Mol.) pecuniosus 01 
‘Oc. (Och.) atropalpus 
‘Oc. (Och.) deserticola 
‘Oc. (Och.) epactius 
‘Oc. (Och.) fluviatilis 
‘Oc. (Och.) muelleri 01 
‘Oc. (Och.) sierrensis 
‘Oc. (Och.) varipalpus 01 
‘Oc. (Pro.) berlini 
‘Oc. (Pro.) brelandi 
‘Oc. (Pro.) burgeri 
‘Oc. (Pro.) galindoi 
‘Oc. (Pro.) hendersoni 
‘Oc. (Pro.) homoeopus 
‘Oc. (Pro.) knabi 01 
‘Oc. (Pro.) kompi 01 
‘Oc. (Pro.) terrens 
‘Oc. (Pro.) triseriatus 
‘Oc. (Pro.) zavortinki 
‘Oc. (Pro.) zoosophus 
Op. (Not.) chathamicus 01 
Op. (Opi.) fuscus 01 01 
Ta. savoryi 
Ta. togoi 
1111111111111111111111111111111111111112
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Cs.  (Cus.) inornata 
Cx. (Cux.) quinquefasciatus 
Ma. (Man.) titillans 
Or. signifera 
Ab. papago 
Ae. cinereus 
Ae. esoensis 
Az. ramirezi 01 
Ge. baisasi 
Ge. longiforceps 
Gy. mediovittata 01 
Hw. sexlineata 
Hw. walkeri 01 
Ia. cavaticus 
Ke. dissimilis 01 
Ke. harbachi 01 
Ko. purpureipes 
Le. geoskusea 
Mu. alternans 
Mu. laniger 01 
‘Oc. (Brh.) alektorovi 
‘Oc. (Brh.) greenii 01 
‘Oc. (Cha.) wattensis 
Oc. (Och.) communis 
Oc. (Och.) infirmatus 
Oc. (Och.) scapularis 
‘Oc. (Psk.) bancroftianus 01 
‘Oc. (Psk.) postspiraculosus 
Oc. (Rus.) refiki 
Oc. (Rus.) rusticus 
Ps. (Gra.) jamaicensis 
Ps. (Jan.) ferox 01 
Ps. (Pso.) ciliata 
Rh. longirostris 01 01 
Rh. wardi 01 
Sc. albolineata 
Sc. arboricola 
St. aegypti 
St. africana 
Za. fulgens 
Za. longipalpis 
Do. leonis 
Do. nivea 
Fl. kochi 
Fl. poicilia 
Ha. australis 
Hg. (Con.) leucocelaenus 
Hg. (Con.) leucotaeniatus 12 
Hg. (Hag.) equinus 01 
Hg. (Hag.) splendens 
‘Oc. (Fin.) albilabris 
‘Oc. (Fin.) alboannulatus 
‘Oc. (Fin.) albocinctus 
‘Oc. (Fin.) albotaeniatus 01 
‘Oc. (Fin.) argenteitarsis 
‘Oc. (Fin.) argyronotum 
‘Oc. (Fin.) assamensis 
‘Oc. (Fin.) banksi 
‘Oc. (Fin.) biocellatus 
‘Oc. (Fin.) candidoscutellum 
‘Oc. (Fin.) chrysolineatus 
‘Oc. (Fin.) chungi 
‘Oc. (Fin.) crossi 
‘Oc. (Fin.) echinus 
‘Oc. (Fin.) elsiae 
‘Oc. (Fin.) embuensis 
‘Oc. (Fin.) geniculatus 
‘Oc. (Fin.) gilli 
‘Oc. (Fin.) gubernatoris 
‘Oc. (Fin.) harperi 01 
‘Oc. (Fin.) hatorii 
‘Oc. (Fin.) ingrami 01 
‘Oc. (Fin.) j. japonicus 
‘Oc. (Fin.) jugaraensis 
‘Oc. (Fin.) keefei 
‘Oc. (Fin.) loi 
‘Oc. (Fin.) lophoventralis 
‘Oc. (Fin.) luteostriatus 
‘Oc. (Fin.) macfarlanei 
‘Oc. (Fin.) notoscriptus 
‘Oc. (Fin.) oreophilus 
‘Oc. (Fin.) papuensis 
‘Oc. (Fin.) prominens 
‘Oc. (Fin.) pseudotaeniatus 
‘Oc. (Fin.) pulchrithorax 
‘Oc. (Fin.) pulchriventer 
‘Oc. (Fin.) rubrithorax 
‘Oc. (Fin.) seoulensis 
‘Oc. (Fin.) sherki 
‘Oc. (Fin.) subalbitarsis 
‘Oc. (Fin.) tubbutiensis 01 
‘Oc. (Mac.) tremulus 
‘Oc. (Mol.) pecuniosus 01 
‘Oc. (Och.) atropalpus 
‘Oc. (Och.) deserticola 
‘Oc. (Och.) epactius 
‘Oc. (Och.) fluviatilis 
‘Oc. (Och.) muelleri 01 
‘Oc. (Och.) sierrensis 
‘Oc. (Och.) varipalpus 01 
‘Oc. (Pro.) berlini 
‘Oc. (Pro.) brelandi 
‘Oc. (Pro.) burgeri 
‘Oc. (Pro.) galindoi 
‘Oc. (Pro.) hendersoni 
‘Oc. (Pro.) homoeopus 
‘Oc. (Pro.) knabi 01 
‘Oc. (Pro.) kompi 01 
‘Oc. (Pro.) terrens 
‘Oc. (Pro.) triseriatus 
‘Oc. (Pro.) zavortinki 
‘Oc. (Pro.) zoosophus 
Op. (Not.) chathamicus 01 
Op. (Opi.) fuscus 01 01 
Ta. savoryi 
Ta. togoi 

APPENDIX 2 Continued
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Cs.  (Cus.) inornata 
Cx. (Cux.) quinquefasciatus – – – 03 
Ma. (Man.) titillans 01 
Or. signifera 01 
Ab. papago 
Ae. cinereus – 
Ae. esoensis – 
Az. ramirezi 
Ge. baisasi 
Ge. longiforceps 01 
Gy. mediovittata 
Hw. sexlineata 
Hw. walkeri 
Ia. cavaticus 
Ke. dissimilis 01 
Ke. harbachi 01 
Ko. purpureipes 
Le. geoskusea 
Mu. alternans 01 
Mu. laniger 
‘Oc. (Brh.) alektorovi 
‘Oc. (Brh.) greenii 
‘Oc. (Cha.) wattensis 
Oc. (Och.) communis 
Oc. (Och.) infirmatus 
Oc. (Och.) scapularis 
‘Oc. (Psk.) bancroftianus 
‘Oc. (Psk.) postspiraculosus 
Oc. (Rus.) refiki 
Oc. (Rus.) rusticus 
Ps. (Gra.) jamaicensis 
Ps. (Jan.) ferox 
Ps. (Pso.) ciliata 
Rh. longirostris 
Rh. wardi 01 
Sc. albolineata 
Sc. arboricola 
St. aegypti 
St. africana 
Za. fulgens 12 
Za. longipalpis 01 
Do. leonis 
Do. nivea 
Fl. kochi 
Fl. poicilia 
Ha. australis 12 
Hg. (Con.) leucocelaenus 
Hg. (Con.) leucotaeniatus 
Hg. (Hag.) equinus 
Hg. (Hag.) splendens 
‘Oc. (Fin.) albilabris 
‘Oc. (Fin.) alboannulatus 02 
‘Oc. (Fin.) albocinctus 
‘Oc. (Fin.) albotaeniatus 
‘Oc. (Fin.) argenteitarsis 
‘Oc. (Fin.) argyronotum 23 
‘Oc. (Fin.) assamensis 
‘Oc. (Fin.) banksi 
‘Oc. (Fin.) biocellatus 
‘Oc. (Fin.) candidoscutellum 23 
‘Oc. (Fin.) chrysolineatus 
‘Oc. (Fin.) chungi 
‘Oc. (Fin.) crossi 
‘Oc. (Fin.) echinus 
‘Oc. (Fin.) elsiae 23 
‘Oc. (Fin.) embuensis 
‘Oc. (Fin.) geniculatus 
‘Oc. (Fin.) gilli 
‘Oc. (Fin.) gubernatoris 
‘Oc. (Fin.) harperi 
‘Oc. (Fin.) hatorii 
‘Oc. (Fin.) ingrami 
‘Oc. (Fin.) j. japonicus 
‘Oc. (Fin.) jugaraensis 
‘Oc. (Fin.) keefei 
‘Oc. (Fin.) loi 
‘Oc. (Fin.) lophoventralis 
‘Oc. (Fin.) luteostriatus 
‘Oc. (Fin.) macfarlanei 23 
‘Oc. (Fin.) notoscriptus 
‘Oc. (Fin.) oreophilus 
‘Oc. (Fin.) papuensis 23 
‘Oc. (Fin.) prominens 
‘Oc. (Fin.) pseudotaeniatus 23 
‘Oc. (Fin.) pulchrithorax 
‘Oc. (Fin.) pulchriventer 23 
‘Oc. (Fin.) rubrithorax 23 
‘Oc. (Fin.) seoulensis 
‘Oc. (Fin.) sherki 
‘Oc. (Fin.) subalbitarsis 23 
‘Oc. (Fin.) tubbutiensis 
‘Oc. (Mac.) tremulus 
‘Oc. (Mol.) pecuniosus 
‘Oc. (Och.) atropalpus 
‘Oc. (Och.) deserticola 
‘Oc. (Och.) epactius 
‘Oc. (Och.) fluviatilis 23 
‘Oc. (Och.) muelleri 01 
‘Oc. (Och.) sierrensis 
‘Oc. (Och.) varipalpus 
‘Oc. (Pro.) berlini 
‘Oc. (Pro.) brelandi 
‘Oc. (Pro.) burgeri 12 
‘Oc. (Pro.) galindoi 
‘Oc. (Pro.) hendersoni 
‘Oc. (Pro.) homoeopus 
‘Oc. (Pro.) knabi 
‘Oc. (Pro.) kompi 
‘Oc. (Pro.) terrens 
‘Oc. (Pro.) triseriatus 
‘Oc. (Pro.) zavortinki 
‘Oc. (Pro.) zoosophus 
Op. (Not.) chathamicus 
Op. (Opi.) fuscus 
Ta. savoryi 
Ta. togoi 
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Cs.  (Cus.) inornata 
Cx. (Cux.) quinquefasciatus – – – 03 
Ma. (Man.) titillans 01 
Or. signifera 01 
Ab. papago 
Ae. cinereus – 
Ae. esoensis – 
Az. ramirezi 
Ge. baisasi 
Ge. longiforceps 01 
Gy. mediovittata 
Hw. sexlineata 
Hw. walkeri 
Ia. cavaticus 
Ke. dissimilis 01 
Ke. harbachi 01 
Ko. purpureipes 
Le. geoskusea 
Mu. alternans 01 
Mu. laniger 
‘Oc. (Brh.) alektorovi 
‘Oc. (Brh.) greenii 
‘Oc. (Cha.) wattensis 
Oc. (Och.) communis 
Oc. (Och.) infirmatus 
Oc. (Och.) scapularis 
‘Oc. (Psk.) bancroftianus 
‘Oc. (Psk.) postspiraculosus 
Oc. (Rus.) refiki 
Oc. (Rus.) rusticus 
Ps. (Gra.) jamaicensis 
Ps. (Jan.) ferox 
Ps. (Pso.) ciliata 
Rh. longirostris 
Rh. wardi 01 
Sc. albolineata 
Sc. arboricola 
St. aegypti 
St. africana 
Za. fulgens 12 
Za. longipalpis 01 
Do. leonis 
Do. nivea 
Fl. kochi 
Fl. poicilia 
Ha. australis 12 
Hg. (Con.) leucocelaenus 
Hg. (Con.) leucotaeniatus 
Hg. (Hag.) equinus 
Hg. (Hag.) splendens 
‘Oc. (Fin.) albilabris 
‘Oc. (Fin.) alboannulatus 02 
‘Oc. (Fin.) albocinctus 
‘Oc. (Fin.) albotaeniatus 
‘Oc. (Fin.) argenteitarsis 
‘Oc. (Fin.) argyronotum 23 
‘Oc. (Fin.) assamensis 
‘Oc. (Fin.) banksi 
‘Oc. (Fin.) biocellatus 
‘Oc. (Fin.) candidoscutellum 23 
‘Oc. (Fin.) chrysolineatus 
‘Oc. (Fin.) chungi 
‘Oc. (Fin.) crossi 
‘Oc. (Fin.) echinus 
‘Oc. (Fin.) elsiae 23 
‘Oc. (Fin.) embuensis 
‘Oc. (Fin.) geniculatus 
‘Oc. (Fin.) gilli 
‘Oc. (Fin.) gubernatoris 
‘Oc. (Fin.) harperi 
‘Oc. (Fin.) hatorii 
‘Oc. (Fin.) ingrami 
‘Oc. (Fin.) j. japonicus 
‘Oc. (Fin.) jugaraensis 
‘Oc. (Fin.) keefei 
‘Oc. (Fin.) loi 
‘Oc. (Fin.) lophoventralis 
‘Oc. (Fin.) luteostriatus 
‘Oc. (Fin.) macfarlanei 23 
‘Oc. (Fin.) notoscriptus 
‘Oc. (Fin.) oreophilus 
‘Oc. (Fin.) papuensis 23 
‘Oc. (Fin.) prominens 
‘Oc. (Fin.) pseudotaeniatus 23 
‘Oc. (Fin.) pulchrithorax 
‘Oc. (Fin.) pulchriventer 23 
‘Oc. (Fin.) rubrithorax 23 
‘Oc. (Fin.) seoulensis 
‘Oc. (Fin.) sherki 
‘Oc. (Fin.) subalbitarsis 23 
‘Oc. (Fin.) tubbutiensis 
‘Oc. (Mac.) tremulus 
‘Oc. (Mol.) pecuniosus 
‘Oc. (Och.) atropalpus 
‘Oc. (Och.) deserticola 
‘Oc. (Och.) epactius 
‘Oc. (Och.) fluviatilis 23 
‘Oc. (Och.) muelleri 01 
‘Oc. (Och.) sierrensis 
‘Oc. (Och.) varipalpus 
‘Oc. (Pro.) berlini 
‘Oc. (Pro.) brelandi 
‘Oc. (Pro.) burgeri 12 
‘Oc. (Pro.) galindoi 
‘Oc. (Pro.) hendersoni 
‘Oc. (Pro.) homoeopus 
‘Oc. (Pro.) knabi 
‘Oc. (Pro.) kompi 
‘Oc. (Pro.) terrens 
‘Oc. (Pro.) triseriatus 
‘Oc. (Pro.) zavortinki 
‘Oc. (Pro.) zoosophus 
Op. (Not.) chathamicus 
Op. (Opi.) fuscus 
Ta. savoryi 
Ta. togoi 
APPENDIX 2 Continued
22222222222222222222222222222222
00000000011111111112222222222333
12345678901234567890123456789012
Cs.  (Cus.) inornata 
Cx. (Cux.) quinquefasciatus – – – 03 
Ma. (Man.) titillans 01 
Or. signifera 01 
Ab. papago 
Ae. cinereus – 
Ae. esoensis – 
Az. ramirezi 
Ge. baisasi 
Ge. longiforceps 01 
Gy. mediovittata 
Hw. sexlineata 
Hw. walkeri 
Ia. cavaticus 
Ke. dissimilis 01 
Ke. harbachi 01 
Ko. purpureipes 
Le. geoskusea 
Mu. alternans 01 
Mu. laniger 
‘Oc. (Brh.) alektorovi 
‘Oc. (Brh.) greenii 
‘Oc. (Cha.) wattensis 
Oc. (Och.) communis 
Oc. (Och.) infirmatus 
Oc. (Och.) scapularis 
‘Oc. (Psk.) bancroftianus 
‘Oc. (Psk.) postspiraculosus 
Oc. (Rus.) refiki 
Oc. (Rus.) rusticus 
Ps. (Gra.) jamaicensis 
Ps. (Jan.) ferox 
Ps. (Pso.) ciliata 
Rh. longirostris 
Rh. wardi 01 
Sc. albolineata 
Sc. arboricola 
St. aegypti 
St. africana 
Za. fulgens 12 
Za. longipalpis 01 
Do. leonis 
Do. nivea 
Fl. kochi 
Fl. poicilia 
Ha. australis 12 
Hg. (Con.) leucocelaenus 
Hg. (Con.) leucotaeniatus 
Hg. (Hag.) equinus 
Hg. (Hag.) splendens 
‘Oc. (Fin.) albilabris 
‘Oc. (Fin.) alboannulatus 02 
‘Oc. (Fin.) albocinctus 
‘Oc. (Fin.) albotaeniatus 
‘Oc. (Fin.) argenteitarsis 
‘Oc. (Fin.) argyronotum 23 
‘Oc. (Fin.) assamensis 
‘Oc. (Fin.) banksi 
‘Oc. (Fin.) biocellatus 
‘Oc. (Fin.) candidoscutellum 23 
‘Oc. (Fin.) chrysolineatus 
‘Oc. (Fin.) chungi 
‘Oc. (Fin.) crossi 
‘Oc. (Fin.) echinus 
‘Oc. (Fin.) elsiae 23 
‘Oc. (Fin.) embuensis 
‘Oc. (Fin.) geniculatus 
‘Oc. (Fin.) gilli 
‘Oc. (Fin.) gubernatoris 
‘Oc. (Fin.) harperi 
‘Oc. (Fin.) hatorii 
‘Oc. (Fin.) ingrami 
‘Oc. (Fin.) j. japonicus 
‘Oc. (Fin.) jugaraensis 
‘Oc. (Fin.) keefei 
‘Oc. (Fin.) loi 
‘Oc. (Fin.) lophoventralis 
‘Oc. (Fin.) luteostriatus 
‘Oc. (Fin.) macfarlanei 23 
‘Oc. (Fin.) notoscriptus 
‘Oc. (Fin.) oreophilus 
‘Oc. (Fin.) papuensis 23 
‘Oc. (Fin.) prominens 
‘Oc. (Fin.) pseudotaeniatus 23 
‘Oc. (Fin.) pulchrithorax 
‘Oc. (Fin.) pulchriventer 23 
‘Oc. (Fin.) rubrithorax 23 
‘Oc. (Fin.) seoulensis 
‘Oc. (Fin.) sherki 
‘Oc. (Fin.) subalbitarsis 23 
‘Oc. (Fin.) tubbutiensis 
‘Oc. (Mac.) tremulus 
‘Oc. (Mol.) pecuniosus 
‘Oc. (Och.) atropalpus 
‘Oc. (Och.) deserticola 
‘Oc. (Och.) epactius 
‘Oc. (Och.) fluviatilis 23 
‘Oc. (Och.) muelleri 01 
‘Oc. (Och.) sierrensis 
‘Oc. (Och.) varipalpus 
‘Oc. (Pro.) berlini 
‘Oc. (Pro.) brelandi 
‘Oc. (Pro.) burgeri 12 
‘Oc. (Pro.) galindoi 
‘Oc. (Pro.) hendersoni 
‘Oc. (Pro.) homoeopus 
‘Oc. (Pro.) knabi 
‘Oc. (Pro.) kompi 
‘Oc. (Pro.) terrens 
‘Oc. (Pro.) triseriatus 
‘Oc. (Pro.) zavortinki 
‘Oc. (Pro.) zoosophus 
Op. (Not.) chathamicus 
Op. (Opi.) fuscus 
Ta. savoryi 
Ta. togoi 
22222222222222222222222222222222
00000000011111111112222222222333
12345678901234567890123456789012
Cs.  (Cus.) inornata 
Cx. (Cux.) quinquefasciatus – – – 03 
Ma. (Man.) titillans 01 
Or. signifera 01 
Ab. papago 
Ae. cinereus – 
Ae. esoensis – 
Az. ramirezi 
Ge. baisasi 
Ge. longiforceps 01 
Gy. mediovittata 
Hw. sexlineata 
Hw. walkeri 
Ia. cavaticus 
Ke. dissimilis 01 
Ke. harbachi 01 
Ko. purpureipes 
Le. geoskusea 
Mu. alternans 01 
Mu. laniger 
‘Oc. (Brh.) alektorovi 
‘Oc. (Brh.) greenii 
‘Oc. (Cha.) wattensis 
Oc. (Och.) communis 
Oc. (Och.) infirmatus 
Oc. (Och.) scapularis 
‘Oc. (Psk.) bancroftianus 
‘Oc. (Psk.) postspiraculosus 
Oc. (Rus.) refiki 
Oc. (Rus.) rusticus 
Ps. (Gra.) jamaicensis 
Ps. (Jan.) ferox 
Ps. (Pso.) ciliata 
Rh. longirostris 
Rh. wardi 01 
Sc. albolineata 
Sc. arboricola 
St. aegypti 
St. africana 
Za. fulgens 12 
Za. longipalpis 01 
Do. leonis 
Do. nivea 
Fl. kochi 
Fl. poicilia 
Ha. australis 12 
Hg. (Con.) leucocelaenus 
Hg. (Con.) leucotaeniatus 
Hg. (Hag.) equinus 
Hg. (Hag.) splendens 
‘Oc. (Fin.) albilabris 
‘Oc. (Fin.) alboannulatus 02 
‘Oc. (Fin.) albocinctus 
‘Oc. (Fin.) albotaeniatus 
‘Oc. (Fin.) argenteitarsis 
‘Oc. (Fin.) argyronotum 23 
‘Oc. (Fin.) assamensis 
‘Oc. (Fin.) banksi 
‘Oc. (Fin.) biocellatus 
‘Oc. (Fin.) candidoscutellum 23 
‘Oc. (Fin.) chrysolineatus 
‘Oc. (Fin.) chungi 
‘Oc. (Fin.) crossi 
‘Oc. (Fin.) echinus 
‘Oc. (Fin.) elsiae 23 
‘Oc. (Fin.) embuensis 
‘Oc. (Fin.) geniculatus 
‘Oc. (Fin.) gilli 
‘Oc. (Fin.) gubernatoris 
‘Oc. (Fin.) harperi 
‘Oc. (Fin.) hatorii 
‘Oc. (Fin.) ingrami 
‘Oc. (Fin.) j. japonicus 
‘Oc. (Fin.) jugaraensis 
‘Oc. (Fin.) keefei 
‘Oc. (Fin.) loi 
‘Oc. (Fin.) lophoventralis 
‘Oc. (Fin.) luteostriatus 
‘Oc. (Fin.) macfarlanei 23 
‘Oc. (Fin.) notoscriptus 
‘Oc. (Fin.) oreophilus 
‘Oc. (Fin.) papuensis 23 
‘Oc. (Fin.) prominens 
‘Oc. (Fin.) pseudotaeniatus 23 
‘Oc. (Fin.) pulchrithorax 
‘Oc. (Fin.) pulchriventer 23 
‘Oc. (Fin.) rubrithorax 23 
‘Oc. (Fin.) seoulensis 
‘Oc. (Fin.) sherki 
‘Oc. (Fin.) subalbitarsis 23 
‘Oc. (Fin.) tubbutiensis 
‘Oc. (Mac.) tremulus 
‘Oc. (Mol.) pecuniosus 
‘Oc. (Och.) atropalpus 
‘Oc. (Och.) deserticola 
‘Oc. (Och.) epactius 
‘Oc. (Och.) fluviatilis 23 
‘Oc. (Och.) muelleri 01 
‘Oc. (Och.) sierrensis 
‘Oc. (Och.) varipalpus 
‘Oc. (Pro.) berlini 
‘Oc. (Pro.) brelandi 
‘Oc. (Pro.) burgeri 12 
‘Oc. (Pro.) galindoi 
‘Oc. (Pro.) hendersoni 
‘Oc. (Pro.) homoeopus 
‘Oc. (Pro.) knabi 
‘Oc. (Pro.) kompi 
‘Oc. (Pro.) terrens 
‘Oc. (Pro.) triseriatus 
‘Oc. (Pro.) zavortinki 
‘Oc. (Pro.) zoosophus 
Op. (Not.) chathamicus 
Op. (Opi.) fuscus 
Ta. savoryi 
Ta. togoi 

APPENDIX 3

New generic-level taxa

The new generic-level taxa recognized in the present study are described below. Additional character data for those species of each genus included in the analysis are contained in the data matrix (Appendix 2). The genera are listed in alphabetical order. Endings of some masculine adjectival species names have been changed to agree in gender with new feminine generic names.

Collessiusreinert, harbach& kitching, gen. nov.
Type species: 

Ochlerotatus macfarlanei Edwards, 1914.

Females
Head:

Vertex with decumbent scales narrow, curved, pale and dark; erect forked scales numerous on occiput and vertex; ocular line narrow, with narrow, curved, pale scales; normally 4 or 5 interocular setae (occasionally 6 in Co. elsiae elsiae); eyes contiguous or separated by less than width of 1 eye facet; antennal pedicel with small, broad, pale scales on mesal surface; clypeus bare; maxillary palpus dark-scaled with apex pale-scaled; proboscis longer than forefemur, dark-scaled with patch of pale scales on ventral and lateral surfaces near midlength (Assemblage A) or entirely dark-scaled (Assemblage B).

Thorax:

Scutum covered with narrow, curved scales except for bare median prescutellar area, scales sparse on scutal fossa, scales dark except for narrow, pale-scaled stripes on acrostichal (anterior and posterior) area, dorsocentral (anterior and posterior) area, outer and posterior margins of scutal fossa, outer margins of prescutellar area mesad of setae, and longitudinal stripe on supraalar area; setae well developed on anterior promontory, acrostichal (anterior and posterior) area, antedorsocentral area, dorsocentral (anterior and posterior) area, scutal fossa, antealar and supraalar areas, and lateral margins of prescutellar area; scutellum with narrow or broad scales on midlobe, narrow scales on lateral lobes; antepronota widely separated, with broad, pale scales, several setae; postpronotum with scales, few posterior setae; prespiracular setae absent; hypostigmal area bare; postspiracular area with scales, several setae; subspiracular area with pale scales (Co. banksi without or with few scales, Co. pseudotaeniatus without scales); upper proepisternum with pale scales, several setae, lower area bare; paratergite narrow, with pale scales; mesokatepisternum with broad, pale scales in upper and lower patches, several upper and posterior setae; prealar area with broad, pale scales on upper (absent in Co. banksi) and lower areas, numerous upper setae; mesepimeron with 1 patch of broad, pale scales on upper area and extending onto middle, several upper setae, lower setae absent.

Wing:

Dark-scaled with elongate, narrow, pale-scaled patch at or near base of anterior margin of costa; upper calypter with several setae on margin; alula with row of narrow, dark scales on margin; remigium with 2 or 3 setae distally on dorsal surface.

Legs:

Midfemur with narrow, longitudinal stripe of white-scales from base to or near apex on middle of anterior surface; hindfemur with subapical white scales but apex fringed with dark scales; hindtarsomeres 1 and 2 with narrow, white-scaled, basal and apical bands; fore- and midtarsi with 2 equal ungues, each with 1 tooth; hindtarsi with 2 equal ungues, simple.

Abdomen:

Tergum I with pale-scaled patch on lateral tergite; terga II–V with narrow, pale-scaled, basal bands.

Genitalia:

Tergum VIII with apex broadly rounded, numerous broad scales on approximately distal 0.30–0.40; sternum VIII sclerotized, width greater than length, apical margin with moderate, median emargination separating broadly rounded lobes (Co. hatorii and Assemblage B with apex nearly straight), numerous short, lanceolate scales and occasionally 1–3 broad scales (Co. hatorii and Assemblage B without lanceolate scales, but with several to numerous broad scales); seta 2-S inserted posterior to 1-S; tergum IX comprising single, moderately to heavily pigmented, elongate plate with median emargination apically in Assemblage A or with plate short and wide in Assemblage B, several setae on each side of midline distally; postgenital lobe relatively long, narrow, apex normally flat or with very shallow, median emargination (Co. hatorii and Assemblage B with postgenital lobe somewhat shorter and wider), several setae distally; upper vaginal sclerite small to moderate size; lower vaginal sclerite absent; insula liplike, with several setae in lateral patches; cercus moderately long, moderately wide throughout length, apex broadly rounded, scales absent (Assemblage B with cercus somewhat shorter and with scales); 3 spermathecal capsules.

Males
Head:

Maxillary palpus with 5 palpomeres, slightly shorter than proboscis, palpomeres 4 and 5 slightly down-turned, palpomeres 3 (distal part) and 4 with moderate number of long setae ventrally; antennal flagellar whorls with numerous long setae directed dorsally and ventrally.

Legs:

Fore- and midtarsi with 2 unequal ungues, larger unguis with 2 teeth, smaller unguis with 1 tooth; hindtarsus with 2 equal ungues, simple.

Genitalia:

Tergum IX with poorly sclerotized, moderate lobe bearing several moderately long, slender setae on each side of midline of posterior margin; gonocoxite moderately long, relatively narrow throughout length, dorsal surface without apical and basal lobes on mesal margin, with few moderately long and long, relatively slender setae scattered over dorsal and ventral surfaces, mesal surface membranous, lateral, outer part of dorsal and most of ventral surfaces with broad scales; gonostylus attached at apex of gonocoxite, long, narrow, slightly curved, gonostylar claw short, narrow, attached at apex, 1 or more shorter, stout setae subapically (these similar to gonostylar claw but somewhat shorter and thinner, especially in Co. hatorii) (Assemblage A) or subapical, stout setae absent (Assemblage B); aedeagus simple, tubelike, distal part wider than middle (Assemblage A) or approximately same width as middle (Assemblage B); proctiger with few tiny, cercal setae; claspette with single, moderately long, columnar stem with small subapical, thumblike lobe (Assemblage A) or subapical lobe absent (Assemblage B), claspette filament attached apically, long, relatively narrow, strongly curved, apex pointed; sternum IX with several setae on posteromesal area.

Pupae
Cephalothorax:

Setae 1,3-CT similarly developed; 11-CT normally single, rarely 2-branched.

Abdomen:

Seta 3-I,II branched (Assemblage A) or normally single (Assemblage B), longer than 6-I,II; 6-I shorter than 7-I; 1-II with several branches; 5-II inserted lateral to 4-II; 3-III single (Co. banksi single or 2-branched); 5-V long, stout, single, longer than tergum VI median length; 2-VI inserted mesal to 1,3-VI (Assemblage A) or 2-VI inserted lateral to 1-VI and mesal to 3-VI (Assemblage B); 6-VII inserted posterior and mesal to 9-VII; 9-VIII inserted slightly anterior to posterolateral corner (Assemblage A) or on corner (Assemblage B), with multiple branches.

Paddle:

Apical margin rounded; midrib extends to apex of paddle; without hairlike spicules on margins; seta 1-Pa short, single.

Fourth-instar larvae
Head:

Seta 1-C slender, single, apex attenuate; 4-C short, slender, with few branches, inserted at approximately same level or slightly anterior to 5,6-C; 5,6-C relatively short, moderately stout, with several branches, 6-C lateral to 5-C, 4–6-C all at approximately same level; 7-C moderately long, relatively slender, with several branches, inserted lateral and posterior to 4–6-C; 12-C short, slender, with several branches, inserted mesal to 13-C; 13-C single, longer than 12-C; 14-C short, stout, normally single or 2-branched, occasionally 3-branched; 19-C absent; antenna moderately long, narrow, with several to numerous spicules, seta 1-A moderately long, normally with 2 or 3 branches (rarely single).

Thorax:

Setae 1–3-P on common setal support plate; 4-P single; 8-P branched, shorter than 4-P; 1-M,T stout, single or 2-branched, inserted on large, pigmented tubercle (Assemblage A) or 1-M,T slender, with few branches, not on tubercle (Assemblage B); 5-M longer than 7-M; 2,6-T single (Lu & Ji, 1997: fig. 44, of Co. ningheensis shows 2-T with 3 branches).

Abdomen:

Seta 6-I,II long, with 2 branches, 6-II longer than 6-III; 7-I,II long, single (rarely 2-branched); 12-I present; 8-II short, single or 2-branched; 1-VII long; 12-VII single; 2,4-VIII single; comb comprising numerous scales in patch; segment X with moderate saddle, incomplete ventrally, with numerous stout spicules on posterior margin dorsal to seta 1-X, acus absent, 1-X inserted on saddle, ventral brush with several long, branched setae with short stems, attached to grid with both transverse and lateral bars, some species with 1 shorter seta anterior to grid.

Siphon:

With attached acus; pecten with numerous closely set spines; seta 1-S single or branched, inserted within distal part of pecten or slightly distal to pecten.

Included species
Assemblage A:

Collessius elsiae elsiae (Barraud), Co. elsiae vicarius (Lien), Co. hatorii (Yamada), Co. macdougalli (Edwards), Co. macfarlanei (Edwards), Co. ningheensis (Lei) and Co. shortti (Barraud). Assemblage B: Collessius banksi (Edwards), Co. pseudotaeniatus (Giles) and Co. tonkinensis (Galliard & Ngu).

Distribution
Assemblage A:

China, India, Indonesia, Japan, Malaysia, Pakistan, Sri Lanka, South Korea, Taiwan, Thailand and Vietnam. Assemblage B: Bangladesh, India, Myanmar, Nepal, Philippine Islands, Sri Lanka and Vietnam.

Bionomics
Assemblage A:

Immatures are normally found in rock pools but also in artificial containers (e.g. concrete water tanks). Females of some species bite humans indoors and outdoors. Assemblage B: Immatures have been collected normally from rock pools but also from tree holes, and artificial containers (e.g. cement sinks and gutters, drains, iron cisterns, tires, etc.). Females of Co. pseudotaeniatus bite humans indoors and outdoors.

Discussion

Species of Collessius can be grouped into Assemblages based on morphological characters (see above). Barraud (1934: figs 37, 39) illustrated the development of the gonostylus and fourth-instar larva of Co. elsiae and Co. shortti. See Tanaka et al. (1979: figs 97, 98, 219) for illustrations of the male genitalia, fourth-instar larva and adults of Co. hatorii, Tanaka (2002: fig. 1) for the pupa and Moriya et al. (1973) for the egg of this species.

Etymology

Collessius is named in honour of Dr Donald H. Colless in recognition of his contributions to the taxonomy of Oriental mosquitoes. The generic name is masculine, formed from his surname and the masculine Latin suffix ‘-ius’. Recommended abbreviation = Co.

Dahlianareinert, harbach& kitching, gen. nov.
Type species:

Culex geniculatus Olivier, 1791.

Females
Head:

Median area of vertex with narrow, curved, pale scales; occiput and vertex with numerous erect forked scales; ocular line narrow, with narrow, white scales; eyes above antennal pedicels separated by width of 2–4 ocular facets; antennal pedicel with few short, fine setae, few small, broad scales sometimes present on mesal surface; clypeus bare; maxillary palpus and proboscis dark-scaled, proboscis longer than forefemur.

Thorax:

Scutum with narrow, curved scales; pale scales forming narrow, longitudinal stripe on anterior and posterior acrostichal area and extending posteriorly to or near prescutellar area, posterior dorsocentral area extending posteriorly laterad of prescutellar setae to posterior margin of scutum, and on anterior and lateral margins of prescutellar bare area; white scales on antealar area and extending posteriorly as narrow, longitudinal stripe on supraalar area; scutal fossa covered with pale scales; acrostichal (anterior and posterior) and dorsocentral (anterior and posterior) areas with setae; scutellum with narrow, curved scales on lateral lobes; paratergite with broad, white scales; antepronota widely separated, with broad and few moderately broad, pale scales; postpronotum with broad, pale and dark scales, Da. echinus normally also with few narrow, curved scales dorsally; broad, white scales on postspiracular area, subspiracular area, upper proepisternum, mesokatepisternum (upper and lower posterior patches), upper and lower prealar areas, mesepimeron (1 large patch); mesepimeron without lower setae.

Wing:

Dark-scaled; upper calypter with several setae on margin; alula with row of narrow scales on margin; remigium with 2 or 3 setae on dorsal surface distally.

Legs:

Ante- and postprocoxal membranes bare; hindfemur with well-developed apical, white-scaled band; hindtibia and tarsus dark-scaled, ungues equal, simple; fore- and midungues equal, each with 1 tooth.

Abdomen:

Tergum I with pale-scaled patch on laterotergite; segment VII laterally compressed.

Genitalia:

Tergum VIII heavily pigmented, basolateral corners expanded into short, proximally curved, thumblike projections, apex broadly rounded, numerous broad scales on distal 0.56–0.58; sternum VIII heavily pigmented, base straight or very slightly concave mesally, apex noticeably wider than base, nearly straight but with shallow, median emargination, broad scales absent or very few (Da. geniculata) or several scales forming lateral patches (Da. echinus), 2 moderately long setae slightly subapical on lateral part of apical margin, width greater than length; tergum IX heavily pigmented with median area slightly lighter, apex with moderately deep, V-shaped, median emargination, about 6 moderately long, stout setae apically on each side of midline; postgenital lobe moderately long, relatively narrow, apex rounded, setae on distal part; upper vaginal sclerite moderately large to large, heavily pigmented; lower vaginal sclerite absent; insula liplike, with few setae in lateral patches; cercus relatively long, moderately wide, apex broadly rounded, without scales; 3 spermathecal capsules.

Males
Head:

Maxillary palpus with 5 palpomeres, slightly shorter than proboscis, dark-scaled, numerous long and moderately long setae ventrolaterally on distal part of palpomere 3 and ventrolaterally on palpomeres 4 and 5; antennal flagellomeres with numerous long setae directed dorsally and ventrally.

Legs:

Foretarsus with 2 unequal ungues, larger unguis with 2 teeth, smaller with 1 tooth; midtarsus with 2 unequal ungues, each with 1 tooth; hindtarsus with 2 equal ungues, simple.

Genitalia:

Tergum IX comprising 2 relatively broad, lateral plates each with short, narrow, strongly sclerotized lobe on posteromesal area bearing 2–6 short, stout, flattened, dark setae; gonocoxite moderately long, relatively narrow, dorsal surface without apical and basal lobes on mesal margin, ventral surface without specialized setae or scales, mesal surface membranous, numerous scales on dorsal, lateral and ventral surfaces; gonostylus attached at apex of gonocoxite, relatively long, narrow, distal part gently curved mesally, single gonostylar claw, long, narrow, attached at apex; aedeagus simple, tubelike, lateral margins gently curved to apex, distal 0.50 narrowest; proctiger with few to several cercal setae; claspette with single moderately long to long, narrow, columnar stem, claspette filament attached apically, long, somewhat flattened, slightly curved to narrow, pointed apex; sternum IX with several setae on posteromedian area.

Pupae (based on Da. geniculata)
Cephalothorax:

Setae 1,3-CT similarly developed, normally with 2 branches; 11-CT single.

Abdomen:

Seta 3-I single, shorter than 6-I; 6-I longer than 7-I; 2-II inserted lateral to 1,3-II; 3-II moderately long, stout, normally single (occasionally 2-forked), shorter than 6-II; 5-II long, stout, single, noticeably longer than 3-II, slightly shorter than 6-II, inserted lateral to 4-II; 3,6-III long, stout, single; 5-V long, stout, single, longer than tergum VI median length; 2-VI inserted anterior and slightly lateral to 1-VI and mesal to 3-VI; 6-VII short, slender, inserted posterior and slightly mesal to 9-VII; 9-VII moderately long, stout, aciculate, usually with 2 branches; 9-VIII moderately long, stout, aciculate, with 2–5 (usually 2 or 3) branches.

Paddle:

Length noticeably greater than width; midrib well developed, reaching apex of paddle; without hairlike spicules on margins; seta 1-Pa short, slender, single.

Fourth-instar larvae
Head:

Seta 1-C single, moderately stout; 4-C moderately long, slender, multiple-branched, inserted mesal and slightly anterior to 6-C; 5-C long, stout, single, inserted posterior to 4,6,7-C; 6-C moderately long to long, stout, single to 3-branched, inserted anterior and approximately same lateral level to 5-C and mesal and slightly anterior to 7-C; 7-C moderately long, moderately stout, with 2 or 3 branches; 12-C branched, inserted mesal to 13-C; 13-C single, longer than 12-C; 14-C short, branched; 19-C absent; antenna moderately long, narrow, without spicules, seta 1-A moderately long, single or 2-branched.

Thorax:

Setae 1–3-P inserted on common setal support plate; 5-P with 2 branches, longer than 6-P; 8-P short, multiple-branched; 4-M single; 5-M longer than 7-M; 2,6-T single.

Abdomen:

Seta 6-I,II with 2 long, stout, aciculate branches; 7-I,II long, stout, single; 12-I present; 8-II single; 1-VII moderately long, stout, stellate, multiple-branched; 12-VII single; 2,4-VIII single; segment VIII with comb comprising several scales in single curved row; segment X with large saddle, incomplete ventrally, acus absent, seta 1-X long, stout, branched, inserted on saddle, ventral brush with several long, normally 2–4-branched setae with long stems, inserted on grid with both transverse and lateral bars, 2 precratal setae.

Siphon:

With attached acus; pecten with several, closely set spines; seta 1-S moderately long, multiple-branched, inserted distal to pecten.

Included species

Dahliana echinus (Edwards), Da. geniculata (Olivier) and Da. gilcolladoi (Sanchez-Covisa Villa, Rodriguez Rodriguez & Guillen Llera).

Distribution

Albania, Algeria, Austria, Belarus, Belgium, Bulgaria, Croatia, Czech Republic, Denmark, France, European Russia, Finland, Germany, Greece, Hungary, Iran, Italy, Lithuania, Macedonia, Morocco, Netherlands, Norway, Poland, Portugal, Romania, Slovenia, Slovakia, Spain, Sweden, Switzerland, Turkey, Ukraine, United Kingdom and Yugoslavia.

Bionomics

Immature stages normally inhabit treeholes but have been found in open tree stumps, root holes of olive trees and pools in rocks.

Discussion

See discussion under Classification of Evaluated Taxa. Marshall (1938: figs 12, 57(g), 69, 94 and pl. IV) provided illustrations of the egg, first- and fourth-instar larvae, male genitalia and female of Da. geniculata, and Reinert (2002e: fig. 3) illustrated the female genitalia of this species. Villa, Rodriguez & Llera (1985) provide photographs of the fourth-instar larvae of Da. echinus, Da. geniculata and Da. gilcolladoi.

Etymology

Dahliana is named in honour of Dr Christine Dahl in recognition of her many important contributions to the taxonomy and bionomics of Palaearctic mosquitoes. The generic name is feminine, formed from her surname and the feminine Latin suffix ‘-iana’. Recommended abbreviation = Da.

Dobrotworskyius Reinert, Harbach& Kitching, gen. nov.
Type species:

Aedes tubbutiensisDobrotworsky (1959).

Females
Head:

Vertex with decumbent scales narrow, curved, pale and dark; erect forked scales numerous on occiput and vertex; ocular line narrow, covered with narrow, curved, pale scales; 2–4 long, interocular setae; eyes above antennal pedicels contiguous (but separated by approximately width of 3.5 eye facets in Db. rubrithorax); antennal pedicels with several small, broad, pale scales on mesal surface; clypeus bare; maxillary palpus dark-scaled with pale scales at apex of palpomeres 3 and 4; proboscis longer than forefemur, dark-scaled (few pale scales intermixed in Db. alboannulatus and Db. tubbutiensis).

Thorax:

Covered with narrow, curved scales (Db. alboannulatus and Db. milsoni also with few broad, pale scales on lateral margins of bare prescutellar area), both dark and pale scales present, pale scales on lateral margin of bare prescutellar area mesal to setae; setae on following: few on anterior promontory, antedorsocentral area, scutal fossa and antealar area, numerous on acrostichal (anterior and posterior), dorsocentral (anterior and posterior), supraalar and prescutellar areas; scutellum with narrow, curved scales on all lobes; antepronota widely separated, with pale scales, normally narrow, curved; postpronotum with both narrow, curved and broad scales, few posterior setae; prespiracular setae absent; hypostigmal area bare; postspiracular area with scales, several setae; subspiracular area with broad, pale scales; upper proepisternum with broad, pale scales, several setae on upper area, lower area bare; paratergite with pale scales; mesokatepisternum with broad, pale scales in upper and lower patches, 3 or 4 upper and several posterior setae; prealar area with broad, pale scales in patch on lower area and few on knob, several upper setae; mesepimeron with 2 patches of broad, pale scales, 1 on upper and 1 on middle, scale patches nearly meet in some specimens, several upper setae, lower setae absent; metameron bare.

Wing:

Dark-scaled, some specimens with 1–3 pale scales at base of costa; upper calypter with several setae on margin; alula with row of narrow, dark scales on margin; remigium with 1–3 setae distally on dorsal surface.

Legs:

Hindfemur with pale-scaled apex; hindtarsus with broad, pale-scaled, basal bands on tarsomeres 1–4; fore-, mid- and hindtarsi with 2 equal ungues, each with 1 tooth.

Abdomen:

Tergum I with patch of pale scales on lateral tergite; terga dark-scaled with dorsomesal pale-scaled areas basally and basolaterally, not forming complete bands.

Genitalia:

Tergum VIII with apex broadly rounded, numerous broad scales on distal half; sternum VIII sclerotized, length equal to or slightly longer than width, apical margin with median emargination separating 2 moderate lobes, several to moderate number of broad scales on lateral areas, seta 2-S inserted posterior to 1-S; tergum IX comprising 2 pigmented, lateral plates normally connected mesally by lightly pigmented, narrow, basal strip, inner margin of plates darkly pigmented distally, setae absent (Db. milsoni occasionally with 1 seta distally on 1 or both plates); postgenital lobe moderately long, moderately wide but distal part narrower, apex normally flat or with minute emargination, several setae distally; upper vaginal sclerite small; lower vaginal sclerite absent; insula liplike, transversely U-shaped (depressed), several setae in lateral patches; cercus moderately long, moderately wide distally but proximal part wider, apex broadly rounded, without scales; 3 spermathecal capsules.

Males
Head:

Maxillary palpus with 5 palpomeres, slightly shorter than proboscis, palpomeres 4 and 5 slightly down-turned, numerous long setae on distal part of palpomere 2 ventrally and palpomere 4 ventrolaterally; antenna flagellar whorls with numerous long setae directed dorsally and ventrally.

Legs:

Fore- and midtarsi with 2 unequal ungues, larger unguis with 2 teeth, smaller unguis with 1 tooth, some species with larger midunguis with 1 tooth; hindtarsus with 2 equal unguis, each with 1 tooth.

Genitalia:

Tergum IX with narrow, median area separating broad, lateral areas, posterior margin with sclerotized, small, short, narrow lobe bearing few short, slender setae on each side of midline; gonocoxite moderately long, moderately wide, mesal margin of dorsal surface with small basal lobe bearing few moderately long and short, relatively slender setae, without apical lobe, mesal surface membranous, lateral, outer part of dorsal and most of ventral surfaces with broad scales; gonostylus attached at apex of gonocoxite, long, narrow, slightly curved, single gonostylar claw, long, narrow, attached at apex; aedeagus simple, tubelike; proctiger with few tiny, cercal setae; claspette with single, moderately long, narrow, columnar stem, claspette filament attached apically, long, somewhat flattened beyond middle, distal part tapering to narrow, curved point; sternum IX with several setae on posteromedian area.

Pupae
Cephalothorax:

Seta 1,3-CT similarly developed; 11-CT single or 2-branched.

Abdomen:

Seta 3-I,II branched, 3-I longer than 6-I, 3-II longer than 6-II; 6-I longer than 7-I; 1-II with multiple branches; 5-II inserted lateral to 4-II; 3-III single, longer than 5-III; 5-V long, stout, single, longer than tergum VI median length; 2-VI inserted mesal to 1,3-VI; 6-VII inserted posterior and slightly mesal to 9-VII, both with multiple branches; 9-VIII inserted on posterolateral corner, with multiple branches.

Paddle:

Apical margin rounded; midrib extending to or near apex of paddle; without hairlike spicules on margins; seta 1-Pa short, slender, single.

Fourth-instar larvae
Head:

Seta 1-C dark, spiniform; 4-C short, slender, with few branches, inserted mesal and approximately in line or slightly posterior or anterior to 5-C and noticeably posterior to 6-C; 5-C moderately long, relatively slender, with several branches, inserted mesal and posterior to 6-C; 6-C moderately long, moderately stout, with several branches, inserted at approximately same level or slightly anterior to 7-C; 7-C moderately long, relatively slender, aciculate, with several branches, inserted lateral and anterior to 5-C; 12-C short, slender, with several branches, inserted mesal to 13-C; 13-C single, longer than 12-C; 14-C short, single; 19-C absent; antenna moderately long, with numerous spicules, seta 1-A moderately long, aciculate, with several branches.

Thorax:

Setae 1–3-P on common setal support plate (in few available species examined); 4,8-P branched; 6-T normally single.

Abdomen:

Seta 6-I,II long, with 2 branches, 6-II longer than 6-III; 7-I long, single; 12-I present; 7-II short, slender, with several branches; 8-II short, branched; 1-VII moderately long, moderately stout; 12-VII normally single; 2,4-VIII single (2-branched in single available specimen of Db. rubrithorax); segment VIII with comb comprising numerous scales in patch; segment X with moderate saddle, incomplete ventrally, acus absent, seta 1-X moderately long, inserted on saddle, ventral brush with several long, multiple-branched setae with short stems, inserted on grid with well-developed transverse and lateral bars, anterior 2 setae shorter and occasionally 1 of these inserted anterior to grid.

Siphon:

With attached acus; pecten with numerous, closely set spines, seta 1-S moderately long, moderately stout, aciculate, with several branches, inserted slightly distal to pecten or rarely level with distal 1 or 2 pecten spines.

Included species

Dobrotworskyius alboannulatus (Macquart), Db. milsoni (Taylor), Db. occidentalis (Skuse), Db. rubrithorax (Macquart), Db. rupestris (Dobrotworsky), Db. subbasalis (Dobrotworsky) and Db. tubbutiensis (Dobrotworsky).

Distribution

Australia.

Bionomics

Immatures are found in rock pools in river and creek beds and occasionally in ground pools with organic matter on the bottom. In addition to the usual habitats some species also were collected from old wells, water collections around houses, artificial containers, rot-hole in a tree and cavities in logs. Females of most species have been collected biting humans during the day, but also in the evening and at night.

Discussion

See Dobrotworsky (1959, 1962, 1965) for illustrations and descriptions of most life stages of species included in Dobrotworskyius. Russell (1996) provided photographs and illustrations of females of Db. alboannulatus, Db. rubrithorax and Db. rupestris. Linley et al. (1991a) provided descriptions and photographs of the eggs of Db. alboannulatus and Db. rubrithorax.

Etymology

Dobrotworskyius is named in honour of Dr Nikolai V. Dobrotworsky in recognition of his important contributions to the taxonomy of Australian mosquitoes, especially of Victoria. The generic name is masculine, formed from his surname and the masculine Latin suffix ‘-ius’. Recommended abbreviation = Db.

Georgecraigius Reinert, Harbach& Kitching, gen. nov.
Type species:

Culex atropalpus Coquillett, 1902.

Females
Head:

Vertex with decumbent scales narrow, curved, pale mesally, broad laterally, few erect forked scales; occiput with numerous erect forked scales; ocular line narrow with narrow, curved, pale scales; eyes moderately to broadly separated above antennal pedicels; antennal pedicel with few, small, broad scales and few short, slender setae mesally; clypeus bare; maxillary palpus and proboscis dark-scaled.

Thorax:

Acrostichal area without anterior and posterior setae; anterior dorsocentral area without setae, posterior dorsocentral setae present; supraalar area with longitudinal patch of pale scales; prescutellar area with pale scales on lateral margins of median, bare space; scutellum with narrow, curved scales on all lobes; scales present on antepronotum, postpronotum, subspiracular area (broad), upper proepisternum (broad), mesokatepisternum (broad in upper and lower posterior patches), lower prealar area (broad) and mesepimeron (broad); mesepimeron without lower setae; metameron bare.

Wing:

Dark-scaled with white-scaled patch at base of costa; upper calypter with several setae on margin; alula with row of narrow scales on margin; remigium without setae.

Legs:

Hindfemur with band of pale scales at apex; hindtarsomeres 1 and 2 with narrow, pale-scaled, basal band, hindungues simple; fore- and midungues each with 1 tooth.

Genitalia:

Tergum VIII with apex broadly rounded, numerous scales on much of surface; sternum VIII sclerotized, width slightly greater than length, apical margin broadly rounded, numerous scales on much of surface, seta 2-S inserted posterior to 1-S; tergum IX comprising 2 lightly to moderately pigmented, short, oval sclerites attached mesally by membrane, without setae; postgenital lobe relatively broad, apex flat or with very shallow, median emargination, numerous setae distally; upper vaginal sclerite small, moderately pigmented; lower vaginal sclerite absent; insula liplike, with few setae in lateral patches; cercus relatively short, moderately wide, apex broadly rounded, without scales; 3 spermathecal capsules.

Males
Head:

Maxillary palpus with 5 palpomeres, shorter than proboscis; antennal flagellar whorls with numerous long setae directed dorsally and ventrally.

Legs:

Fore- and midtarsi with 2 unequal ungues, larger unguis with 2 teeth, smaller with 1 tooth; hindtarsus with 2 equal ungues, simple.

Genitalia:

Tergum IX with pair of lateral lobes, relatively narrowly connected mesally, posterior margin relatively flat but each side gently sloping to midline, without setae; gonocoxite moderately long, relatively narrow, mesal margin of dorsal surface without apical lobe, with small, darkened, basal lobe bearing 2–4 long, relatively stout setae and several slender setae, ventral surface with row of long, relatively stout, more or less evenly spaced setae on mesal margin, mesal surface membranous; gonostylus attached at apex of gonocoxite, long, narrow, curved mesally, with single, relatively short, narrow, gonostylar claw at apex; aedeagus simple, tubelike, distal half slightly wider, apex convex; proctiger with cercal setae; claspette with single moderately long to long, narrow, columnar stem, claspette filament attached apically, long, somewhat flattened, slightly curved; sternum IX with 2–12 setae.

Pupae
Cephalothorax:

Seta 11-CT single, longer than 10,12-CT.

Abdomen:

Seta 3-I long, stout, single (rarely 2-branched on 1 side of Gc. atropalpus), noticeably longer than 6-I; 7-I longer than 6-I; 1-II very slender, multiple-branched; 2-II short, lateral to 1,3-II; 3-III single, long, stout, inserted at same level as 1-III; 5-III inserted mesal to 4-III; 6-VII short, slender, single or 2-branched, inserted mesal and posterior to 9-VII; 9-VII stout, branched, longer than 6-VII; 9-VIII moderately long, stout, multiple-branched.

Paddle:

Wide; without hairlike spicules on lateral margin; midrib extends to apex of paddle; seta 1-Pa single.

Fourth-instar larvae
Head:

Seta 1-C slender to moderately stout, apex attenuate, single; 4-C short, slender, branched, inserted mesal and slightly posterior to 6-C; 5-C moderately long to long, stout, inserted posterior to 6-C; 6-C moderately long, stout; 7-C moderately long, moderately stout, branched, inserted approximately at same level or slightly posterior to 5-C; 12-C short, branched, shorter than and inserted mesal to 13-C; 13-C single; 14-C short, single; 19-C absent; antenna moderately long, with few to several spicules.

Thorax:

Seta 4-P single; 5-M longer than 7-M; 2,6-T single.

Abdomen:

Seta 12-I present; 6-I–III long, aciculate, branched; 7-I long, stout, aciculate; 7,10,12-VII single; segment VIII with comb comprising numerous scales in patch, seta 2-VIII longer than 1-VIII, 2,4-VIII single; segment X with relatively small saddle, incomplete ventrally, acus absent, seta 1-X short, slender, single, inserted ventral to saddle (occasionally on ventral margin of saddle in Gc. fluviatilis), ventral brush attached to grid with both transverse and lateral bars, without precratal setae.

Siphon:

With acus attached; pecten with numerous spines, evenly spaced or with distal 1 or 2 spines more widely spaced; seta 1-S moderately long, aciculate, multiple-branched.

Included species

Georgecraigius atropalpus (Coquillett), Gc. epactius (Dyar & Knab) and Gc. fluviatilis (Lutz).

Distribution

Argentina (Misiones), Bolivia (Cochabamba), Brazil (Bahia, Goias, Minas Gerais, Para, Rio de Janeiro, Sao Paulo, Sergipe), Canada (Labrador, Newfoundland, Ontario, Quebec), Colombia (Boyaca, Cundinamarca, Meta), Costa Rica (Alajuela, Cartago, Guanacaste, San Jose), Ecuador (Zamora), El Salvador, French Guiana (Guyane, Inini), Guatemala (Guatemala), Guyana (Essequibo), Honduras (Colon), Mexico (Baja California Sur, Chiapas, Guerrero, Morelos, Nuevo Leon, Oaxaca, Puebla, Veracruz), Nicaragua (Zelaya), Panama (Canal Zone, Chiriqui, Cocle, Colon, Darien, Panama), Surinam (Suriname), United States (Alabama, Arizona, Arkansas, Colorado, Connecticut, District of Columbia, Georgia, Indiana, Kansas, Kentucky, Louisiana, Maine, Maryland, Massachusettus, Michigan, Minnesota, Missouri, Nebraska, New Hampshire, New Jersey, New Mexico, New York, North Carolina, Ohio, Oklahoma, Pennsylvania, Rhode Island, South Carolina, Tennessee, Texas, Utah, Vermont, Virginia, West Virginia, Wisconsin) and Venezuela (Aragua, Distrito Federal, Territorio Amazonas).

Bionomics

Immature stages usually are collected from rockholes and rock pools, but also are found in artificial containers, and occasionally treeholes, ground pools, stream pools and cavities in agave leaves. Wood et al. (1979) provide a summary of egg development for Gc. atropalpus.

Discussion

Georgecraigius includes two subgenera, Georgecraigius and Horsfallius (see descriptions below). Zavortink (1972) provided descriptions and illustrations of the members of this genus (as Atropalpus Section). Yamaguti & LaCasse (1951), Carpenter & LaCasse (1955) and Wood et al. (1979) also provide descriptions and illustrations of Gc. atropalpus. See Linley & Craig (1994) for descriptions and photographs of Gc. atropalpus and Gc. epactius eggs.

Etymology

Georgecraigius is named in honour of Dr George Brownlee Craig, Jr in recognition of his pioneering contributions to the genetics of Nearctic mosquitoes and descriptions of the eggs of aedine species of this region. The generic name is masculine, formed from his first name, surname and the masculine Latin suffix ‘-ius’. Recommended abbreviation = Gc.

GeorgecraigiussubgenusGeorgecraigius Reinert, Harbach& Kitching
Females
Thorax:

Antealar scales pale; paratergite without scales; postpronotum with broad scales; postspiracular area with patch of broad, white scales.

Legs:

Postprocoxal membrane without scales; hindtarsomere 2 with narrow, apical, pale-scaled band.

Abdomen:

Tergum I without scales on laterotergite.

Males
Head:

Maxillary palpus with few short setae ventrally on distal part of palpomere 3 and ventrolaterally on palpomere 4, palpomeres 3 and 4 not down-turned.

Pupae
Abdomen:

Seta 5-II inserted mesal to 4-II; 6-II shorter than 7-II; 6-III-V short; 5-IV–VI each less than or approximately equal to following tergum; 2-VI,VII inserted lateral to or at same level as 1-VI,VII; 8-VI,VII inserted ventrally.

Fourth-instar larvae
Head:

Seta 5-C long, single.

Thorax:

Setae 1–3-P and 5,6-P not inserted on common setal support plate; 4-M single.

Abdomen:

Seta 7-II long, developed similar to 7-I; 13-III–V moderately long, branched; 1-VII short, slender, single.

Siphon:

Pecten with distal 1 or 2 spines more widely spaced; seta 1-S inserted within pecten.

Included species

Georgecraigus atropalpus and Gc. epactius.

Distribution

Canada (Labrador, Newfoundland, Ontario, Quebec), Costa Rica (Alajuela, Guanacaste), El Salvador, Guatemala (Guatemala), Mexico (Baja California Sur, Chiapas, Guerrero, Morelos, Nuevo Leon, Oaxaca, Puebla, Veracruz), Panama (Panama) and United States (Alabama, Arizona, Arkansas, Colorado, Connecticut, District of Columbia, Georgia, Indiana, Kansas, Kentucky, Louisiana, Maine, Maryland, Massachusetts, Michigan, Minnesota, Missouri, Nebraska, New Hampshire, New Jersey, New Mexico, New York, North Carolina, Ohio, Oklahoma, Pennsylvania, Rhode Island, South Carolina, Tennessee, Texas, Utah, Vermont, Virginia, West Virginia, Wisconsin).

Bionomics

Immature stages usually occur in rockholes and rock pools, but also have been found in artificial containers, and occasionally in treeholes, ground pools, stream pools and cavities in agave leaves.

Recommended abbreviation = Gec.

GeorgecraigiussubgenusHorsfallius Reinert, Harbach& Kitching, subgen. nov.
Type species:

Culex fluviatilis Lutz, 1904.

Females
Thorax:

Antealar scales dark; paratergite with broad, pale scales (rarely absent?); postpronotum with narrow, curved scales dorsally; postspiracular area without scales.

Legs:

Postprocoxal membrane with pale scales; hindtarsomere 2 without apical, pale-scaled band.

Abdomen:

Tergum I with broad, pale scales on laterotergite.

Males
Head:

Maxillary palpus with several to numerous moderate to long setae ventrally on distal part of palpomere 3 and ventrolaterally on palpomere 4, palpomeres 3 and 4 down-turned.

Pupae
Abdomen:

Seta 5-II inserted lateral to 4-II; 6-II longer than 7-II; 6-III-V relatively long; 5-IV-VI very long, stout, each noticeably longer than following tergum; 2-VI,VII inserted mesal to 1-VI,VII; 8-VI,VII inserted dorsally.

Fourth-instar larvae
Head:

Seta 5-C moderately long, multiple-branched.

Thorax:

Setae 1–3-P and 5,6-P inserted on common setal support plates; 4-M branched.

Abdomen:

Seta 7-II short, developed differently than 7-I; 13-III–V long, single; 1-VII long, stout, aciculate, with 2 branches.

Siphon:

Pecten with spines evenly spaced; seta 1-S inserted slightly distal to pecten.

Included species

Georgecraigius fluviatilis.

Distribution

Argentina (Misiones), Bolivia (Cochabamba), Brazil (Bahia, Goias, Minas Gerais, Para, Rio de Janeiro, Sao Paulo, Sergipe), Colombia (Boyaca, Cundinamarca, Meta), Costa Rica (Alajuela, Cartago, San Jose), Ecuador (Zamora), French Guiana (Guyane, Inini), Guyana (Essequibo), Honduras (Colon), Mexico (Veracruz), Nicaragua (Zelaya), Panama (Canal Zone, Chiriqui, Cocle, Colon, Darien, Panama), Surinam (Suriname) and Venezuela (Aragua, Distrito Federal, Territorio Amazonas).

Bionomics

Immature stages usually are collected from rockholes, rock pools and stream pools. They are found occasionally in artificial containers, ground pools and treeholes.

Etymology

Subgenus Horsfallius is named in honour of Dr William Robert Horsfall in recognition of his outstanding contributions to the biology of New World mosquitoes and descriptions of the eggs of aedine species of the Nearctic Region. The name is masculine, formed from his surname and the masculine Latin suffix ‘-ius’. Recommended abbreviation = Hor.

Gilesius Reinert, Harbach& Kitching, gen. nov.
Type species:

Culex pulchriventer Giles, 1901.

Females
Head:

Vertex with decumbent scales narrow and curved, several broad scales present or absent on anterolateral area; erect forked scales numerous on occiput and vertex; ocular line narrow, with narrow, curved, pale scales; 4 or 5 interocular setae; eyes separated by approximately width of 3 ocular facets; antennal pedicel with small, broad, pale scales on mesal surface; clypeus bare; maxillary palpus dark-scaled, with or without pale scales on subbasal area; proboscis longer than forefemur, dark-scaled.

Thorax:

Scutum covered with narrow, curved scales except bare median prescutellar area; scales pale on following: combined anterior acrostichal and anterior dorsocentral areas, moderate stripe on posterior acrostichal area, narrow stripe on posterior dorsocentral area, narrow stripe on outer margins mesad of setae on prescutellar area, and moderate, longitudinal stripe on supraalar area; acrostichal (anterior and posterior) and dorsocentral (anterior and posterior) setae short; prescutellar area with several setae laterally; scutellum with narrow, curved scales on all lobes, median lobe also with or without few broad scales; antepronota widely separated, with broad, pale scales, several setae; postpronotum with scales, few posterior setae; prespiracular setae absent; hypostigmal area with broad, pale scales; postspiracular area with pale scales, several setae; subspiracular area with broad, pale scales; upper proepisternum with broad, pale scales, several setae, lower area bare; paratergite with pale scales; mesokatepisternum with upper and lower patches of broad, pale scales, upper and posterior setae; prealar area with broad, pale scales on upper and lower areas, numerous upper setae; mesepimeron with 2 patches of broad, pale scales, 1 on upper and 1 on middle areas, several upper setae, lower setae absent.

Wing:

Dark-scaled, with or without small, pale-scaled patch on anterobasal area of costa; upper calypter with several setae on margin; alula with row of narrow, dark scales on margin; remigium with 2 or 3 setae distally on dorsal surface.

Legs:

Femora without subapical pale-scaled bands; midfemur without median, longitudinal white-scaled stripe on anterior surface; hindfemur with apex with white scales; tibiae and tarsi dark-scaled; fore- and midtarsi with 2 equal ungues, each with 1 tooth; hindtarsi with 2 equal ungues, simple.

Abdomen:

Tergum I with pale-scaled patch on laterotergite; terga dark-scaled with basal pale-scaled areas; segment VII dorsoventrally flattened.

Genitalia:

Tergum VIII with apex broadly rounded, nearly covered with broad scales; sternum VIII sclerotized, width slightly greater than length, apical margin with small, median emargination, several broad scales on lateral areas, seta 2-S inserted posterior to 1-S; tergum IX comprising 2 moderately pigmented, lateral plates connected mesally by lightly pigmented area, several short setae distally on each plate; postgenital lobe relatively long, moderately wide, apex flat, several setae distally; upper vaginal sclerite small to moderate size; lower vaginal sclerite absent; insula liplike, few setae in lateral patches; cercus moderately long, moderately wide throughout length, apex broadly rounded, numerous broad scales on dorsal surface; 3 spermathecal capsules.

Males
Head:

Maxillary palpus with 5 palpomeres, shorter than proboscis, palpomeres 4 and 5 slightly down-turned; antennal flagellar whorls with numerous long setae directed dorsally and ventrally.

Legs:

Fore- and midtarsi with 2 unequal ungues, larger unguis with 2 teeth, smaller unguis with 1 tooth; hindtarsus with 2 equal ungues, simple.

Genitalia:

Tergum IX with median part relatively narrow with bilobed, pincushion-like structure bearing numerous, short to moderately long, very slender, curved setae, lateral area developed as large, oblong lobe projecting cephalad; gonocoxite moderately long, dorsal surface without apical and basal lobes on mesal surface, middle part of dorsal surface wide with mesal area broad, curved, projecting ventrally with row of short, stout, closely set, comb-like setae on mesal margin, much of dorsal surface covered with short to moderately long, very slender setae, ventral surface with elongate patch of numerous, long, moderately wide, fusiform scales on most of distal 0.50 of mesal margin, mesal surface membranous, lateral, outer part of dorsal and most of ventral surfaces with broad scales; gonostylus attached at apex of gonocoxite, long, narrow but proximal part somewhat thicker, somewhat sigmoid-shaped, single gonostylar claw short, curved, spiniform; aedeagus simple, tubelike; proctiger with few tiny, cercal setae; claspette with single, short to moderately long, narrow, columnar stem with small subapical lobe, claspette filament attached apically, long, curved, relatively slender throughout length, apex pointed; sternum IX with few setae (Gi. pulchriventer also with 2 or 3 broad scales) on posteromesal area.

Pupae
Cephalothorax:

Setae 1,3-CT similarly developed; 11-CT single, longer than 10-CT.

Abdomen:

Seta 3-I,II single, longer than 6-I,II; 6-I shorter than 7-I; 1-II long, with 3 branches; 5-II inserted lateral to 4-II; 3-III relatively short, single, approximately equal to 5-III length; 5-V moderately long, stout, single, shorter than tergum VI median length; 2-VI inserted mesal to 1,3-VI; 6-VII single, inserted slightly posterior and mesal to 9-VII; 9-VII with 3 branches; 9-VIII inserted on posterolateral corner, with multiple branches.

Paddle:

Apical margin rounded; midrib extends to apex of paddle; without hairlike spicules on margin; seta 1-Pa short, single.

Fourth-instar larvae
Head:

Seta 1-C slender, single, apex attenuate; 4-C short, with few slender branches, inserted mesal and slightly posterior to 6-C; 5-C moderately long, stout, lightly aciculate, with several branches, inserted mesal to 6-C; 6-C moderately long, stout, with 3 or 4 aciculate branches, inserted lateral and approximately same level to 5-C; 4–6-C inserted noticeably anterior to 7-C; 7-C moderately long, moderately stout, lightly aciculate, with 4 or 5 branches; 12-C short, with 2 or 3 slender branches, inserted mesal to 13-C; 13-C single, longer than 12-C; 14-C short, moderately stout, single or 2-branched; 19-C absent; antenna moderately long, narrow, with several small spicules, seta 1-A moderately long, with 3 or 4 (rarely 2) slender branches.

Thorax:

Setae 1–3-P on common setal support plate, 3-P multiple-branched; 4-P relatively long; 5-P long, stout, with 2 or 3 aciculate branches; 8-P short, stellate, with 5 or 6 branches, shorter than 4-P; 4-M single; 5-M longer than 7-M; 2,6-T single.

Abdomen:

Seta 6-I,II long, stout, aciculate, with 2 branches, 6-II longer than 6-III; 7-I,II long, stout, aciculate, single; 12-I present; 8-II short, with 2 branches; 1-VII long, stout, aciculate, single; 12-VII single; 2,4-VIII single; segment VIII with comb comprising numerous scales in patch; segment X with moderate saddle, incomplete ventrally, with numerous stout spicules on posterior margin dorsal to seta 1-X, acus absent, 1-X moderately long to long, stout, aciculate, single, inserted on saddle, ventral brush with several long, 2–5-branched setae with long stems, attached to grid with both transverse and lateral bars, anterior few setae somewhat shorter.

Siphon:

With attached acus; pecten with numerous closely set spines; seta 1-S relatively long with several aciculate branches, inserted very slightly distal to pecten.

Included species

Gilesius alius (Lien) and Gi. pulchriventer (Giles).

Distribution

China, India, Nepal, Pakistan and Taiwan.

Bionomics

Immature stages inhabit rock pools and treeholes but also have been collected in fresh water ground pools, a seepage pool, mud pools, barrels, tree stump holes, and holes in a fallen tree. Females of Gi. pulchriventer have been collected from humans at dusk.

Discussion

See Lien (1968b: figs 1, 6) for illustrations of the male genitalia, pupa and fourth-instar larva of Gi. alius.

Etymology

Gilesius is named in honour of Dr (Lt-Col) George Michael James Giles in recognition of his contributions to the taxonomy of mosquitoes, primarily of India and surrounding areas. The generic name is masculine, formed from his surname and the masculine Latin suffix ‘-ius’. Recommended abbreviation =Gi.

Himalaius Reinert, Harbach& Kitching, gen. nov.
Type species:

Finlaya gilliBarraud, 1924.

Females
Head:

Vertex with decumbent, narrow, curved, golden-white to white scales mesally, broad, golden-white to white scales laterally; occiput with numerous erect forked scales; clypeus bare; maxillary palpus and proboscis dark brown-scaled, proboscis approximately equal to forefemur length.

Thorax:

Scutum with narrow, curved scales, golden-white scales covering approximately anterior 0.60 of scutum, similar scales forming narrow stripe on posterior acrostichal area extending from anterior patch to anterior part of prescutellar area where it forks and extends along lateral margins of prescutellar bare area, similar narrow stripe on posterior dorsocentral area extending posteriorly along lateral margin of prescutellar setae, longitudinal stripe on supraalar area, scutal fossa with small, posteromedian patch of white scales, prealar area with small patch of somewhat broader, narrow, white scales; setae on posterior dorsocentral area; scutellum with narrow, curved, golden-white scales on all lobes; paratergite without scales; antepronota widely separated, with numerous moderately broad to broad, golden-white scales; postpronotum with golden-white scales, broad scales on most of area except narrow, curved scales dorsally; broad, white scales on upper proepisternum, mesokatepisternum (upper and lower posterior patches), lower prealar area and mesepimeron (single large patch), other areas without scales; mesepimeron without lower setae.

Wing:

Dark-scaled; upper calypter with several setae on margin; alula with row of narrow, dark scales on margin; remigium with 2 or 3 setae on dorsal surface distally.

Legs:

Ante- and postprocoxal membranes without scales; hindfemur with apex dark-scaled; hindtibia dark-scaled with basal band of white scales, band narrow dorsally, broader on anterior, posterior and ventral surfaces; hindtarsomeres 1–3 with broad, basal bands of white scales, hindungues equal, simple; fore- and midungues equal, each with 1 tooth.

Abdomen:

Tergum I with white-scaled patch on laterotergite; segment VII laterally compressed.

Genitalia:

Tergum VIII with broad scales covering all but narrow basal strip; sternum VIII sclerotized, slightly wider than long, several broad scales on proximal area, apical margin nearly straight with several moderately stout setae decreasing in length mesally, seta 2-S inserted posterior to 1-S; tergum IX comprising lightly to moderately pigmented sclerite with relatively deep, median, apical emargination separating pair of narrow lobes each bearing 3 or 4 short, slender setae; postgenital lobe relatively long and narrow, apex rounded, several setae distally; upper vaginal sclerite small, lightly pigmented; lower vaginal sclerite absent; insula liplike, with few setae in lateral patches; cercus relatively moderately long, moderately wide, apex rounded, mesal margin gently concave, without scales; 3 spermathecal capsules.

Males
Head:

Maxillary palpus with 5 palpomeres, 0.86–0.95 length of proboscis, dark-scaled, distal part of palpomere 3 bearing several long setae ventrolaterally, palpomeres 4 and 5 short, slightly down-turned, with several short and moderately long setae primarily on ventral and distal areas.

Legs:

Fore- and midtarsi with 2 unequal ungues, larger unguis with 2 teeth, smaller unguis with 1 tooth; hindungues equal, simple.

Genitalia:

Tergum IX comprising 2 moderately pigmented, lateral plates, each with small, short, narrow, heavily sclerotized lobe on posterior margin bearing 5 or 6 short, stout setae; gonocoxite moderately long, relatively narrow, dorsal surface without apical and basal lobes on mesal margin, several scales present, ventral surface without specialized setae or scales, mesal surface membranous; gonostylus long, attached at apex of gonocoxite, narrow but distal part narrower, nearly straight, with single relatively short, narrow, gonostylar claw at apex; aedeagus simple, tubelike, moderately long, relatively narrow; proctiger with distal part heavily pigmented, apex flattened with 2 or 3 short, stout teeth, cercal setae present; claspette with single long, narrow columnar stem, claspette filament attached at apex of stem, long, relatively narrow, slightly flattened, distal part terminating in long, slender, pointed apex; sternum IX relatively short, with several setae on posteromedian area.

Pupae
Cephalothorax:

Seta 1-CT very long, stout, with 2 branches; 3-CT very long, with 3–5 branches; 4-CT shorter than 5-CT; 7-CT long, with 3–5 branches; 11-CT relatively long, normally with 2 branches (single on 1 side of 1 specimen).

Abdomen:

Seta 3-I long, moderately stout, with 3 or 4 branches; 6-I long, slender, single or 2-branched, longer than 7-I; 2-II inserted lateral to 1-II and mesal to 3-II; 3-II long, stout, aciculate, with 2 or 3 branches; 4-II inserted mesal to 5-II; 6-II long, slender, single to 3-branched, longer than 7-II; 3-III long, stout, with 2 branches; 5-IV,V exceptionally long; 2-IV inserted mesal to 1,3-IV; 6-VII short, slender, inserted mesal and slightly anterior to 9-VII; 9-VII moderately long, stout, aciculate, with 3–5 branches; 9-VIII moderately long, stout, aciculate, with 10 or 11 branches, inserted noticeably far anteriorly from posterior margin.

Paddle:

Ovoid; midrib well developed, reaching apex of paddle; without hairlike spicules on lateral margins; seta 1-Pa very long, 0.65–1.02 length of paddle, single.

Fourth-instar larvae
Head:

With scooplike pouch anteroventral to antennal base; seta 1-C relatively short, moderately thickened throughout length but distal part slightly wider, apex bluntly pointed, single; 4-C very short, slender, with 6–9 branches, inserted near, mesal and slightly posterior to 5-C; 5,6-C very long, stout, dark, single, 5-C inserted near, mesal and slightly posterior to 6-C; 7-C long, stout, lightly aciculate, with 2 branches (3-branched on 1 side of 1 specimen), inserted anterior and lateral to 4–6-C and posterior to antennal base; 12-C short, slender, with 6–9 branches, inserted mesal to 13-C; 13-C moderately long, slender, with 3–5 branches; 19-C absent; antenna long, narrow, with few small spicules, seta 1-A single, extending beyond apex of antennal shaft.

Thorax:

Seta 2-P longer than 1,3-P; 4-P moderately long, slender, with 3 or 4 branches; 6-P longer than 7-P; 6,7-P longer than 5-P; 8-P relatively short, slender, with 2 or 3 branches; 1-M moderately long, slender, with 3–6 branches, approximately 2.9 times length of 2-M; 4-M with 3–6 branches; 5-M longer than 7-M; 2-T with 8–12 branches; 6-T with 3–5 branches.

Abdomen:

Seta 12-I present; 6,7-I moderately long, stout, 6-I with 4–6 branches, 7-I with 4 or 5 branches, only slightly shorter than 6-I; 6-II moderately long, stout, with 3 or 4 branches; 7-II relatively moderately long, slender, with 4–6 branches; 8-II single; 1-VII very long, moderately stout, with 2 or 3 branches; 3-VII short, slender with 3–8 branches, inserted lateral and slightly anterior to 1-VII; 7,10,12-VII branched; 1-VIII with 5 or 6 branches; 2,4-VIII branched; segment VIII with comb comprising 80–94 scales in patch; segment X with moderately large saddle, incomplete ventrally, acus absent, 1-X moderately long, slender, with 3 branches, inserted on saddle, 3-X very long, stout, single, ventral brush inserted on grid with both transverse and lateral bars, 2 precratal setae.

Siphon:

Acus attached; pecten comprising 24–36 evenly spaced spines; seta 1-S short, slender, with 6–8 branches, inserted distal to pecten.

Included species

Himalaius gilli (Barraud) and Hi. simlensis (Edwards). The latter species is provisionally placed in Himalaius.

Distribution

Himalaius gilli: India (Western Himalayas, Himachal Pradesh State, Krol Mountains near Solan, Naini Tal District, Uttar Pradesh State, Himachal Pradesh State, Uttaranchal State) and Nepal (Okaldunga District, Ramechhap District). Himalaius simlensis: India (Western Himalayas, Bhowali, Kumaon, Lansdowne, Muktesar, Simla) (Barraud, 1934), and Himachal Pradesh and Uttaranchal (Kaur, 2003).

Bionomics

Barraud (1924, 1934) reported that the immature stages of Hi. gilli (including those of the type series) were collected from water in treeholes. However, Kaur (1992) states that Barraud's paratypes deposited in the National Institute of Communicable Diseases, New Delhi, India, were reared from larvae collected in bamboo. Darsie, Pradhan & Vaidya (1992) collected pupae from turbid, chocolate-coloured water in treeholes. The immatures of Hi. simlensis are not known.

Discussion

Few specimens of Hi. gilli were available for examination but these bear distinct characters in all life stages that readily distinguish the genus (see above description based on Hi. gilli). Several abdominal segments of the larval specimens are in poor condition and could not be described. The species simlensis is provisionally included in Himalaius based on the characters mentioned in the brief and incomplete descriptions of Edwards (1922) and Barraud (1924).

Etymology

The generic name Himalaius is formed from ‘Himalaya’, the mountains, and the masculine Latin suffix ‘-ius’, meaning ‘pertaining to the Himalayas’. Recommended abbreviation = Hi.

Jarnellius Reinert, Harbach& Kitching, gen. nov.
Type species:

Culex varipalpus Coquillett, 1902.

Females
Head:

Vertex with decumbent scales narrow (few broad scales occasionally present laterally in some species); occiput and vertex with erect forked scales; eyes moderately separated above antennal pedicels; antennal pedicel with few to several, often partially overlapping, small, broad scales on mesal surface; clypeus bare; maxillary palpus dark with pale-scaled areas (all dark-scaled in Ja. muelleri); proboscis without median, pale-scaled band.

Thorax:

Scutum with moderately wide to wide, median, pale-scaled stripe or patch extending over anterior acrostichal area laterally to inner dorsocentral area or wider and extending laterally to mesal margin of scutal fossa, pale-scaled, median area normally continuing posteriorly to prescutellar area (except in Ja. varipalpus); narrow, curved, white scales on scutal fossa (lateral and posterior margins), supraalar area (elongate longitudinal line) and prescutellar space (lateral and anterior margins of median, bare area); acrostichal and dorsocentral areas with both anterior and posterior setae; paratergite with broad, pale scales; postpronotum with narrow, curved scales dorsally and broad scales ventrally (except all narrow scales in Ja. muelleri); postspiracular, upper proepisternal, lower prealar, mesokatepisternal (large upper patch and moderate lower posterior patch), mesepimeral (upper and lower patches, except 1 large patch in Ja. muelleri) and metameral (except Ja. deserticola) areas with broad, pale scales.

Wing:

Dark-scaled with small white-scaled patch at base of costa (pale-scaled patch reduced to 2 or 3 scales or occasionally absent in Ja. muelleri); upper calypter with several setae on margin; alula with row of narrow scales on margin; remigium without setae ventrally and normally absent dorsally (1 or 2 setae distally in Ja. muelleri and Ja. varipalpus).

Legs:

Anteprocoxal membrane bare, postprocoxal membrane normally with broad, pale scales (absent in Ja. deserticola and Ja. sierrensis); hindtarsomere 1 with broad, basal and apical, white-scaled bands, hindtarsomeres 2 and 3 with broad, apical, white-scaled bands (hindtarsus entirely dark-scaled in Ja. muelleri), ungues simple.

Abdomen:

Tergum I with white-scaled patch on laterotergite; segment VIII dorsoventrally flattened.

Genitalia:

Tergum VIII sclerotized, width greater than length, setae on distal 0.70 or more of tergum; sternum VIII sclerotized, width greater than length, seta 2-S inserted posterolateral to 1-S, scales normally present (not seen in only available specimen of Ja. monticola); tergum IX comprising single sclerite with moderately deep, median, apical emargination, 3–12 setae distally on each side of midline; postgenital lobe moderately long, relatively broad, apex broad with very small, median emargination, 16–25 setae distally; upper and lower vaginal sclerites absent; insula liplike, with setae in lateral patches; cercus moderately long, relatively narrow with apex sharply rounded, without scales (moderately wide with apex broadly rounded and 1–3 scales/cercus in Ja. muelleri); 3 spermathecal capsules.

Males
Head:

Maxillary palpus with 5 palpomeres, longer than proboscis, moderate number of moderately long to long setae ventrolaterally on distal part of palpomere 3 and ventrolaterally on palpomere 4; antennal flagellar whorls with numerous long setae directed dorsally and ventrally.

Legs:

Fore- and midtarsi with 2 unequal ungues, each with 1 tooth (smaller ungues simple in Ja. muelleri); hindungues equal, both simple.

Genitalia:

Tergum IX with posterior margin with pair of short, relatively narrow, heavily sclerotized lobes bearing several short, stout setae; gonocoxite moderately long, relatively narrow, mesal margin of dorsal surface without apical lobe but with small basal lobe bearing several long setae, 1 or more of these setae stout and somewhat flattened, ventral surface without specialized scales or setae, mesal surface membranous; gonostylus attached at apex of gonocoxite, moderately long, narrow, distal part slender and slightly curved mesally, with single moderately long, narrow gonostylar claw at apex; aedeagus simple, tubelike, moderately long, apex broadly rounded; proctiger with cercal setae; claspette with single moderately long, slender, columnar stem, claspette filament attached apically, relatively long, relatively narrow, somewhat flattened with distal part narrowing to pointed, slightly recurved apex; sternum IX with setae on posteromedian area.

Pupae
Cephalothorax:

Seta 1-CT longer than 3-CT, single or 2-branched; 11-CT single.

Abdomen:

Setae 3,6-I single, moderately stout; 2-II shorter and inserted mesal to 1-II; 3-II single, moderately long, stout, inserted mesal to 2-II; 5-II less than 3-II length, inserted mesal to 4-II (lateral to 4-II in Ja. muelleri); 3-III single, stout, longer than 5-III; 5-V single, less than tergum VI median length (slightly greater than tergum length in Ja. muelleri); 2,3-VI inserted lateral to 1-VI (? in Ja. laguna); 6-VII inserted posterior and mesal to 9-VII (at same level in Ja. muelleri); 9-VII longer and stouter than 6-VII; 9-VIII stout, with 2 or 3 branches.

Paddle:

Length greater than width; apex broadly rounded or with minute, broad, median emargination; midrib extending 0.60–0.70 from base; without fringe of hairlike spicules; seta 1-Pa short, single.

Fourth-instar larvae
Head:

Seta 1-C single, spiniform; 4-C short (barely moderately long in Ja. muelleri), multiple-branched, inserted mesal and slightly anterior to 6-C; 5-C moderately long (approaching long), single, inserted mesal and slightly anterior to 7-C and posterior to 4,6-C; 6-C moderately long, with 2 branches (single in Ja. muelleri); 7-C moderately long, multiple-branched; 12-C branched, inserted mesal to 13-C; 13-C single; 19-C very short; antenna without spicules (with few spicules in Ja. muelleri), seta 1-A single, 2,3-A inserted apically.

Thorax:

Setae 1–3-P inserted on common setal support plate; 4-P branched, longer than 8-P; 5-P branched; 4-M single (rarely 2-branched in Ja. sierrensis); 5-M longer than 7-M; 2,6-T single.

Abdomen:

Setae 3-I–VI and 2,4-VIII single; 6-I–III long, aciculate, branched; 7-I moderately long to long, stout, aciculate, single; 12-I absent (present in Ja. deserticola and Ja. sierrensis); 7-II moderately long, single, except relatively short and branched in Ja. muelleri; 1-VII longer than dorsal length of segment X (see method of measurement for character 49 in Character Descriptions); 2-VII inserted near 1-VII; 3-VII moderately long, inserted anterolaterad of 1-VII; segment VIII with comb comprising several scales in 1 or 2 irregular rows; segment X with saddle incomplete ventrally, without acus, seta 1-X single or 2-branched, inserted on saddle in small unsclerotized area (reduced in Ja. muelleri), 2-X branched, 3-X single, ventral brush attached to grid with only transverse bars (Ja. muelleri with both transverse and lateral bars), seta 4a-X long, except relatively short in Ja. muelleri, 4 relatively long, broad anal papillae with apices broadly rounded.

Siphon:

Acus present; pecten comprising several evenly spaced spines, on proximal 0.50 of siphon; seta 1-S branched, aciculate, inserted distal to pecten; 6-S short, single.

Included species

Jarnellius deserticola (Zavortink), Ja. laguna (Arnell & Nielsen), Ja. monticola (Belkin & McDonald), Ja. muelleri (Dyar), Ja. sierrensis (Ludlow) and Ja. varipalpus (Coquillett).

Distribution

Canada (British Columbia), Mexico (Baja California Sur, Distrito Federal, Durango, Hidalgo, Mexico, Sonora, Zacatecas) and United States (Arizona, California, Idaho, Montana, Nevada, New Mexico, Oregon, Texas, Utah, Washington).

Bionomics

Immature stages are usually found in treeholes. Jarnellius muelleri are found occasionally in artificial containers and were once collected from leaf axils of maguey (Zavortink, 1972). Insight into the ecology of treeholes in which most species of Jarnellius are found is given in Zavortink (1985).

Discussion

The above generic description of Jarnellius is additional to the description (Varipalpus Group) given by Arnell & Nielsen (1972), who should be referred to for the systematic background, descriptions and illustrations of the included species. Zavortink (1972) provided a description and illustrations of Ja. muelleri (as Ae. (Och). muelleri), the type species of the new subgenus Lewnielsenius. The genus includes two subgenera (see below). Myers (1967) described and illustrated eggs of Ja. sierrensis and Ja. varipalpus.

Etymology

Jarnellius is named in honour of Dr J. Hal Arnell in recognition of his important work on species of this genus and other New World aedines. The generic name is masculine, formed from his first initial ‘J’, surname and the masculine Latin suffix ‘-ius’. Recommended abbreviation = Ja.

Jarnellius SubgenusJarnellius Reinert, Harbach& Kitching
Females
Head:

Maxillary palpus dark-scaled with pale-scaled areas.

Thorax:

Scutal fossa with white-scaled lines on lateral and posterior margins that are not expanded mesally; postpronotum with narrow, curved scales dorsally and broad scales ventrally; hypostigmal area without scales; mesepimeron with upper and lower patches of white scales.

Legs:

Hindtibia dark-scaled with white scales basally; hindtarsomere 1 with broad basal and apical, white-scaled bands, hindtarsomeres 2 and 3 with broad, apical, white-scaled band; fore- and midungues each with 1 tooth.

Genitalia:

Cercus moderately long, moderately wide, apex sharply rounded, without scales.

Males
Legs:

Fore- and midungues each with 1 tooth.

Pupae
Abdomen:

Seta 5-II inserted lateral to 4-II; 5-V shorter than median length of tergum VI; 6-VII inserted posterior and mesal to 9-VII.

Fourth-instar larvae
Head:

Seta 6-C with 2 branches; antenna without spicules.

Abdomen:

Seta 7-II developed similarly to 7-I, single (occasionally 2-branched in Ja. sierrensis), moderately long to long; ventral brush attached to grid with only transverse grid bars, seta 4a-X long.

Distribution

Canada (British Columbia), Mexico (Baja California Sur, Sonora) and United States (Arizona, California, Idaho, Montana, Nevada, New Mexico, Oregon, Utah, Washington).

Included species

Jarnellius deserticola, Ja. laguna, Ja. monticola, Ja. sierrensis and Ja. varipalpus.

Recommended abbreviation = Jar.

Jarnellius SubgenusLewnielsenius Reinert, Harbach& Kitching, subgen. nov.
Type species:

Aedes (Heteronycha) muelleri Dyar, 1920.

Females
Head:

Maxillary palpus dark-scaled.

Thorax:

Scutal fossa with white-scaled areas on lateral and posterior margins that are expanded mesally; postpronotum with narrow scales; hypostigmal area with pale scales; mesepimeron with single, large patch of white scales extending over upper and lower areas.

Legs:

Hindtibia and hindtarsus entirely dark-scaled; fore- and midungues simple.

Genitalia:

Cercus moderately long, moderately wide, apex broadly rounded, with 1–3 scales.

Males
Head:

Maxillary palpus dark-scaled.

Legs:

Fore- and midtarsi with larger ungues with 1 tooth, smaller ungues simple.

Pupae
Abdomen:

Seta 5-II inserted mesal to 4-II; 5-V longer than median length of tergum VI; 6-VII inserted mesal and at approximately same level as 9-VII.

Fourth-instar larvae
Head:

Seta 6-C single; antenna with few spicules.

Abdomen:

Seta 7-II developed differently from 7-I, relatively short, branched; ventral brush attached to grid with both transverse and lateral grid bars, seta 4a-X relatively short.

Distribution

Mexico (Distrito Federal, Durango, Hidalgo, Mexico, Zacatecas) and United States (Arizona, New Mexico, Texas).

Included species

Jarnellius muelleri (Dyar).

Etymology

Subgenus Lewnielsenius is named in honour of Dr Lewis T. Nielsen in recognition of his many contributions to the taxonomy, distribution and bionomics of North American mosquitoes, especially aedine species. The subgenus is masculine, formed from the combination of his first name, surname and the Latin masculine suffix ‘-ius’. Recommended abbreviation = Lew.

Jihlienius Reinert, Harbach& Kitching, gen. nov.
Type species:

Aedes (Finlaya) chungiLien, 1968.

Females
Head:

Vertex with decumbent scales broad, pale on median area and dark on lateral areas; erect forked scales numerous on occiput, few semierect scales on vertex caudal to ocular setae; ocular line narrow, covered with broad, pale scales; 2 long interocular setae; eyes above antennal pedicels contiguous or separated by less than width of 1 eye facet; antennal pedicels with few small, broad, dark scales on mesal surface; clypeus bare; maxillary palpus and proboscis dark-scaled; proboscis longer than forefemur.

Thorax:

Covered with narrow, relatively straight scales except bare, median, prescutellar area and elongate strip between supraalar and prescutellar setae, scales dark except for following pale-scaled areas: narrow stripe on anterior promontory and extending caudally on acrostichal area to approximately level of scutal angle where it becomes less distinct for short distance and then distinct on caudal part of posterior acrostichal area, narrow line on caudal part of anterior dorsocentral area and extending over posterior dorsocentral area to caudal margin of scutum, stripe broadened at juncture of prescutal suture into small triangular patch which connects with narrow stripe extending outward along scutal suture and cephalad along margin of scutal fossa onto antedorsocentral area, more or less elongate stripe on supraalar area, and indistinct stripe on lateral margins of prescutellar area mesad of setae; setae on following: few on anterior promontory, antedorsocentral area, dorsocentral (anterior and posterior) area, scutal fossa and antealar area, numerous on supraalar and prescutellar areas; scutellum with broad or broad and narrow scales on lobes; antepronota widely separated, with broad, pale scales, several setae; postpronotum with pale scales, few posterior setae; prespiracular setae absent; hypostigmal and subspiracular areas bare; postspiracular area with few setae, scales absent; upper proepisternum with broad, pale scales, several setae on upper area, lower area bare; paratergite narrow, with broad, pale scales; mesokatepisternum with broad, pale scales in upper and lower patches, 3 or 4 upper and several posterior setae; prealar area with broad, pale scales in patch on lower area and extending onto lower part of knob, several upper setae; mesepimeron with 1 patch of broad, pale scales on upper area and extending onto middle, several upper setae, lower setae absent.

Wing:

Dark-scaled with small pale-scaled patch at base of costa; upper calypter with several setae on margin; alula with row of narrow, dark scales on margin (Ji. chungi); remigium with 2 or 3 setae distally on dorsal surface.

Legs:

Hindfemur with narrow basal and moderate preapical bands dark-scaled, remainder pale-scaled; hindtibia and hindtarsus dark-scaled except tarsomere 1 with broad, pale-scaled, basal band; fore- and midtarsi with 2 equal ungues, each with 1 tooth; hindtarsus with 2 equal ungues, simple.

Abdomen:

Tergum I with pale-scaled patch on lateral tergite; terga dark-scaled with small, basolateral, pale-scaled patch; segment VII laterally compressed.

Genitalia:

Tergum VIII with apex broadly rounded, numerous broad scales on most of surface; sternum VIII sclerotized, width greater than length, apical margin with median emargination, scales absent, seta 2-S inserted posterior to 1-S; tergum IX comprising 2 moderately pigmented, moderately long, lateral plates connected mesally by basal pigmented strip, base wider than apex, few short setae distally on each plate; postgenital lobe long, relatively narrow, apex rounded, several setae distally; upper vaginal sclerite small; lower vaginal sclerite absent; insula liplike, with few setae in lateral patches; cercus moderately long, relatively wide, outer distal margin gently oblique, without scales; 3 spermathecal capsules.

Males
Head:

Maxillary palpus with 5 slender palpomeres in nearly straight line, shorter than proboscis, few moderately long setae on distal part of palpomere 3 ventrally and palpomere 4 ventrolaterally; antennal flagellar whorls with numerous long setae directed dorsally and ventrally.

Thorax:

Scaled areas of scutum with narrow, relatively long, pale scales except small area of scutal fossa with narrow, dark scales.

Legs:

Fore- and midtarsi with 2 unequal ungues, larger unguis with 2 teeth, smaller unguis with 1 tooth; hindtarsus with 2 equal ungues, simple.

Genitalia:

Tergum IX with median area narrow, lateral area developed as broad, rounded lobe projecting cephalad, posterior margin with very small, sclerotized lobe on each side of midline bearing few short, slender, curved setae; gonocoxite moderately long, relatively narrow, dorsal surface without apical and basal lobes on mesal margin, however, with small, dark, oval area basomesally bearing straight setae, few of these setae on most proximal part long and somewhat flattened, middle part with several moderately long setae and distal part with several short setae, ventral surface with narrow, elongate patch of moderately long, slender, lanceolate setae on distal area of mesal margin (Lu & Ji, 1997: fig. 34, shows narrow fusiform scales in this area of Ji. gonguoensis), mesal surface membranous, lateral and outer parts of dorsal and ventral surfaces with broad scales; gonostylus attached at apex of gonocoxite, relatively short, narrow, single gonostylar claw, long, narrow, attached at apex; aedeagus simple, tubelike, apex concave; proctiger with few tiny, cercal setae; claspette with single, moderately long, narrow, columnar stem, claspette filament attached apically, long, somewhat flattened beyond midlength, with transverse striations on much of median area, apex curved into narrow point; sternum IX with few setae on posteromedian area.

Pupae
Cephalothorax:

Setae 1,3-CT similarly developed; 10-CT single; 11-CT with 2 branches, shorter than 10-CT.

Abdomen:

Seta 3-I,II single, shorter than 6-I,II; 6-I longer than 7-I; 1-II short, slender, with 2 branches; 5-II inserted lateral to 4-II; 3-III single, longer than 5-III; 5-V long, stout, single, longer than tergum VI median length; 2-VI inserted mesal to 1,3-VI; 6-VII inserted slightly posterior and mesal to 9-VII, 6-VII single, 9-VII with 2 or 3 branches; 9-VIII inserted on posterolateral corner, with 3–5 branches.

Paddle:

Apical margin rounded; midrib extends to apex of paddle; without hairlike spicules on margins; seta 1-Pa short, single or 2-forked.

Fourth-instar larvae
Head:

Seta 1-C stout, spiniform; 4-C short, slender, multiple-branched, inserted slightly mesal and approximately in line with 6-C; 5-C moderately long, stout, single, inserted posterior to 4,6,7-C; 6-C moderately long but shorter than 5-C, moderately stout, single, inserted anterior and approximately same lateral level to 5-C, mesal and anterior to 7-C; 7-C relatively short, with 3–5 slender branches, inserted lateral and slightly anterior to 5-C; 12-C short, with few branches, inserted mesal to 13-C; 13-C single, longer than 12-C; 14-C short, slender, single; 19-C absent; antenna moderately long, narrow, without spicules, seta 1-A moderately long, slender, single.

Thorax:

Setae 1–3-P inserted on common setal support plate; 4-P single or occasionally 2-branched; 5-P normally with 2 branches; 8-P short, with few branches, shorter than 4-P; 4-M single; 5-M longer than 7-M; 2,6-T single.

Abdomen:

Seta 6-I,II with 2 or 3 moderately long, moderately stout branches; 7-I moderately long, moderately stout, with 2 branches; 6-II longer than 6-III; 12-I present; 7-II short, slender, with 3 or 4 branches; 8-II with 2 or 3 branches; 1-VII moderately long, stout, single; 12-VII single; 2,4-VIII single; comb on segment VIII comprising patch of more than 20 scales; segment X with moderately large saddle, incomplete ventrally, with several moderately long, stout spicules on posterior margin dorsal to insertion of seta 1-X, acus absent, seta 1-X moderately long, slender, single or 2-branched, inserted on saddle, ventral brush with long, normally 2–4-branched setae with long stems, inserted on grid with both transverse and lateral bars, 2 shorter, precratal setae.

Siphon:

With attached acus; pecten with numerous, closely set spines; seta 1-S moderately long, slender, multiple-branched, inserted slightly distal to pecten.

Included species

Jihlienius chungi (Lien) and Ji. unicinctus (Edwards). Jihlienius gonguoensis (Gong & Lu) is provisionally included based on illustrations in Lu & Ji (1997).

Distribution

China, India, Nepal and Taiwan.

Bionomics

Immatures have been collected from bamboo stumps and treeholes.

Discussion

Much of the generic description is based on Ji. chungi because published descriptions of other species were inadequate, several life stages are unknown and/or specimens were not available for study. See Lien (1968a) for illustrations of the male genitalia, pupa and fourth-instar larva of Ji. chungi.

Etymology

Jihlienius is named in honour of Dr Jih Ching Lien in recognition of his important contributions to the taxonomy of the mosquitoes of Taiwan. The generic name is masculine, formed from his first name, surname and the Latin suffix ‘-ius’. Recommended abbreviation = Ji.

Patmarksia Reinert, Harbach& Kitching, gen. nov.
Type species:

Leucomyia australiensis variety papuensis Taylor, 1914.

Females
Head:

Vertex with median area covered with narrow, curved, pale scales; occiput and vertex with erect forked scales; interocular space with patch of narrow, curved, pale scales projecting ventrad over mesal margins of eyes; eyes contiguous or nearly so; antennal pedicel with few short, fine setae and small, broad, nonoverlapping scales on mesal surface; clypeus bare; maxillary palpus and proboscis dark-scaled.

Thorax:

Scutum with narrow, curved, dark scales except for moderate to broad median, longitudinal patch of white scales on approximately anterior 0.55–0.70 (white-scaled patch on acrostichal area and extending laterally over mesal part of dorsocentral area or to near lateral part of scutal fossa, but not on lateral part of scutal fossa), other scutal areas normally dark-scaled including supraalar and prescutellar areas (but not median, prescutellar bare area) (few species with white scales along lateral margin of prescutellar bare area); scutellum with narrow, curved scales on all lobes; acrostichal (both anterior and posterior) areas without setae, anterior dorsocentral area normally without setae (occasionally 1 seta on caudal part), posterior dorsocentral area with several setae; paratergite with broad, pale scales; postpronotum normally with patch of broad, scales ventrally and narrow, curved scales dorsally (no narrow scales seen in Pm. dobodura but this area rubbed in specimens examined; only narrow, curved, pale scales in Pm. anggiensis); mesokatepisternum with upper and lower posterior patches of broad, pale scales; metameron bare.

Wing:

Costa dark-scaled; upper calypter with several setae on margin; alula with row of narrow scales on margin; remigium with dorsal setae distally.

Legs:

Ante- and postprocoxal membranes bare; hindfemur with dorsoapical patch of pale scales; tibiae dark-scaled (except narrow, pale-scaled, basal band in Pm. anggiensis); hindtarsus with broad, basal, pale-scaled bands on tarsomeres 1–4.

Genitalia:

Tergum VIII sclerotized, with numerous broad scales on distal part; sternum VIII sclerotized, length greater than width, normally moderate to large number of broad scales in lateral patches, apex with small to moderately deep, median emargination separating pair of small to broad lobes, apical margin with numerous short, curved setae, seta 2-S inserted posterior to seta 1-S; tergum IX comprising 2 moderately pigmented, narrow, lateral plates connected by basal or subbasal pigmented or membranous band, distal part of mesal surfaces of plates darkly pigmented and strongly fused to basolateral margins of cerci, without setae; postgenital lobe with several setae distally; upper vaginal sclerite small, moderately pigmented; lower vaginal sclerite absent; insula liplike, V-shaped medially (depressed), with setae in lateral patches; cercus moderately long, moderately wide, apex broadly rounded, without scales (rarely 1 or 2 scales present); 3 spermathecal capsules.

Males
Head:

Maxillary palpus with 5 palpomeres, slightly shorter to slightly longer than proboscis, numerous long setae ventrolaterally on distal part of palpomere 3 and ventrolaterally on palpomere 4; antennal flagellar whorls with numerous long setae directed dorsally and ventrally.

Legs:

Fore- and midtarsi with 2 ungues, both toothed.

Genitalia:

Tergum IX comprising 2 narrow, lateral plates narrowly joined mesally, without setae; gonocoxite moderately long, moderately wide, mesal margin of dorsal surface without apical lobe but with small basal lobe bearing several short and moderately long, slender setae, ventral surface without specialized setae or scales on mesal margin, mesal surface membranous; gonostylus attached at apex of gonocoxite, moderately long, narrow, with single, short, narrow, gonostylar claw at apex; aedeagus simple, tubelike, moderately long, distal 0.50 widest; proctiger with cercal setae; claspette with single moderately long, slender, columnar stem, claspette filament attached apically, moderately long to long, somewhat flattened, relatively narrow with distal part tapering to slender apex, middle part with transparent sheaf with transverse striations encircling spiniform; sternum IX with setae on posteromedian area.

Pupae
Cephalothorax:

Seta 11-CT single (rarely 2-branched in Pm. argyronotum).

Abdomen:

Seta 3-I long, stout, single, longer than 6-I; 6-I single, longer than 7-I; 2-II short, inserted lateral to 1-II; 3-II long, stout, single, longer than 6-II; 3-III long, stout, single, longer than 5-III; 5-IV,V long, stout, longer than following tergum; 6,9-VII both relatively stout, aciculate, branched; 9-VIII stout, aciculate, multiple-branched.

Paddle:

Broad; apex broadly rounded or flattened; midrib nearly reaching apex; without hairlike spicules on margin; seta 1-Pa single.

Fourth-instar larvae
Head:

Seta 1-C relatively stout, single; 4-C short, branched, inserted slightly posterior and mesal to 5-C; 5,6-C moderately long, multiple-branched, displaced far anteriorly, 6-C inserted lateral and slightly anterior to 5-C; 7-C moderately long, multiple-branched, inserted lateral and posterior to 4–6-C; 12-C short, branched, inserted mesal to 13-C; 13-C single; 14-C short, normally single; 19-C absent; antenna with few to numerous spicules, 1-A normally branched.

Thorax:

Setae 1–3-P inserted on common setal support plate; 8-P short, normally branched; 1-M,T not on tubercle, 1-M moderately long, 1-T short; 6-T single.

Abdomen:

Seta 7-I long, stout, normally single; 12-I present; 7-II short, branched; 7,10,12-VII single; 2,4-VIII single; segment VIII with comb normally with numerous scales in patch (some species with several scales in 1 or 2 irregular rows); segment X with saddle incomplete ventrally, moderately developed spicules on posterior margin dorsal to seta 1-X (occasionally spicules smaller), ventral brush with setae attached to grid with both transverse and lateral bars, seta 1-X single (except 2 or 3 branches in Pm. argenteitarsis and Pm. dobodura), inserted on saddle.

Siphon:

With attached acus; pecten with spines evenly spaced; seta 1-S branched, inserted distal to pecten.

Included species

Patmarksia anggiensis (Bonne-Wepster), Pm. argenteitarsis (Brug), Pm. argyronotum (Belkin), Pm. buxtoni (Belkin), Pm. clintoni (Taylor), Pm. derooki (Brug), Pm. dobodura (King & Hoogstraal), Pm. hollandia (King & Hoogstraal), Pm. mackerrasi (Taylor), Pm. novalbitarsis (King & Hoogstraal), Pm. palmarum (Edwards), Pm. papuensis (Taylor) and Pm. subalbitarsis (King & Hoogstraal).

Distribution

Australia (Queensland), Indonesia (Irian Jaya, Moluccas), Papua New Guinea and Solomon Islands (Guadalcanal, New Georgia).

Bionomics

Immature stages of Patmarksia species are found in plant containers (e.g. fallen palm fronds, coconut husks and leaves, treeholes, holes in fallen logs, cut bamboo, funnel-shaped fungi), artificial containers (e.g. metal cans, drum heads, flower bowls), and small puddles, ruts, borrow pits and rock pools in stream beds.

Discussion

See King & Hoogstraal (1946: pls 20, 21, 23) for illustrations of the male genitalia, fourth-instar larva and adults of Pm. papuensis, Penn (1949: fig. 35) for the pupa and Reinert (2002e: fig. 9) for the female genitalia.

Etymology

Patmarksia is named in honour of Dr Elizabeth Nesta Marks in recognition of her pioneering and outstanding contributions to the taxonomy of Australasian mosquitoes. The generic name is feminine, formed from her nickname Pat’, her surname and the feminine Latin suffix ‘-ia’. Recommended abbreviation =Pm.

Rampamyia Reinert, Harbach& Kitching, gen. nov.
Type species:

Culex notoscriptus Skuse, 1889.

Females
Head:

Vertex with decumbent scales both broad and narrow; occiput and vertex with erect forked scales; eyes contiguous or separated by less than or equal to 1 eye facet; antennal pedicel with patch of overlapping, broad, silvery scales on mesal surface; clypeus bare; maxillary palpus with apex pale-scaled; proboscis with well-developed pale-scaled band at about midlength.

Thorax:

Acrostichal area with narrow stripe of pale scales on anterior and posterior areas; posterior dorsocentral area with narrow stripe of pale scales; scutal fossa with narrow stripe of pale scales on lateral and usually posterior margins; supraalar area with narrow, longitudinal patch of pale scales; prescutellar median, bare area with pale scales laterally mesal to setae; acrostichal and dorsocentral areas with both anterior and posterior setae; scutellum with broad scales on all lobes; scutal fossa with 1 or 2 posterior setae; paratergite with broad, pale scales; broad scales on antepronotum, postpronotum, subspiracular area, upper proepisternum, lower prealar area, upper and lower posterior mesokatepisternum, and upper and lower mesepimeron, other pleural areas without scales; mesepimeron without lower setae.

Wing:

Dark-scaled; upper calypter with several setae on margin; alula with row of narrow scales on margin; remigium without dorsal and ventral setae.

Legs:

Ante- and postprocoxal membranes bare; mid- and hindfemora with median stripe of pale scales on anterior surface; hindtarsus with wide, basal, pale-scaled bands on tarsomeres 1–4; fore- and midungues equal, each with 1 tooth; hindungues equal, simple.

Abdomen:

Tergum I with silvery-scaled patch on lateral tergite; segment VII laterally compressed.

Genitalia:

Tergum VIII sclerotized, width greater than length, without (rarely with 1 or 2 adventitious) scales, setae on distal 0.30 or less of tergum; sternum VIII sclerotized, width greater than length, without (rarely with 1 or 2 adventitious) scales, apex broad, each side nearly flat and very gently sloping to midline, seta 2-S inserted posterior to seta 1-S; tergum IX comprising single sclerite with moderately deep, median, apical emargination, 2–7 setae distally on each side of midline; postgenital lobe relatively long, narrow, with spicules well developed on lateral margins, few setae distally; upper vaginal sclerite well developed, moderately to heavily pigmented; lower vaginal sclerite absent; insula liplike, with 5–7 setae in lateral patches; cercus moderately long, moderately wide, apex broadly rounded, without scales; 3 spermathecal capsules.

Males
Head:

Maxillary palpus with 5 palpomeres, longer than proboscis, numerous long setae ventrolaterally on distal part of palpomere 3 and ventrolaterally on palpomere 4; antennal flagellar whorls with numerous long setae directed dorsally and ventrally.

Legs:

Fore- and midtarsi with 2 unequal ungues, both toothed; hindtarsus with 2 equal ungues, simple.

Genitalia:

Posterior margin of tergum IX with pair of short, narrow, acute lobes bearing few slender to moderately stout setae; gonocoxite relatively long, relatively narrow, mesal margin of dorsal surface without apical and basal lobes, ventral surface with patch of long, narrow, lanceolate setae on distal part of mesal margin, mesal surface membranous; gonostylus attached at apex of gonocoxite, relatively short, less than or equal to 0.25 length of gonocoxite, narrow, distal part slender and slightly curved mesally, with single long, narrow gonostylar claw at apex; aedeagus simple, tubelike, noticeably long and narrow; paramere relatively long and narrow; proctiger with cercal setae; claspette with single, long, slender, columnar stem, claspette filament attached apically, flattened, curved with distal part tapering to bluntly pointed apex; sternum IX with setae on posteromedian area.

Pupae
Cephalothorax:

Seta 11-CT single.

Abdomen:

Setae 3,6,7-I single; 6-I longer than 7-I; 3-II long, single, inserted mesal to 2-II; 5-II long, single; 5-IV,V stout, aciculate, each longer than the following tergum; 2-VI inserted mesal to 1-VI; 6-VII inserted posterior and mesal to 9-VII, noticeably shorter and thinner than 9-VII; 9-VIII with multiple, aciculate branches.

Paddle:

Apex broadly rounded; without fringe of hairlike spicules; seta 1-Pa short, single.

Fourth-instar larvae
Head:

Seta 1-C stout, single, sharply curved, spiniform; 4-C short, with very slender branches, inserted slightly mesal and slightly posterior to 6-C; 4,6-C displaced far anterior; 5-C inserted posterior and slightly lateral to 4,6-C and mesal and slightly anterior to 7-C; 7-C moderately long, with few branches; 5,6,13-C single; 19-C present; antenna without spicules, 1-A short, single (rarely 2-branched), inserted on distal third of antennal shaft.

Thorax:

Setae 1–3-P not on common setal support plate; 8-P short, multiple-branched; 1-M,T slender, branched, not on tubercle; 5-M very long, single, noticeably longer than 7-M; 6-T single.

Abdomen:

Seta 7-I long, single; 12-I present; 7-II short, branched; 1,3,7,10,12-VII single; 2,4-VIII single; segment VIII with comb comprising numerous scales in patch; segment X with saddle incomplete ventrally, with well-developed spicules on posterior margin dorsal to seta 1-X, ventral brush with several long, branched setae with relatively long stems, attached to grid with both transverse and lateral bars, precratal setae absent, 1-X multiple-branched, inserted on saddle.

Siphon:

Relatively short; pecten with spines evenly spaced; seta 1-S branched, inserted distal to pecten.

Included species

Rampamyia albiabris (Edwards), Ra. notoscripta notoscripta (Skuse), Ra. notoscripta montana (Brug) and Ra. quinquelineata (Edwards).

Distribution

Australia, Bismarck Archipelago (New Britain, New Ireland), Indonesia (Java, Moluccas), New Caledonia (Ile des Pins, Loyalty Islands), New Zealand (North Island, South Island), Papua New Guinea and Solomon Islands (Bellona, Bougainville, Florida, Guadalcanal, Kolombangara, Malaita, New Georgia, Rendova, Rennell, Roviana, Sterling, Ugi).

Bionomics

Immature stages occur in plant containers, including treeholes, bamboo, plant leaf axils, palm fronds, coconut shells and husks, and various small and large artificial containers made of metal, cement and wood, and rockholes.

Discussion

See Belkin (1962: figs 226–229) for illustrations of the male genitalia, pupa and fourth-instar larva of Ra. notoscripta, Russell (1996: 73) for photographs of the female and Linley et al. (1991a) for a description and photographs of the egg.

Etymology

Rampamyia is named in honour of Dr Rampa Rattanarithikul in recognition of her outstanding contributions to the taxonomy of Oriental mosquitoes. The generic name is feminine, formed from her first name and the feminine Greek noun ‘-myia’, meaning ‘fly’. Recommended abbreviation = Ra.

Vansomerenis Reinert, Harbach& Kitching, gen. nov.
Type species:

Aedes (Finlaya) pulchrithorax Edwards, 1939.

Females
Head:

Vertex with median, pale-scaled patch; vertex and occiput with erect forked scales; ocular line narrow, with broad, silvery scales; eyes contiguous; clypeus bare; maxillary palpus and proboscis dark-scaled.

Thorax:

Scutum with narrow, curved scales; pale scales forming relatively narrow stripe on anterior and posterior acrostichal areas and extending to prescutellar area, similar stripe on posterior dorsocentral area extending caudally laterad of prescutellar setae to posterior margin of scutum, elongate, longitudinal patch on supraalar area, prescutellar area with narrow patches on lateral and anterior parts of median, bare area; scutal fossa covered with narrow, curved, dark scales; dorsocentral area with anterior and posterior setae; acrostichal area without setae; scutellum with narrow, curved scales on all lobes; mesopostnotum bare; paratergite with broad, silvery scales; broad, silvery scales on antepronotum, postpronotum (large patch), upper proepisternum, mesokatepisternum (small upper and lower posterior patches), lower prealar area and mesepimeron (single large patch extending over upper and lower areas); subspiracular and postspiracular areas without scales; upper proepisternum with 3–5 setae; mesepimeron without lower setae.

Wing:

Dark-scaled; upper calypter with several setae on margin; alula with row of narrow scales on margin; remigium with few dorsal setae distally.

Legs:

Ante- and postprocoxal membranes bare; hindfemur without pale scales dorsoapically; hindtibia with broad, basal, white-scaled patch; hindtarsus with white-scaled, basal bands on tarsomeres 1 and 2, narrow on tarsomere 1, broad on tarsomere 2; fore-, mid- and hindungues equal, each with 1 tooth.

Abdomen:

Tergum I with silvery-scaled patch on lateral tergite; segment VII laterally compressed.

Genitalia:

Tergum VIII with numerous broad scales on much of surface; sternum VIII sclerotized, width greater than length, apical margin nearly flat or with very shallow, broad, median emargination, with several stout setae decreasing in length mesally, few to several broad scales, seta 2-S inserted posterior to 1-S; tergum IX comprising 2 long, very narrow, darkly pigmented sclerites connected on approximately mesal 0.70 by moderately pigmented band, 2 or 3 short setae apically on each side of midline; postgenital lobe long, relatively narrow, with several setae distally; upper and lower vaginal sclerites absent; insula liplike, with few setae in lateral patches; cercus moderately long, relatively narrow, apex sharply rounded, without scales; 3 spermathecal capsules.

Males
Head:

Maxillary palpus with 5 palpomeres, all relatively slender and in approximately straight line, dark-scaled, slightly longer to slightly shorter than proboscis, palpomeres 4 and 5 with few setae; antennal flagellar whorls with numerous long setae directed dorsally and ventrally.

Legs:

Fore-, mid- and hindtarsi each with 2 ungues, each with 1 tooth.

Genitalia:

Tergum VIII with small, median lobe on posterior margin; tergum IX comprising 2 very small sclerites connected mesally by tonguelike, median, cephalad-directed lobe of tergum X, each sclerite with small, strongly sclerotized lobe on posterior margin bearing few short, stout setae, lateral part of lobe connected by narrow band with sternum IX; gonocoxite moderately long, moderately wide, dorsal surface with scales all approximately same length, mesal margin without apical lobe, with moderate-size, basal lobe, distal part of lobe narrow and bearing 2 or 3 moderately long to long, moderately stout setae whereas proximal part of lobe is expanded with series of wrinkles at base and bearing several short to moderately long setae (some of which may be narrowly lanceolate), ventral surface without specialized scales or setae, mesal surface membranous; gonostylus attached at apex of gonocoxite, relatively short, slender, single gonostylar claw moderately long, narrow, attached at apex; aedeagus simple, tubelike, moderately long; claspette with single moderately long, slender, columnar stem, claspette filament long, proximal part narrow, distal part expanded, flattened but terminating in pointed apex; proctiger with few cercal setae; sternum IX with few setae on posteromedian area.

Pupae
Cephalothorax:

Setae 1,3-CT branched; 7-CT very long, greater than 6.0 times length of 6-CT; 8-CT moderately long, moderately stout, aciculate, with 5–14 branches; 9-CT moderately long, moderately stout, with 2–6 branches; 10-CT relatively short, very slender, with 6–19 branches; 11-CT long, stout, single.

Abdomen:

Seta 3-I long, with 2 or 3 branches; 6-I very long, stout, aciculate, single, noticeably longer than 7-I; 7-I long, stout, aciculate, normally with 2 branches (rarely single or 3-branched); 2-II shorter and inserted lateral to 1-II and mesal to 3-II; 3-II,III long, stout, single; 5-II shorter than 3-II; 6-II long, stout, aciculate, single, longer than 7-II; 7-II long, branched; 5-IV–VI long, stout, aciculate, longer than following tergum; 2-VI inserted mesal to 1,3-VI; 6-VII short, slender, with 4–10 branches, inserted posterior and mesal to 9-VII; 9-VII stout, aciculate, branched, longer than 6-VII; 9-VIII stout, aciculate, multiple-branched, inserted near posterolateral corner of segment VIII.

Paddle:

Length greater than width, index 1.20–1.42; without hairlike spicules on margins; midrib reaching apex of paddle; seta 1-Pa single.

Fourth-instar larvae
Head:

Seta 1-C single, spiniform; 4-C short, very slender, branched, inserted mesal and slightly posterior to 6-C; 5-C long, stout, aciculate, often single (range single to 3-branched), inserted posterior and mesal to 6-C and posterior and lateral to 4-C; 6-C long, stout, aciculate, longer than 5-C, inserted mesal and very slightly anterior to 7-C; 7-C moderately long, relatively stout, aciculate, with 3–6 branches; 12-C branched, inserted mesal to and shorter than 13-C; 13-C branched; 19-C absent; antenna relatively long, slender, slightly curved mesally, with few scattered spicules, seta 1-A moderately long, single or 2-branched (occasionally 3-branched).

Thorax:

Seta 5-P branched, shorter than 6-P; 8-P short; 4-M branched; 6-T single.

Abdomen:

Setae 6-I,II long, branched (6-II rarely single in Va. luteostriata); 6-III branched, shorter than 6-II; 7-I long, with 2 or 3 branches; 7-II short, with 4–15 branches, differently developed than 7-I; segment VIII with comb comprising numerous scales in patch; 2-VIII branched; segment X with saddle incomplete ventrally, acus absent, setae 1-X short, branched, inserted on saddle, ventral brush with several long, branched setae with short stems, attached to grid with well developed transverse grid bars, lateral bars absent but represented by faint depression, 2 precratal setae.

Siphon:

With acus attached; pecten with number of evenly spaced spines; seta 1-S branched, inserted distal to pecten.

Included species

Vansomerenis hancocki (van Someren), Va. luteostriata (Robinson) and Va. pulchrithorax (Edwards).

Distribution

Angola, Kenya, Tanzania, Uganda, Zaire and Zambia.

Bionomics

Immature stages have been found in treeholes, a rot hole in the fork of a tree, a tree buttress and bored bamboo.

Discussion

See Edwards (1941: figs 6m, 176 and pl. III-14) for illustrations of the female genitalia, fourth-instar larva and female of Va. pulchrithorax. Hopkins (1952: figs 62, 63) also illustrated the fourth-instar larva of this species.

Etymology

Vansomerenis is named in honour of Mrs E. C. C. van Someren in recognition of her many important contributions to the taxonomy of Afrotropical mosquitoes, especially those in tribe Aedini. The generic name is feminine, formed from her surname and the feminine Greek suffix ‘-is denoting ‘daughter of’, i.e. daughter of van Someren. Recommended abbreviation = Va.

APPENDIX 4

Checklist and conspectus of reclassification of aedine taxa

‘Finlaya’ and certain allied taxa were the focus of the present study, which resulted in the recognition of 17 genera, of which 11 are new to science, three elevated from subgeneric status and three restored from synonymy. As in our previous study (Reinert et al., 2004), some species of ‘Finlaya’ and their allies cannot be assigned to the new genera with confidence because of a lack of specimens for examination and/or inadequate published descriptions. Consequently, these species (followed by an asterisk in the list below) are only provisionally placed in the newly proposed genera. As previously, where it is not possible to assign species to a genus formally recognized herein or by Reinert et al. (2004) they are retained for the time being in the old broad concept of the genus-level taxon in which they were previously placed. For example, unassigned species previously placed in Ochlerotatus (Ochlerotatus) are listed under this genus and subgenus of previous authors (sensu auctorum). The following checklist and conspectus includes all aedine species and subspecies recognized as valid at the end of March 2005. Where necessary, the terminations of species names have been changed to agree in gender with generic names.

Aedini sensu Belkin, 1962

Abraedes Zavortink, 1970

papago (Zavortink, 1970)

Aedes Meigen, 1818

cinereus Meigen, 1818

dahuricus Danilov, 1987

esoensis Yamada, 1921

geminus Peus, 1970

mubiensis Luh & Shih, 1958

rossicus Dolbeskin, Gorickaja & Mitrofanova, 1930

sasai Tanaka, Mizusawa & Saugstad, 1975

yamadai Sasa, Kano & Takahasi, 1950

‘Aedes’sensu auctorum

Subgenus Aedimorphus Theobald, 1903 sensu auctorum

abnormalis (Theobald, 1909)

ssp. kabwachensis Edwards, 1941

adami Geoffroy, 1971

aerarius McIntosh, 1975

albocephalus (Theobald, 1903)

albodorsalis Fontenille & Brunhes, 1985

alboscutellatus (Theobald, 1905)

alboventralis (Theobald, 1910)

apicoannulatus (Edwards, 1912)

argenteopunctatus (Theobald, 1901)

argenteoscutellatus Carter & Wijesundara, 1948

bambiotai Geoffroy, 1987

bancoi Geoffroy, 1987

bedfordi Edwards, 1936

bevisi (Edwards, 1915)

boneti Gil Collado, 1936

ssp. kumbae Chwatt, 1948

caecus (Theobald, 1901)

caliginosus (Graham, 1910)

centropunctatus (Theobald, 1913)

chamboni Cornet, 1968

culicinus Edwards, 1922

cumminsii (Theobald, 1903)

issp. mediopunctata (Theobald, 1909)

dalzieli (Theobald, 1910)

davidi Basio, 1971

dentatus (Theobald, 1904)

dialloi Hamon & Brengues, 1965

domesticus (Theobald, 1901)

durbanensis (Theobald, 1903)

ssp. angolae Ribeiro & Ramos, 1974

ebogoensis Rickenbach & Ferrara, 1965

eritreae Lewis, 1942

falabreguesi Hamon, 1957

filicis Ingram & de Meillon, 1927

fowleri (Charmoy, 1908)

gandarai da Cunha Ramos, Capela & Ribeiro, 1995

gibbinsi Edwards, 1935

gouldi Reinert, 1972

grenieri Hamon, Service, Adam & Taufflieb, 1961

grjebinei Hamon, Taufflieb & Maillot, 1957

hirsutus (Theobald, 1901)

ssp. adenensis Edwards, 1941

holocinctus Edwards, 1941

hopkinsi Edwards, 1936

insolens Edwards, 1936

irritans (Theobald, 1901)

jamesi (Edwards, 1914)

karooensis Muspratt, 1961

lamborni Edwards, 1923

leesoni Edwards, 1932

leptolabis Edwards, 1936

leucarthrius (Speiser, 1909)

lokojoensis Service, 1959

longiseta Edwards, 1936

lottei Hamon & Brengues, 1965

lowisii (Theobald, 1910)

mansouri Qutubuddin, 1959

masoalensis Fontenille & Brunhes, 1985

mathioti Fontenille & Brunhes, 1985

mattinglyi Hamon & Rickenbach, 1954

mediolineatus (Theobald, 1901)

microstictus Edwards, 1936

minutus (Theobald, 1901)

mixtus Edwards, 1936

mutilus Edwards, 1936

natronius Edwards, 1932

nigricephalus (Theobald, 1901)

nigrostriatus (Barraud, 1927)

nyounae Hamon & Adam, 1959

oakleyi Stone, 1939

ochraceus (Theobald, 1901)

orbitae Edwards, 1922

ovazzai Hamon & Adam, 1959

pachyurus Edwards, 1936

pallidostriatus (Theobald, 1907)

pampangensis (Ludlow, 1905)

phyllolabis Edwards, 1929

pipersalatus (Giles, 1902)

pseudotarsalis van Someren, 1946

pubescens Edwards, 1925

punctifemoris (Ludlow, 1921)

punctothoracis (Theobald, 1909)

quasiunivittatus (Theobald, 1901)

reali Hamon & Adam, 1959

rickenbachi Hamon & Adam, 1959

semlikiensis van Someren, 1950

senyavinensis Knight & Hurlbut, 1949

seychellensis (Theobald, 1912)

simulans (Newstead & Carter, 1911)

smithburni van Someren, 1950

stenoetrus (Theobald, 1907)

stenoscutus Edwards, 1912

subdentatus Edwards, 1936

syntheticus Barraud, 1928

taeniorhynchoides (Christophers, 1911)

tarsalis (Newstead, 1907)

tauffiebi Rickenbach & Ferrara, 1965

tricholabis Edwards, 1941

ssp. bwamba van Someren, 1950

trimaculatus (Theobald, 1905)

trukensis Bohart, 1957

veeniae McIntosh, 1975

vexans (Meigen, 1830)

ssp. arabiensis (Patton, 1905)

ssp. nipponii (Theobald, 1907)

wendyae Service, 1959

wigglesworthi Edwards, 1941

yangambiensis de Meillon & Lavoipierre, 1944

yvonneae Edwards, 1941

Subgenus Cancraedes Edwards, 1929 sensu auctorum

cancricomes Edwards, 1922

curtipes Edwards, 1915

indonesiae Mattingly, 1958

kohkutensis Mattingly, 1958

mamoedjoensis Mattingly, 1958

masculinus Mattingly, 1958

palawanicus Mattingly, 1958

penghuensis Lien, 1968

simplex (Theobald, 1903)

thurmanae Mattingly, 1958

Alanstonea Mattingly, 1960

brevitibia (Edwards, 1914)

treubi (de Meijere, 1910)

Albuginosus Reinert, 1987

capensis (Edwards, 1924)

gilliesi (van Someren, 1962)

haworthi (Edwards, 1923)

kapretwae (Edwards, 1941)

kennethi (Muspratt, 1956)

marshallii (Theobald, 1901)

ngong (van Someren, 1950)

stokesi (Evans, 1929)

teesdalei (van Someren, 1954)

Armigeres Theobald, 1901

Subgenus Armigeres

alkatirii Toma, Miyagi & Syafruddin, 1995

apoensis Bohart & Farner, 1944

aureolineatus (Leicester, 1908)

azurini Basio, 1971

baisasi Stone & Thurman, 1958

bhayungi Thurman & Thurman, 1958

breinli (Taylor, 1914)

candelabrifer Brug, 1939

confusus Edwards, 1915

conjungens Edwards, 1914

denbesteni Brug, 1925

durhami (Edwards, 1917)

ejercitoi Baisas, 1935

fimbriatus Edwards, 1930

foliatus Brug, 1931

giveni Edwards, 1926

hybridus Edwards, 1914

joloensis (Ludlow, 1904)

jugraensis (Leicester, 1908)

kesseli Ramalingam, 1987

kinabaluensis Ramalingam, 1972

kuchingensis Edwards, 1915

lacuum Edwards, 1922

laoensis Toma & Miyagi, 2003

maiae (Edwards, 1917)

malayi (Theobald, 1901)

manalangi Baisas, 1935

maximus Edwards, 1922

milnensis Lee, 1944

moultoni Edwards, 1914

obturbans (Walker, 1859)

pallithorax Dong, Zhou & Dong, 2004

papuensis Peters, 1963

setifer Delfinado, 1966

seticoxitus Luh & Li, 1981

subalbatus (Coquillett, 1898)

var. chrysocorporis Hsieh & Liao, 1956

theobaldi Barraud, 1934

yunnanensis Dong, Zhou & Dong, 1995

Subgenus Leicesteria Theobald, 1904

annulipalpis (Theobald, 1910)

annulitarsis (Leicester, 1908)

balteatus Macdonald, 1960

cingulatus (Leicester, 1908)

dentatus Barraud, 1927

digitatus (Edwards, 1914)

dolichocephalus (Leicester, 1908)

flavus (Leicester, 1908)

inchoatus Barraud, 1927

lepidocoxitus Dong, Zhou & Dong, 1995

longipalpis (Leicester, 1904)

magnus (Theobald, 1908)

menglaensis Dong, Zhou & Dong, 2002

omissus (Edwards, 1914)

pectinatus (Edwards, 1914)

pendulus Edwards, 1914

traubi Macdonald, 1960

vimoli Thurman & Thurman, 1958

Nomina dubia

striocrurus (Giles, 1904)

ventralis (Walker, 1860)

Ayurakitia Thurman, 1954

griffithi Thurman, 1954

peytoni (Reinert, 1972)

Aztecaedes Zavortink, 1972

ramirezi (Vargas & Downs, 1950)

Belkinius Reinert, 1982

aurotaeniatus (Edwards, 1922)

Bothaella Reinert, 1973

alongi (Galliard & Ngu, 1947), comb. nov.

brownscutuma (Dong, Zhu & Dong, 1999)

eldridgei (Reinert, 1973)

helenae (Reinert, 1973)

kleini (Reinert, 1973)

Bruceharrisonius Reinert, 2003, stat. nov., raised to genus rank

alektorovi (Stackelberg, 1943), comb. nov.

aureostriatus (Doleschall, 1857), comb. nov.

christophersi (Edwards, 1922), comb. nov.

doonii (Wattal, Bhatia & Kalra, 1958), comb. nov.

greenii (Theobald, 1903), comb. nov.

hurlbuti (Lien, 1967), comb. nov.

okinawanus (Bohart, 1946), comb. nov.

taiwanus (Lien, 1968), comb. nov.

Christophersiomyia Barraud, 1923

annulirostris (Theobald, 1905)

chionodes (Belkin, 1962)

gombakensis (Mattingly, 1959)

ibis (Barraud, 1931)

thomsoni (Theobald, 1905)

Collessius Reinert, Harbach & Kitching, gen. nov.

banksi (Edwards, 1922), comb. nov.

elsiae (Barraud, 1923), comb. nov.

ssp. vicarius (Lien, 1968), comb. nov.

hatorii (Yamada, 1921), comb. nov.

macdougalli (Edwards, 1922), comb. nov.

macfarlanei (Edwards, 1914), comb. nov.

ningheensis (Lei, 1989), comb. nov.

pseudotaeniatus (Giles, 1901), comb. nov.

shortti (Barraud, 1923), comb. nov.

tonkinensis (Galliard & Ngu, 1947), comb. nov.

Dahliana Reinert, Harbach & Kitching, gen. nov.

echinus (Edwards, 1920), comb. nov.

geniculata (Olivier, 1791), comb. nov.

gilcolladoi (Sanchez-Covisa Villa, Rodriguez Rodriguez & Guillen Llera, 1985), comb. nov.

Diceromyia Theobald, 1911

adersi (Edwards, 1917)

agastyai (Tewari & Hiriyan, 1992)

bananea (Wolfs, 1958)

cordellieri (Huang, 1986)

coulangesi (Rodhain & Boutonnier, 1983)

fascipalpis (Edwards, 1912)

flavicollis (Edwards, 1928)

franciscoi (Mattingly, 1959)

furcifer (Edwards, 1913)

grassei (Doucet, 1951)

iyengari (Edwards, 1923)

kanarensis (Edwards, 1934)

madagascarensis (van Someren, 1949)

mefouensis (Ferrara, 1974)

meronephada (Dyar & Shannon, 1925)

micropterus (Giles, 1901)

nummata (Edwards, 1923)

periskelata (Giles, 1902)

pseudonummata (Reinert, 1973)

punctipes (Edwards, 1921)

ramachandrai (Reuben, 1967)

reginae (Edwards, 1922)

reubenae (Tewari & Hiriyan, 1992)

scanloni (Reinert, 1970)

sylvatica (Brunhes, 1983)

taylori (Edwards, 1936)

tiptoni (Grjebine, 1953)

whartoni (Mattingly, 1965)

zethus (de Meillon & Lavoipierre, 1944)

Dobrotworskyius Reinert, Harbach & Kitching, gen. nov.

alboannulatus (Macquart, 1850), comb. nov.

milsoni (Taylor, 1915), comb. nov.

occidentalis (Skuse, 1889), comb. nov.

rubrithorax (Macquart, 1850), comb. nov.

rupestris (Dobrotworsky, 1959), comb. nov.

subbasalis (Dobrotworsky, 1962), comb. nov.

tubbutiensis (Dobrotworsky, 1959), comb. nov.

Downsiomyia Vargas, 1950

albolateralis (Theobald, 1908)

albonivea (Barraud, 1934)

axitiosa (Kulasekera, Knight & Harbach, 1990)

dorseyi (Knight, 1946)

ganapathi (Colless, 1958)

harinasutai (Knight, 1978)

idjenensis (Brug, 1934)

inermis (Colless, 1958)

lactea (Knight, 1946)

laoagensis (Knight, 1946)

leonis (Colless, 1958)

litorea (Colless, 1958)

mikrokopion (Knight & Harrison, 1988)

mjobergi (Edwards, 1926), comb. nov.

mohani (Knight, 1969)

nipponica (LaCasse & Yamaguti, 1948)

nipoononivea (Sasa & Nakahashi, 1952)

nishikawai (Tanaka, Mizusawa & Saugstad, 1979)

nivea (Ludlow, 1903)

niveoides (Barraud, 1934)

novonivea (Barraud, 1934)

omorii (Lien, 1968)

pexa (Colless, 1958)

pseudonivea (Theobald, 1905)

saperoi (Knight, 1946)

shehzadae (Qutubuddin, 1972), comb. nov.

sinensis (Chow, 1950)

subnivea (Edwards, 1922)

vana (Colless, 1958)

watteni (Lien, 1968)

Edwardsaedes Belkin, 1962

bekkui (Mogi, 1977)

imprimens (Walker, 1860)

pingpaensis (Chang, 1965)

Eretmapodites Theobald, 1901

adami Ferrara & Eouzan, 1974

angolensis da Cunha Ramos & Ribeiro, 1992

argyrurus Edwards, 1936

brenguesi Rickenbach & Lombrici, 1975

brottesi Rickenbach, 1967

caillardi Rickenbach, Ferrara & Eouzan, 1968

chrysogaster Graham, 1909

corbeti Hamon, 1962

dracaenae Edwards, 1916

dundo da Cunha Ramos & Ribeiro, 1992

eouzani Rickenbach & Lombrici, 1974

ferrarai Rickenbach & Eouzan, 1970

forcipulatus Edwards, 1936

germaini Rickenbach & Eouzan, 1970

gilletti van Someren, 1949

grahami Edwards, 1911

grenieri Hamon & van Someren, 1961

haddowi van Someren, 1949

hamoni Grjebine, 1972

harperi van Someren, 1949

hightoni van Someren, 1947

inornatus Newstead, 1907

intermedius Edwards, 1936

jani Rickenbach & Lombrici, 1976

lacani Rickenbach & Eouzan, 1970

leucopous Graham, 1909

mahaffyi van Someren, 1949

marcellei Adam & Hamon, 1959

mattinglyi Hamon & van Someren, 1961

melanopous Graham, 1909

mortiauxi Cunha Ramos & Ribeiro, 1990

oedipodeios Graham, 1909

parvipluma Edwards, 1941

pauliani Grjebine, 1950

penicillatus Edwards, 1941

plioleucus Edwards, 1941

ssp. brevis Edwards, 1941

productus Edwards, 1941

quinquevittatus Theobald, 1901

ravissei Rickenbach & Eouzan, 1970

rickenbachi Ferrara & Eouzan, 1974

salauni Rickenbach, Ferrara & Eouzan, 1968

semisimplicipes Edwards, 1914

silvestris Ingram & de Meillon, 1927

ssp. conchobius Edwards, 1941

subsimplicipes Edwards, 1914

tendeiroi da Cunha Ramos, Ribeiro & Machado, 1992

tonsus Edwards, 1941

vansomereni Hamon, 1962

wansoni Edwards, 1941

ssp. douceti Adam & Hamon, 1959

Finlaya Theobald, 1903

alocasicola (Marks, 1947)

ananae (Knight & Laffoon, 1946)

avistylus (Brug, 1939)

bougainvillensis (Marks, 1947)

burnetti (Belkin, 1962)

crocea (Knight & Laffoon, 1946)

dobrotworskyi (Marks, 1958)

fijiensis (Marks, 1947)

flavipennis Giles, 1904

franclemonti (Belkin, 1962)

freycinetiae (Laird, 1957)

fuscipalpis (Belkin, 1962)

fuscitarsis (Belkin, 1962)

gahnicola (Marks, 1947)

gani (Bonne-Wepster, 1940)*

gressitti (Bohart, 1947)

hollingsheadi (Belkin, 1962)

horotoi (Taylor, 1972)

hui (Bohart, 1957)

josephinae (Marks, 1958)

knighti (Stone & Bohart, 1944)

kochi (Dönitz, 1901)

lewelleni (Starkey & Webb, 1946)

lutea (Ludlow, 1905)

maffii (Taylor & Tenorio, 1974)

medleri (Knight & Laffoon, 1946)

neogeorgiana (Belkin, 1962)

oceanica (Belkin, 1962)

poicilia Theobald, 1903

samoana Grünberg, 1913

schlosseri (Belkin, 1962)

solomonis (Stone & Bohart, 1944)

sorsogonensis (Banez & Jueco, 1966)

stonei (Knight & Laffoon, 1946)

tutuilae (Ramalingam & Belkin, 1965)

wallacei (Edwards, 1926)

Fredwardsius Reinert, 2000

vittatus (Bigot, 1861)

Georgecraigius Reinert, Harbach & Kitching, gen. nov.

Subgenus Georgecraigius

atropalpus (Coquillett, 1902), comb. nov.

epactius (Dyar & Knab, 1908), comb. nov.

Subgenus Horsfallius Reinert, Harbach & Kitching, subgen. nov.

fluviatilis (Lutz, 1904), comb. nov.

Geoskusea Edwards, 1929

baisasi (Knight & Hull, 1951)

becki (Belkin, 1962)

daggyi (Stone & Bohart, 1944)

fimbripes (Edwards, 1924)

kabaenensis (Brug, 1939)

longiforceps (Edwards, 1929)

lunulata (King & Hoogstraal, 1946)

perryi (Belkin, 1962)

timorensis (Miyagi, Toma & Lien, 2004), comb. nov.

tonsa (Edwards, 1924)

Gilesius Reinert, Harbach & Kitching, gen. nov.

alius (Lien, 1968), comb. nov.

pulchriventer (Giles, 1901), comb. nov.

Gymnometopa Coquillett, 1905

mediovittata (Coquillett, 1906)

Haemagogus Williston, 1896

Subgenus Conopostegus Dyar, 1925

clarki (Galindo, Carpenter & Trapido, 1953)

leucocelaenus (Dyar & Shannon, 1924)

leucophoebus (Galindo, Carpenter & Trapido, 1953)

leucotaeniatus (Komp, 1938)

Subgenus Haemagogus

acutisentis Arnell, 1973

aeritinctus Galindo & Trapido, 1967

albomaculatus Theobald, 1903

anastasionis Dyar, 1921

andinus Osorno-Mesa, 1944

argyromeris Dyar & Ludlow, 1921

baresi Cerqueira, 1960

boshelli Osorno-Mesa, 1944

capricornii Lutz, 1904

celeste Dyar & Nunez Tovar, 1927

chalcospilans Dyar, 1921

chrysochlorus Arnell, 1973

equinus Theobald, 1903

iridicolor Dyar, 1921

janthinomys Dyar, 1921

lucifer (Howard, Dyar & Knab, 1913)

mesodentatus Komp & Kumm, 1938

nebulosus Arnell, 1973

panarchys Dyar, 1921

regalis Dyar & Knab, 1906

soperi Levi-Castillo, 1955

spegazzinii Brethes, 1912

splendens Williston, 1896

tropicalis Cerqueira & Antunes, 1938

Halaedes Belkin, 1962

ashworthi (Edwards, 1921)

australis (Erichson, 1842)

wardangensis (Brust, Ballard, Driver, Hartley, Galway & Curran, 1998)

Heizmannia Ludlow, 1905

Subgenus Heizmannia

aurea Brug, 1932

aureochaeta (Leicester, 1908)

carteri Amerasinghe, 1993

chandi Edwards, 1922

chengi Lien, 1968

communis (Leicester, 1908)

complex (Theobald, 1910)

covelli Barraud, 1929

demeilloni Mattingly, 1970

funerea (Leicester, 1908)

greenii (Theobald, 1905)

heterospina Gong & Lu, 1986

himalayensis Edwards, 1922

indica (Theobald, 1905)

kana Tanaka, Mizusawa & Saugstad, 1979

kanhsienensis Tung, 1955

lii Wu, 1936

macdonaldi Mattingly, 1957

mattinglyi Thurman, 1959

menglianensis Lu & Gong, 1986

menglianeroides Dong, Dong & Zhou, 2003

persimilis Mattingly, 1970

propinqua Mattingly, 1970

proxima Mattingly, 1970

reidi Mattingly, 1957

ruiliensis Dong, Zhou & Wang, 1997

scanloni Mattingly, 1970

scintillans Ludlow, 1905

taiwanensis Lien, 1968

tengchongensis Dong, Wang & Zhou, 2002

thelmae Mattingly, 1970

viridis Barraud, 1929

Subgenus Mattinglyia Lien, 1968

achaetae (Leicester, 1908)

catesi (Lien, 1968)

discrepans (Edwards, 1922)

occidentayunnana (Gong & Lu, 1991)

tripunctata (Theobald, 1908)

Himalaius Reinert, Harbach & Kitching, gen. nov.

gilli (Barraud, 1924), comb. nov.

simlensis (Edwards, 1922)*, comb. nov.

Howardina Theobald, 1903

albonotata (Coquillett, 1906)

allotecnon (Kumm, Komp & Ruiz, 1940)

arborealis (Bonne-Wepster & Bonne, 1920)

argyriti (Dyar & Nunez Tovar, 1927)

aureolineata (Berlin, 1969)

auriti Theobald, 1907

aurivittata (Cerqueira, 1943)

bahamensis (Berlin, 1969)

brevis (Berlin, 1969)

brevivittata (Berlin, 1969)

busckii (Coquillett, 1906)

cozumelensis (Diaz Najera, 1966)

ecuadoriensis (Berlin, 1969)

eleanorae (Berlin, 1969)

fulvithorax (Lutz, 1904)

grabhami (Berlin, 1969)

guatemala (Berlin, 1969)

guerrero (Berlin, 1969)

inaequalis Grabham, 1907

ioliota (Dyar & Knab, 1913)

leei (Berlin, 1969)

lorraineae (Berlin, 1969)

marinkellei (Berlin, 1969)

martinezi (Berlin, 1969)

osornoi (Berlin, 1969)

pseudodminicii (Komp, 1936)

quadrivittata (Coquillett, 1902)

septemstriata (Dyar & Knab, 1907)

sexlineata (Theobald, 1901)

spinosa (Berlin, 1969)

stenei (Thompson, 1956)

vanemdeni (Martini, 1931)

walkeri (Theobald, 1901)

whitmorei (Dunn, 1918)

Huaedes Huang, 1968

medialis (Brug, 1932)

variepictus (King & Hoogstraal, 1946)

wauensis (Huang, 1968)

Hulecoeteomyia Theobald, 1904, stat. nov., restored to genus rank

chrysolineata (Theobald, 1907), comb. nov.

formosensis (Yamada, 1921), comb. nov.

harveyi (Barraud, 1923), comb. nov.

japonica (Theobald, 1901), comb. nov.

ssp. shintienensis (Tsai & Lien, 1950), comb. nov.

ssp. amamiensis (Tanaka, Mizusawa & Saugstad, 1979), comb. nov.

ssp. yaeyamensis (Tanaka, Mizusawa & Saugstad, 1979), comb. nov.

jugraensis Leicester, 1908, comb. nov.

koreica (Edwards, 1917), comb. nov.

nigrorhynchus (Brug, 1931), comb. nov.

pallirostris (Edwards, 1922), comb. nov.

reinerti (Rattanarithikul & Harrison, 1988), comb. nov.

rizali (Banks, 1906), comb. nov.

saxicola (Edwards, 1922), comb. nov.

sherki (Knight, 1948), comb. nov.

yunnanensis (Gaschen, 1934)*, comb. nov.

Indusius Edwards, 1934

pulverulentus (Edwards, 1922)

Isoaedes Reinert, 1979

cavaticus (Reinert, 1979)

Jarnellius Reinert, Harbach & Kitching, gen. nov.

Subgenus Jarnellius

deserticola (Zavortink, 1969), comb. nov.

laguna (Arnell & Nielsen, 1972), comb. nov.

monticola (Belkin & McDonald, 1957), comb. nov.

sierrensis (Ludlow, 1905), comb. nov.

varipalpus (Coquillett, 1902), comb. nov.

Subgenus Lewnielsenius Reinert, Harbach & Kitching, subgen. nov.

muelleri (Dyar, 1920), comb. nov.

Jihlienius Reinert, Harbach & Kitching, gen. nov.

chungi (Lien, 1968), comb. nov.

gonguoensis (Gong & Lu, 1986), comb. nov.

unicinctus (Edwards, 1922), comb. nov.

Kenknightia Reinert, 1990

dissimilierodes (Dong, Zhou & Dong, 2002)

dissimilis (Leicester, 1908)

gaffigani (Reinert, 1990)

harbachi (Reinert, 1990)

karwari (Barraud, 1924)

lerozeboomi (Reinert, 1990)

leucomeres (Giles, 1904)

litwakae (Reinert, 1990)

luzonensis (Rozeboom, 1946)

paradissimilis (Rozeboom, 1946)

pecori (Reinert, 1990)

wilkersoni (Reinert, 1990)

Kompia Aitken, 1941

purpureipes (Aitken, 1941)

Leptosomatomyia Theobald, 1905

aurimargo (Edwards, 1922)

Levua Stone & Bohart, 1944

dufouri (Hamon, 1953)

fryeri (Theobald, 1912)

geoskusea (Amos, 1944)

Lorrainea Belkin, 1962

amesii (Ludlow, 1903)

celebica (Mattingly, 1959)

dasyorrhus (King & Hoogstraal, 1946)

fumida (Edwards, 1928)

lamellifera (Bohart & Ingram, 1946)

Macleaya Theobald, 1903, stat. nov., restored to genus rank

Subgenus Chaetocruiomyia Theobald, 1910, transferred from Ochlerotatus

calabyi (Marks, 1963), comb. nov.

elchoensis (Taylor, 1929), comb. nov.

humeralis (Edwards, 1922), comb. nov.

macmillani (Marks, 1964), comb. nov.

moloiensis (Taylor, 1929), comb. nov.

spinosipes Edwards, 1922, comb. nov.

tulliae (Taylor, 1929), comb. nov.

wattensis (Taylor, 1929), comb. nov.

Subgenus Macleaya, transferred from Ochlerotatus

littlechildi (Taylor, 1933), comb. nov.

stoneorum (Marks, 1977), comb. nov.

tremula Theobald, 1903, comb. nov.

Molpemyia Theobald, 1910, stat. nov., restored to genus rank

auridorsum (Edwards, 1922), comb. nov.

pecuniosa (Edwards, 1922), comb. nov.

purpurea Theobald, 1910, comb. nov.

Mucidus Theobald, 1901

alternans (Westwood, 1835)

aurantius (Theobald, 1907)

ssp. chrysogaster Taylor, 1927

ferinus (Knight, 1947)

grahamii Theobald, 1909

laniger (Wiedemann, 1820)

lucianus (Muspratt, 1959)

mucidus (Karsch, 1887)

nigerrimus Theobald, 1913

painei (Knight, 1948)

quadripunctis (Ludlow, 1910)

quasiferinus (Mattingly, 1961)

scatophagoides Theobald, 1901

sudanensis Theobald, 1908

tonkingi (Gebert, 1948)

Neomelaniconion Newstead, 1907

albicosta (Edwards, 1913)

albothorax (Theobald, 1907)

aurovenatum (Worth, 1960)

bequaerti (Wolfs, 1947)

bergerardi (Pajot & Geoffroy, 1971)

bolense (Edwards, 1936)

carteri (Edwards, 1936)

circumluteolum (Theobald, 1908)

crassiforceps (Edwards, 1927)

ellinorae (Edwards, 1941)

flavimargo (Edwards, 1941)

fuscinerve (Edwards, 1914)

jamoti (Hamon & Rickenbach, 1954)

lineatopenne (Ludlow, 1905)

ssp. aureum (Gutsevich, 1955)

luridum (McIntosh, 1971)

luteolaterale (Theobald, 1901)

mcintoshi (Huang, 1985)

monotrichus (Edwards, 1936)

palpale Newstead, 1907

pogonurum (Edwards, 1936)

punctocostale (Theobald, 1909)

taeniarostre (Theobald, 1909)

unidentatum (McIntosh, 1971)

Ochlerotatus Lynch Arribalzaga, 1891

Subgenus Rusticoidus Shevchenko & Prudkina, 1973

albescens (Edwards, 1921)

bicristatus (Thurman & Winkler, 1950)

krymmontanus (Alekseev, 1989)

lepidonotus (Edwards, 1920)

provocans (Walker, 1848)

quasirusticus (Torres Canamares, 1951)

refiki (Medschid, 1928)

rusticus (Rossi, 1790)

var. subtrichurus (Martini, 1927)

subdiversus (Martini, 1926)

Unassigned to subgenus

atlanticus (Dyar & Knab, 1906)

communis (de Geer, 1776)

dorsalis (Meigen, 1830)

fulvus (Wiedemann, 1828)

ssp. pallens (Ross, 1943)

infirmatus (Dyar & Knab, 1906)

scapularis (Rondani, 1848)

sollicitans (Walker, 1856)

Ochlerotatussensu auctorum

Subgenus ‘Finlaya’ Theobald, 1903 sensu auctorum

albocinctus (Barraud, 1924)

albotaeniata (Leicester, 1904)

alticola (Bonne-Wepster, 1948)

auronitens (Edwards, 1922)

australiensis (Theobald, 1910)

barnardi (Edwards, 1924)

biocellatus (Taylor, 1915)

britteni (Marks & Hodgkin, 1958)

candidoscutellum (Marks, 1947)

crossi (Lien, 1967)

eatoni Edwards, 1916

embuensis (Edwards, 1930)

fengi (Edwards, 1935)

gracilelineatus (Bonne-Wepster, 1937)

harperi (Knight, 1948)

ingrami (Edwards, 1930)

keefei (King & Hoogstraal, 1946)

koreicoides (Sasa, Kano & Hayashi, 1950)

lauriei Carter, 1920

lepchana (Barraud, 1923)

loi (Lien, 1968)

mallochi (Taylor, 1944)

monocellatus (Marks, 1948)

nyasae (Edwards, 1930)

oreophilus Edwards, 1916

peipingensis (Feng, 1938)

plagosus (Marks, 1959)

quasirubithorax (Theobald, 1910)

roai (Belkin, 1962)

rubiginosus (Belkin, 1962)

seoulensis (Yamada, 1921)

sintoni (Barraud, 1924)

stanleyi (Peters, 1963)

subauridorsum (Marks, 1948)

suffusus (Edwards, 1922)

toxopeusi (Bonne-Wepster, 1948)

tsiliensis (King & Hoogstraal, 1946)

versicolor (Barraud, 1924)

wasselli (Marks, 1947)

wellmanii (Theobald, 1905)

Subgenus Ochlerotatus Lynch Arribalzaga, 1891 sensu auctorum

aboriginis (Dyar, 1917)

abserratus (Felt & Young, 1904)

aculeatus (Theobald, 1903)

aenigmaticus (Cerqueira & Costa, 1946)

akkeshiensis (Tanaka, 1998)

albifasciatus (Macquart, 1838)

albineus (Seguy, 1923)

aloponotum (Dyar, 1917)

ambreensis (Rodhain & Boutonnier, 1983)

andersoni (Edwards, 1926)

angustivittatus (Dyar & Knab, 1907)

annulipes (Meigen, 1830)

antipodeus Edwards, 1920

atactavittatus (Arnell, 1976)

auratus (Grabham, 1906)

aurifer (Coquillett, 1903)

behningi (Martini, 1926)

bejaranoi (Martinez, Carcavallo & Prosen, 1960)

berlandi (Seguy, 1921)

bimaculatus (Coquillett, 1902)

biskraensis (Brunes, 1999)

bogotanus (Arnell, 1976)

breedensis (Muspratt, 1953)

burjaticus (Kuchartshuk, 1973)

burpengaryensis (Theobald, 1905)

caballus (Theobald, 1912)

cacozelus (Marks, 1963)

calcariae (Marks, 1957)

calumnior (Belkin, Heinemann & Page, 1970)

campestris (Dyar & Knab, 1907)

camptorhynchus (Thomson, 1869)

canadensis (Theobald, 1901)

ssp. mathesoni (Middlekauff, 1944)

cantans (Meigen, 1818)

var. subvexans (Martini, 1922)

cantator (Coquillett, 1903)

caspius (Pallas, 1771)

ssp. meirai Ribeiro, da Cunha Ramos, Capela & Pires, 1980

var. hargreavesi Edwards, 1920

cataphylla (Dyar, 1916)

var. rostochiensis (Martini, 1920)

chelli Edwards, 1915

churchillensis (Ellis & Brust, 1973)

clelandi (Taylor, 1914)

clivis (Lanzaro & Eldridge, 1992)

coluzzii (Rioux, Guilvard & Pasteur, 1998)

comitatus (Arnell, 1976)

condolescens (Dyar & Knab, 1907)

continentalis (Dobrotworsky, 1960)

crinifer (Theobald, 1903)

cunabulanus (Edwards, 1924)

cyprioides (Danilov & Stupin, 1982)

cyprius (Ludlow, 1920)

decticus (Howard, Dyar & Knab, 1917)

deficiens (Arnell, 1976)

detritus (Haliday, 1833)

diantaeus (Howard, Dyar & Knab, 1913)

duplex (Martini, 1926)

dupreei (Coquillett, 1904)

dzeta (Seguy, 1924)

edgari (Stone & Rosen, 1952)

eidsvoldensis (Mackerras, 1927)

eucephalaeus (Dyar, 1918)

euedes (Howard, Dyar & Knab, 1913)

euiris (Dyar, 1922)

euplocamus (Dyar & Knab, 1906)

excrucians (Walker, 1856)

explorator (Marks, 1964)

fitchii (Felt & Young, 1904)

flavescens (Müller, 1764)

flavifrons (Skuse, 1889)

grossbecki (Dyar & Knab, 1906)

gutzevichi (Dubitsky & Deshevykh, 1978)

hakusanensis (Yamaguti & Tamaboko, 1954)

harrisoni (Muspratt, 1953)

hastatus (Dyar, 1922)

hesperonotius (Marks, 1959)

hexodontus (Dyar, 1916)

hodgkini (Marks, 1959)

hokkaidensis (Tanaka, Mizusawa & Saugstad, 1979)

hortator (Dyar & Knab, 1907)

hungaricus (Mihalyi, 1955)

imperfectus (Dobrotworsky, 1962)

impiger (Walker, 1848)

ssp. daisetsuzanus (Tanaka, Mizusawa & Saugstad, 1979)

implicatus (Vockeroth, 1954)

incomptus (Arnell, 1976)

increpitus (Dyar, 1916)

inexpectatus (Bonne-Wepster, 1948)

intermedius (Danilov & Gornostaeva, 1987)

intrudens (Dyar, 1919)

jacobinae (Serafim & Davis, 1933)

jorgi (Carpintero & Leguizamón, 2000)

juppi (McIntosh, 1973)

kasachstanicus (Gutsevich, 1962)

lasaensis (Meng, 1962)

ssp. gyirongensis (Ma, 1982)

lepidus (Cerqueira & Paraense, 1945)

leucomelas (Meigen, 1804)

linesi (Marks, 1964)

longifilamentus (Su & Zhang, 1988)

luteifemur (Edwards, 1926)

macintoshi (Marks, 1959)

mariae (Sergent & Sergent, 1903)

martineti (Senevet, 1937)

mcdonaldi (Belkin, 1962)

melanimon (Dyar, 1924)

meprai (Martinez & Prosen, 1953)

mercurator (Dyar, 1920)

milleri (Dyar, 1922)

mitchellae (Dyar, 1905)

montchadskyi (Dubitsky, 1968)

nevadensis (Chapman & Barr, 1964)

nigrinus (Eckstein, 1918)

nigripes (Zetterstedt, 1838)

nigrithorax (Macquart, 1847)

nigrocanus (Martini, 1927)

nigromaculis (Ludlow, 1906)

niphadopsis (Dyar & Knab, 1918)

nivalis (Edwards, 1926)

normanensis (Taylor, 1915)

nubilis (Theobald, 1903)

obturbator (Dyar & Knab, 1907)

oligopistus (Dyar, 1918)

patersoni (Shannon & del Ponte, 1928)

pectinatus (Arnell, 1976)

pennai (Antunes & Lane, 1938)

perkinsi (Marks, 1949)

pertinax (Grabham, 1906)

perventor (Cerqueira & Costa, 1946)

phaecasiatus (Marks, 1964)

phaeonotus (Arnell, 1976)

phoeniciae (Coluzzi & Sabatini, 1968)

pionips (Dyar, 1919)

procax Skuse, 1889

pseudonormanensis (Marks, 1949)

pulcritarsis (Rondani, 1872)

ssp. asiaticus (Edwards, 1926)

pullatus (Coquillett, 1904)

punctodes (Dyar, 1922)

punctor (Kirby, 1837)

purpuraceus (Brug, 1932)

purpureifemur (Marks, 1959)

purpuriventris (Edwards, 1926)

ratcliffei (Marks, 1959)

raymondi (del Ponte, Castro & Garcia, 1951)

rempeli (Vockeroth, 1954)

rhyacophilus (da Costa Lima, 1933)

riparioides (Su & Zhang, 1987)

riparius (Dyar & Knab, 1907)

sagax (Skuse, 1889)

sapiens (Marks, 1964)

schizopinax (Dyar, 1929)

schtakelbergi (Shingarev, 1928)

scutellalbum (Boshell-Manrique, 1939)

sedaensis (Lei, 1989)

sergievi (Danilov, Markovich & Proskuryakova, 1978)

serratus (Theobald, 1901)

shannoni (Vargas & Downs, 1950)

silvestris (Dobrotworsky, 1961)

simanini (Gutsevich, 1966)

sinkiangensis (Hsiao, 1977)

spencerii (Theobald, 1901)

ssp. idahoensis (Theobald, 1903)

spilotus (Marks, 1963)

squamiger (Coquillett, 1902)

sticticus (Meigen, 1838)

stigmaticus (Edwards, 1922)

stimulans (Walker, 1848)

stramineus (Dubitzky, 1970)

stricklandi Edwards, 1912

subalbirostris (Klein & Marks, 1960)

surcoufi (Theobald, 1912)

synchytus (Arnell, 1976)

taeniorhynchus (Wiedemann, 1821)

tahoensis (Dyar, 1916)

thelcter (Dyar, 1918)

theobaldi (Taylor, 1914)

thibaulti (Dyar & Knab, 1910)

tormentor (Dyar & Knab, 1906)

tortilis (Theobald, 1903)

trivittatus (Coquillett, 1902)

turneri (Marks, 1963)

upatensis (Anduze & Hecht, 1943)

ventrovittis (Dyar, 1916)

vigilax (Skuse, 1889)

ssp. ludlowae (R. Blanchard, 1905)

ssp. vansomerenae (Mattingly, 1955)

vittiger (Skuse, 1889)

washinoi (Lanzaro & Eldridge, 1992)

zammitii (Theobald, 1903)

Subgenus Protomacleaya Theobald, 1907 sensu auctorum

aitkeni (Schick, 1970)

alboapicus (Schick, 1970)

amabilis (Schick, 1970)

argyrothorax (Bonne-Wepster & Bonne, 1920)

berlini (Schick, 1970)

bertrami (Schick, 1970)

braziliensis (Gordon & Evans, 1922)

brelandi (Zavortink, 1972)

buenaventura (Schick, 1970)

burgeri (Zavortink, 1972)

campana (Schick, 1970)

casali (Schick, 1970)

chionotum (Zavortink, 1972)

daryi (Schick, 1970)

diazi (Schick, 1970)

gabriel (Schick, 1970)

galindoi (Schick, 1970)

hendersoni (Cockerell, 1918)

heteropus (Dyar, 1921)

homoeopus (Dyar, 1922)

idanus (Schick, 1970)

impostor (Schick, 1970)

insolitus (Coquillett, 1906)

knabi (Coquillett, 1906)

kompi (Vargas & Downs, 1950)

metoecopus (Dyar, 1925)

niveoscutum (Zavortink, 1972)

podographicus (Dyar & Knab, 1906)

sandrae (Zavortink, 1972)

schicki (Zavortink, 1972)

schroederi (Schick, 1970)

sumidero (Schick, 1970)

tehuantepec (Schick, 1970)

terrens (Walker, 1856)

thorntoni (Dyar & Knab, 1907)

triseriatus (Say, 1823)

vargasi (Schick, 1970)

zavortinki (Schick, 1970)

zoosophus (Dyar & Knab, 1918)

Subgenus uncertain

daliensis (Taylor, 1916)

Opifex Hutton, 1902

Subgenus Nothoskusea Dumbleton, 1962

chathamicus (Dumbleton, 1962)

Subgenus Opifex

fuscus Hutton, 1902

Paraedes Edwards, 1934

barraudi Edwards, 1934

bonneae (Mattingly, 1958)

chrysoscuta (Theobald, 1910)

collessi (Mattingly, 1958)

menoni (Mattingly, 1958)

ostentatio (Leicester, 1908)

pagei (Ludlow, 1911)

thailandensis (Reinert, 1976)

Patmarksia Reinert, Harbach & Kitching, gen. nov.

anggiensis (Bonne-Wepster, 1937), comb. nov.

argenteitarsis (Brug, 1932), comb. nov.

argyronotum (Belkin, 1962), comb. nov.

buxtoni (Belkin, 1962), comb. nov.

clintoni (Taylor, 1946), comb. nov.

derooki (Brug, 1932), comb. nov.

dobodura (King & Hoogstraal, 1946), comb. nov.

hollandia (King & Hoogstraal, 1946), comb. nov.

mackerrasi (Taylor, 1927), comb. nov.

novalbitarsis (King & Hoogstraal, 1946), comb. nov.

palmarum (Edwards, 1924), comb. nov.

papuensis (Taylor, 1914), comb. nov.

subalbitarsis (King & Hoogstraal, 1946), comb. nov.

Phagomyia Theobald, 1905, stat. nov., restored to genus rank

assamensis (Theobald, 1908), comb. nov.

cacharana (Barraud, 1923), comb. nov.

cogilli (Edwards, 1922), comb. nov.

deccana (Barraud, 1923), comb. nov.

feegradei (Barraud, 1934), comb. nov.

gubernatoris (Giles, 1901), comb. nov.

var. kotiensis (Barraud, 1934), comb. nov.

inquinata (Edwards, 1922)*, comb. nov.

iwi (Marks, 1955)*, comb. nov.

khazani (Edwards, 1922), comb. nov.

kiangsiensis (Tung, 1955)*, comb. nov.

lophoventralis (Theobald, 1910), comb. nov.

melanoptera (Giles, 1904), comb. nov.

plumifera (King & Hoogstraal, 1946), comb. nov.

prominens (Barraud, 1923), comb. nov.

stevensoni (Barraud, 1923), comb. nov.

watasei (Yamada, 1921), comb. nov.

Pseudoarmigeres Stone & Knight, 1956

albomarginatus (Newstead, 1907)

argenteoventralis (Theobald, 1909)

ssp. dunni (Evans, 1928)

kummi (Edwards, 1930)

michaelikati (van Someren, 1946)

ssp. gurneri (van Someren, 1946)

natalensis (Edwards, 1930)

Pseudoskusea Theobald, 1907, stat. nov., restored to genus rank

bancroftiana (Edwards, 1921), comb. nov.

culiciformis (Theobald, 1905), comb. nov.

multiplex (Theobald, 1903), comb. nov.

postspiraculosa (Dobrotworsky, 1961), comb. nov.

Psorophora Robineau-Desvoidy, 1827

Subgenus Grabhamia Theobald, 1903

cingulata (Fabricius, 1805)

columbiae (Dyar & Knab, 1906)

confinnis (Lynch Arribalzaga, 1891)

dimidiata Cerqueira, 1943

discolor (Coquillett, 1903)

infinis (Dyar & Knab, 1906)

insularia (Dyar & Knab, 1906)

jamaicensis Theobald, 1901

leucocnemis Martini, 1931

paulli Paterson & Shannon, 1927

pruinosa Martini, 1935

pygmaea (Theobald, 1903)

santamarinai Broche, 2000

signipennis (Coquillett, 1904)

varinervis Edwards, 1922

Subgenus Janthinosoma Lynch Arribalzaga, 1891

albigenu (Peryassu, 1908)

albipes (Theobald, 1907)

amazonica Cerqueira, 1960

champerico (Dyar & Knab, 1906)

circumflava Cerqueira, 1943

cyanescens (Coquillett, 1902)

discrucians (Walker, 1856)

ferox (von Humboldt, 1819)

fiebrigi Edwards, 1922

forceps Cerqueira, 1939

horrida (Dyar & Knab, 1908)

johnstonii (Grabham, 1905)

lanei Shannon & Cerqueira, 1943

longipalpus Randolph & O'Neill, 1944

lutzii (Theobald, 1901)

mathesoni Belkin & Heinemann, 1975

melanota Cerqueira, 1943

mexicana (Bellardi, 1859)

pilosa Duret, 1971

pseudoalbipes Duret, 1971

pseudomelanota Barata & Cotrim, 1971

totonaci Lassmann, 1951

varipes (Coquillett, 1904)

Subgenus Psorophora

ciliata (Fabricius, 1794)

cilipes (Fabricius, 1805)

holmbergii Lynch Arribalzaga, 1891

howardii Coquillett, 1901

lineata (von Humboldt, 1819)

ochripes (Macquart, 1850)

pallescens Edwards, 1922

pilipes (Macquart, 1834)

saeva Dyar & Knab, 1906

stonei Vargas, 1956

Nomina dubia

goeldii (Giles, 1904)

marmorata (Philippi, 1865)

perterrens (Walker, 1856)

Rampamyia Reinert, Harbach & Kitching, gen. nov.

albilabris (Edwards, 1925), comb. nov.

notoscripta (Skuse, 1889), comb. nov.

ssp. montana (Brug, 1939), comb. nov.

quinquelineata (Edwards, 1922), comb. nov.

Rhinoskusea Edwards, 1929

longirostris (Leicester, 1908)

pillaii (Mattingly, 1958)

portonovoensis (Tewari & Hiriyan, 1992)

wardi (Reinert, 1976)

Scutomyia Theobald, 1904

albolineata Theobald, 1904

arboricola (Knight & Rozeboom, 1946)

bambusicola Knight & Rozeboom, 1946

boharti (Knight & Rozeboom, 1946)

hoogstraali (Knight & Rozeboom, 1946)

impatibilis (Walker, 1859)

laffooni (Knight & Rozeboom, 1946)

platylepida (Knight & Hull, 1951)

pseudalbolineata (Brug, 1939)

Skusea Theobald, 1903

cartroni (Ventrillon, 1906)

lambrechti (van Someren, 1971)

moucheti (Ravaonjanahary & Brunhes, 1977)

pembaensis (Theobald, 1901)

Stegomyia Theobald, 1901

aegypti (Linnaeus, 1762)

ssp. formosus (Walker, 1848)

africana Theobald, 1901

agrihanensis (Bohart, 1957)

albopicta (Skuse, 1895)

alcasidi (Huang, 1972)

alorensis (Bonne-Wepster & Brug, 1932)

amalthea (de Meillon & Lavoipierre, 1944)

andrewsi (Edwards, 1926)

angusta (Edwards, 1935)

annandalei Theobald, 1910

var. quadricincta (Barraud, 1923)

aobae (Belkin, 1962)

apicoargentea Theobald, 1909

bambusae (Edwards, 1935)

blacklocki (Evans, 1925), comb. nov.

bromeliae Theobald, 1911

burnsi (Basio & Reisen, 1971)

calceata (Edwards, 1924)

chaussieri (Edwards, 1923)

chemulpoensis (Yamada, 1921)

contigua (Edwards, 1936)

cooki (Belkin, 1962)

corneti (Huang, 1986)

cozi (Cornet, 1973)

craggi Barraud, 1923

cretina (Edwards, 1921)

daitensis (Miyagi & Toma, 1981)

deboeri (Edwards, 1926)

demeilloni (Edwards, 1936)

denderensis (Wolfs, 1949)

dendrophila (Edwards, 1921)

desmotes Giles, 1904

dybasi (Bohart, 1957)

ealaensis (Huang, 2004), comb. nov.

edwardsi Barraud, 1923

ethiopiensis (Huang, 2004), comb. nov.

flavopicta (Yamada, 1921)

ssp. downsi (Bohart & Ingram, 1946)

ssp. miyarai (Tanaka, Mizusawa & Saugstad, 1979)

fraseri Edwards, 1912

futunae (Belkin, 1962)

galloisi (Yamada, 1921)

galloisiodes (Liu & Lu, 1984)

gandaensis (Huang, 2004), comb. nov.

gardnerii Ludlow, 1905

ssp. imitator Leicester, 1908

grantii Theobald, 1901

guamensis (Farner & Bohart, 1944)

gurneyi (Stone & Bohart, 1944)

hakanssoni (Knight & Hurlbut, 1949)

hansfordi (Huang, 1997)

hebridea (Edwards, 1926)

heischi (van Someren, 1951)

hensilli (Farner, 1945)

hogsbackensis (Huang, 2004), comb. nov.

hoguei (Belkin, 1962)

horrescens (Edwards, 1935)

josiahae (Huang, 1988)

katherinensis (Woodhill, 1949)

keniensis (van Someren, 1946)

kenyae (van Someren, 1946)

kesseli (Huang & Hitchcock, 1980)

kivuensis (Edwards, 1941)

krombeini (Huang, 1975)

lamberti (Ventrillon, 1904)

langata (van Someren, 1946)

ledgeri (Huang, 1981)

lilii Theobald, 1910

luteocephala Newstead, 1907

maehleri (Bohart, 1957)

malayensis (Colless, 1962)

malikuli (Huang, 1973)

marshallensis (Stone & Bohart, 1944)

mascarensis (MacGregor, 1924)

masseyi (Edwards, 1923)

mattinglyorum (Huang, 1994)

maxgermaini (Huang, 1990)

mediopunctata Theobald, 1905

var. sureilensis (Barraud, 1934)

metallica (Edwards, 1912)

mickevichae (Huang, 1988)

mpusiensis (Huang, 2004), comb. nov.

muroafcete (Huang, 1997)

neoafricana (Cornet, Valade & Dieng, 1978)

neogalloisi (Chen & Chen, 2000)

neopandani (Bohart, 1957)

njombiensis (Huang, 1997)

novalbopicta (Barraud, 1931)

opok (Corbet & van Someren, 1962)

palauensis (Bohart, 1957)

pandani (Stone, 1939)

patriciae (Mattingly, 1954)

paullusi (Stone & Farner, 1945)

pernotata (Farner & Bohart, 1944)

perplexa Leicester, 1908

polynesiensis (Marks, 1951)

poweri Theobald, 1905

pseudalbopicta Borel, 1928

pseudoafricana (Chwatt, 1949)

pseudonigeria Theobald, 1910

pseudoscutellaris Theobald, 1910

quasiscutellaris (Farner & Bohart, 1944)

rhungkiangensis (Chang & Chang, 1974)

riversi (Bohart & Ingram, 1946)

robinsoni (Belkin, 1962)

rotana (Bohart & Ingram, 1946)

rotumae (Belkin, 1962)

ruwenzori (Haddow & van Someren, 1950)

saimedres (Huang, 1988)

saipanensis (Stone, 1945)

sampi (Huang, 2004), comb. nov.

schwetzi (Edwards, 1926)

scutellaris (Walker, 1858)

scutoscripta (Bohart & Ingram, 1946)

seampi (Huang, 1974)

seatoi (Huang, 1969)

segermanae (Huang, 1997)

sibirica (Danilov & Filippova, 1978)

simpsoni Theobald, 1905

soleata (Edwards, 1924)

strelitziae (Muspratt, 1950)

subalbopicta (Barraud, 1931)

subargentea (Edwards, 1925)

tabu (Ramalingam & Belkin, 1965)

tongae (Edwards, 1926)

tulagiensis (Edwards, 1926)

unilineata (Theobald, 1906)

upolensis (Marks, 1957)

usambara (Mattingly, 1953)

varuae (Belkin, 1962)

vinsoni (Mattingly, 1953)

wadai (Tanaka, Mizusawa & Saugstad, 1979)

w-albus Theobald, 1905

woodi (Edwards, 1922)

Nomina dubia

ciliaris (Linnaeus, 1767)

niveus (von Eichwald, 1837)

rimandoi (Basio, 1971)

Tanakaius Reinert, Harbach & Kitching, 2004

savoryi (Bohart, 1957)

togoi (Theobald, 1907)

Udaya Thurman, 1954

argyrurus (Edwards, 1934)

lucaris Macdonald & Mattingly, 1960

subsimilis (Barraud, 1927)

Vansomerenis Reinert, Harbach & Kitching, gen. nov.

hancocki (van Someren, 1962), comb. nov.

luteostriata (Robinson, 1950), comb. nov.

pulchrithorax (Edwards, 1939), comb. nov.

Verrallina Theobald, 1903

Subgenus Harbachius Reinert, 1999

abdita (Barraud, 1931)

consonensis (Reinert, 1973)

fragilis Leicester, 1908

hamistylus (Laffoon, 1946)

indecorabilis Leicester, 1908

nobukonis (Yamada, 1932)

pahangi (Delfinado, 1968)

ramalingami (Reinert, 1974)

robertsi (Laffoon, 1946)

srilankensis (Reinert, 1977)

stunga (Klein, 1973)

uniformis (Theobald, 1910)

yusafi (Barraud, 1931)

Subgenus Neomacleaya Theobald, 1907

adusta (Laffoon, 1946)

agrestis (Barraud, 1931)

andamanensis (Edwards, 1922)

assamensis Bhattacharyya, Tewari, Prakash, Mohapatra & Mahanta, 2004

atriisimilis (Tanaka & Mizusawa, 1973)

atria (Barraud, 1928)

campylostylus (Laffoon, 1946)

cauta (Barraud, 1928)

clavata (Barraud, 1931)

comata (Barraud, 1931)

comosa (Reinert, 1974)

cretata (Delfinado, 1967

cyrtolabis (Edwards, 1928)

gibbosa (Delfinado, 1967)

harrisonica (Reinert, 1974)

hispida (Delfinado, 1967)

incerta (Edwards, 1922)

indica (Theobald, 1907)

johnsoni (Laffoon, 1946)

johorensis (Reinert, 1974)

komponga (Klein, 1973)

lankaensis (Stone & Knight, 1958)

latipennis (Delfinado, 1967)

leicesteri (Edwards, 1917)

macrodixoa (Dyar & Shannon, 1925)

margarsen (Dyar & Shannon, 1925)

neomacrodixoa (King & Hoogstraal, 1947)

nigrotarsis (Ludlow, 1908)

notabilis (Delfinado, 1967)

nubicola (Laffoon, 1946)

panayensis (Ludlow, 1914)

petroelephantus (Wijesundara, 1951)

philippinensis (Delfinado, 1968)

phnoma (Klein, 1973)

prioekanensis (Brug, 1931)

protuberans (Delfinado, 1967)

pseudodiurna (Theobald, 1910)

pseudomediofasciata (Theobald, 1910)

pseudovarietas (Reinert, 1974)

rami (Barraud, 1928)

rara (Delfinado, 1968)

sabahensis (Reinert, 1974)

seculata (Menon, 1950)

singularis (Leicester, 1908)

sohni (Reinert, 1974)

spermathecus (Wijesundara, 1951)

torosa (Delfinado, 1967)

uncus (Theobald, 1901)

vallistris (Barraud, 1928)

varietas (Leicester, 1908)

virilis Leicester, 1908

yerburyi (Edwards, 1917)

Subgenus Verrallina

azureosquamata (Bonne-Wepster, 1948)

bifoliata (King & Hoogstraal, 1947)

butleri (Theobald, 1901)

carmenti (Edwards, 1924)

cuccioi (Belkin, 1962)

cunninghami (Taylor, 1944)

dux (Dyar & Shannon, 1925)

embiensis (Huang, 1968)

foliformis (King & Hoogstraal, 1947)

funerea (Theobald, 1903)

iriomotensis (Tanaka & Mizusawa, 1973)

killertonis (Huang, 1968)

leilae (King & Hoogstraal, 1947)

lineata (Taylor, 1914)

lugubris (Barraud, 1928)

mccormicki (Belkin, 1962)

milnensis (King & Hoogstraal, 1947)

multifolium (King & Hoogstraal, 1947)

obsoleta (Huang, 1968)

parasimilis (King & Hoogstraal, 1947)

pipkini (Bohart, 1957)

quadrifolium (Brug, 1934)

quadrispinata (King & Hoogstraal, 1947)

reesi (King & Hoogstraal, 1947)

sentania (King & Hoogstraal, 1947)

similis (Theobald, 1910)

simpla (King & Hoogstraal, 1947)

trispinata (King & Hoogstraal, 1947)

vanapa (Huang, 1968)

variabilis (Huang, 1968)

Nomen dubium

perdita Leicester, 1908

Zavortinkius Reinert, 1999

brunhesi (Reinert, 1999)

brygooi (Brunhes, 1971)

fulgens (Edwards, 1917)

geoffroyi (Reinert, 1999)

huangae (Reinert, 1999)

interruptus (Reinert, 1999)

longipalpis (Grünberg, 1905)

monetus (Edwards, 1935)

mzooi (van Someren, 1962)

phillipi (van Someren, 1949)

pollinctor (Graham, 1910)

Zeugnomyia Leicester, 1908

aguilari Baisas & Feliciano, 1953

fajardoi Baisas & Feliciano, 1953

gracilis Leicester, 1908

lawtoni Baisas, 1946

APPENDIX 5

Abbreviations of aedine genera

Recommended two-letter abbreviations of formally recognized genera of Aedini.

Abraedes Zavortink = Ab.

Aedes Meigen = Ae.

Alanstonea Mattingly = As.

Albuginosus Reinert = Al.

Armigeres Theobald = Ar.

Ayurakitia Thurman = Ay.

Aztecaedes Zavortink = Az.

Belkinius Reinert = Be.

Bothaella Reinert = Bo.

Bruceharrisonius Reinert = Br.

Christophersiomyia Barraud = Cr.

Collessius Reinert, Harbach & Kitching = Co.

Dahliana Reinert, Harbach & Kitching = Da.

Diceromyia Theobald = Di.

Dobrotworskyius Reinert, Harbach & Kintching = Db.

Downsiomyia Vargas = Do.

Edwardsaedes Belkin = Ed.

Eretmapodites Theobald = Er.

Finlaya Theobald = Fl.

Fredwardsius Reinert = Fr.

Georgecraigius Reinert, Harbach & Kitching = Gc.

Geoskusea Edwards = Ge.

Gilesius Reinert, Harbach & Kitching = Gi.

Gymnometopa Coquillett = Gy.

Haemagogus Williston = Hg.

Halaedes Belkin = Ha.

Heizmannia Ludlow = Hz.

Himalaius Reinert, Harbach & Kitching = Hi.

Howardina Theobald = Hw.

Huaedes Huang = Hu.

Hulecoeteomyia Theobald = Hl.

Indusius Edwards = In.

Isoaedes Reinert = Ia.

Jarnellius Reinert, Harbach & Kitching = Ja.

Jihlienius Reinert, Harbach & Kitching = Ji.

Kenknightia Reinert = Ke.

Kompia Aitken = Ko.

Leptosomatomyia Theobald = Lp.

Levua Stone & Bohart = Le.

Lorrainea Belkin = Lo.

Macleaya Theobald = Mc.

Molpemyia Theobald = Mo.

Mucidus Theobald = Mu.

Neomelaniconion Newstead = Ne.

Ochlerotatus Lynch Arribalzaga = Oc.

Opifex Hutton = Op.

Paraedes Edwards = Pr.

Patmarksia Reinert, Harbach & Kitching = Pm.

Phagomyia Theobald = Ph.

Pseudarmigeres Stone & Knight = Pa.

Pseudoskusea Theobald = Pk.

Psorophora Robineau-Desvoidy = Ps.

Rampamyia Reinert, Harbach & Kitching = Ra.

Rhinoskusea Edwards = Rh.

Scutomyia Theobald = Sc.

Skusea Theobald = Sk.

Stegomyia Theobald = St.

Tanakaius Reinert, Harbach & Kitching = Ta.

Udaya Thurman = Ud.

Vansomerenis Reinert, Harbach & Kitching = Va.

Verrallina Theobald = Ve.

Zavortinkius Reinert = Za.

Zeugnomyia Leicester = Ze.