Diptera: Culicidae: Aedini) Based on Morphological Data from All Life Stages

Al 14I Zoological Journal of the Linnean Society, 2006, 148, 1-101. With 2 figures

Phylogeny and classification of Finlaya and allied taxa (Diptera: Culicidae: Aedini) based on morphological data from all life stages

JOHN F. REINERTI, RALPH E. HARBACH2 and IAN J. IUTCHING2 Center for Medical, Agricultural and Veterinary Entomology, United States Department of Agriculture, Agricultural Research Service, 1600/1700 SW 23rd Drive, Gainesville, FL 32608-1067, USA 2Department of Entomology, The Natural History Museum, Cromwell Road, London SW7 5BD, UK

Received July 2005; accepted for publication March 2006

The phylogenetic relationships and generic assignments ofFinlaya 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 Ochierotatus 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 Coquil- lett), Jihlienius gen. nov. (Aedes chungi Lien), Patmarksia gen. nov. (Leucomyia australiensis variety papuensis Taylor), Rampamyia gen. nov. (Culex notoscriptus Skuse) and Vansoinerenis gen. nov. (Aedes pulchrithorax Edwards) are introduced for the type species indicated in parentheses, and their allies. Additionally, Hors fallius 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. Journal compilation © 2006 The Linnean Society of London, Zoological Journal of the Linnean Society, 2006, 148, 1-101. No claim to original US government works. ADDITIONAL KEYWORDS: cladistics - mosquitoes - systematics - taxonomy.

INTRODUCTION ognized species, 32 subspecies, nine infrasubspecific forms/varieties and nine nomina dubia. Reinert, Har- Aedini (sensu Belkin, 1962) is the largest tribe in fam- bach & Kitching (2004) conducted a phylogenetic anal- ily Culicidae (Diptera). It includes 1235 currently rec- ysis of higher-level relationships within the tribe using 172 morphological characters from all life stages Corresponding author. E-mail: and 119 exemplar species representing the 12 genera [email protected] and 56 subgenera that were recognized at the time. All

Journal compilation © 2006 The Linnean Society of London, Zoological Journal of the Linnean Society, 2006, 148, 1-101 No claim to original US government works. 2 J. F. REINERT ET AL. of the genera except Aedes and Ochierotatus were and subgroups are by no means sacrosanct and we recovered as monophyletic. (The authors of all aedine already know, from life history studies, that reassess- taxa are cited in Appendix 4.) In addition to 12 for- ment of the present placing of some species will be merly recognized genera, Reinert et cii. (2004) pro- needed. This applies particularly to rockpool species. posed generic status for 32 subgenera of Aedes and Belkin (1962) followed the classification of Knight & Ochierot at us, raised a dade consisting of Oc. (Finiaya) Marks (1952) but stated that Finiaya as understood at and its relatives (Kochi Group) to generic rank as present is a large heterogeneous assemblage of genus Faniaya, elevated Downsiomyia from synonymy species . . . and I believe that a natural classification with Finiaya as the generic name for a dade compris- of Finiaya cannot be developed until the immature ing Oc. (Fin.) niveus and its relatives (Niveus Group), stages are carefully studied. Tanaka, Mizusawa & and described a new genus, Tanakaius, to accommo- Saugstad (1979) recognized that previous attempts at date Oc. (Fin.) togoi and Oc. (Fin.) savoryi (Togoi classification of Finiaya did not reflect a natural clas- Group). Genus Aedes was restricted to the former sub- sification and proposed nine groups for species of Fin- genus Aedes and genus Ochierotatus was limited to a iaya occurring in the Japanese and Korean area based dade consisting of the type species, six related species primarily on characters of the male genitalia and lar- and members of subgenus Rusticoidus. These actions vae. Schick (1970) revised the Terrens Group and left species previously included in subgenera Aedimor- Arnell & Nielsen (1972) revised the Varipalpus Group phus and Cci ncraedes of Aedes and subgenera Bruce- based on characters of the adult and immature stages. harrisonius, Chaetocruiomyia, Finiaya, Macieaya, Zavortink (1972) removed all New World species from Moipemyia, Ochierotatu.s, Protomacieaya and Pseu- Finiaya and placed them in subgenera Aztecaedes, doskusea of Ochierotatus without generic assign- Gymnometopa, Protomacleaya and Ochierotatus of ments. The present study is the first step toward Aedes and subgenus Conopostegus of Haemagogus resolving the relationships and generic placements of based on features of the adults, pupae and larvae. these taxa. It treats species and species-groups of Reinert (1990, 1993, 1999b, 2003) transferred some uncertain taxonomic position (incertae sedis) that species to other subgenera, i.e. Kenknightia, Moipe- were previously included in subgenus Finiaya of myia, Zavortinkius and Bruceharrisonius. Reinert Ochierotatus, and species previously removed from (2000a) elevated Ochierotatus to generic rank and Finiaya by Zavortink (1972) and placed in subgenera transferred subgenus Finiaya to that genus. Reinert Protomacleaya and Ochierotatus of Ochierot at us. (2002e) separated subgenus Finiaya and related sub- Edwards (1932) provided the first comprehensive genera of Ochierotatus into Assemblages and Sub- classification of Finiaya Theobald (as a subgenus of assemblages based on features of the female genitalia. Aedes). He recognized eight groups within Finiaya He mentioned that Subgenus Finiaya, as currently (designated groups A—H) based on a few morphological configured, contains several diverse groups that are characters of adults. Barraud (1934) followed markedly different in the female genitalia; therefore, Edwards arrangement of genera and subgenera. the above definition of the subgenus is somewhat Knight (1948) recognized that Edwards Group F con- imprecise and indicates the need for a revision of the tained more or less unrelated forms and therefore entire taxon. However, this revision should include all divided it into eight divisions (denoted by Roman life stages as suggested by Belkin (1962). Finally, numerals). Knight & Marks (1952) revised and Reinert et al. (2004), as noted above, formally limited expanded the classification of Edwards and added genus Finiaya to the Kochi Group. numerous subgroup categories. They also based their Few molecular phylogenetic studies of aedine taxa classification almost exclusively on adult characters have been published. Wesson, Porter & Collins (1992) and indicated that a number of the groups were unnat- examined the relationships of six aedine species (four ural assemblages. Whereas Knight & Hull (1951), Aedes, one Haemagogus, one Psorophora) based on when examining the Philippine fauna, utilized the rDNA ITS2 sequence data. Besansky & Fahey (1997) group categories of Knight & Marks (1952) (they noted included three Aedes and one Haemagogus among 13 this paper was in press), subsequent authors, i.e. mosquito species in a study of relationships based on Bohart (1957), Marks & Hodgkin (1958), Macdonald & sequence data for the nuclear protein-coding white Traub (1960), Lee et cii. (1982) and Lu & Ji (1997), pro- gene. Kumar, Black & Rai (1998) investigated the vided some modifications to their classification, again relationships of 15 culicine species representing six based primarily on adult characters. Dobrotworsky genera, including nine species of Aedes and one (1965) followed Knight & Marks arrangement and each of Armigeres and Haemagogus. More recently, applied it to the ten species occurring in Victoria, Aus- Cook et al. (2005) investigated the relationships tralia. Marks (in Lee et al., 1982) stated that Knight & among 20 species of A edes representing six tradition- Marks.. . subdivision of the groups of subgenus Fin- ally recognized subgenera, including Aedes (1 spe- iaya was based on adult characters only. These groups cies), Aedimorphus (1), Diceromyia (2), Haiaedes (3),

Ochierotatus ( 10) and Stegomyia (3), based on As noted above, Reinert et al. (2004) formally recog- sequenceIT data for the cytochrome oxidase c subunits I nized 46 genera for most of the species previously and (COT, COIl) of mtDNA. Finally, Behbahani included in 12 genera. Many species previously et al. (2005) used ITS2 sequences to construct a phy- included in two subgenera of former genus Aedes and logenetic tree for five species of Stegomyia. The eight subgenera of former genus Ochierotatus, includ- results of Wesson et al. (1992), Besansky & Fahey ing subgenus Fi niaya , could not be placed in any of the (1997) and Kumar et al. (1998) indicated a paraphyl- 46 genera, and are regarded as incertae sedis (i.e. of etic Aedini, but these studies provided little insight uncertain placement) within Aedini. To avoid confu- into the phylogeny and classification of the tribe sion, species of uncertain generic and subgeneric because they included very few generic-level taxa and placement, and generic-level names used in the previ- few species. For example, whereas St. aegypti and ous sense that include these species, are enclosed St. aibopicta formed a strongly supported sister pair within single quotation marks herein to distinguish in the white-gene phylogeny of Besansky & Fahey them from formally recognized monophyletic taxa. (1997), they were not placed in a congeneric relation- The objective of the present study is to clarify the rela- ship in the ITS2 phylogeny of Wesson et al. (1992) or tionships and generic assignments of Finiaya and the RFLP phylogeny of Kumar et al. (1998). All three related taxa of uncertain taxonomic position in the of these studies included Ochierotatus (Protoma- classification proposed by Reinert et al. (2004). Herein cieaya) triseriatus, which was strongly paired with we present the results of cladistic analysis of these Psorophora in the phylogeny of Wesson et al. (1992) taxa and propose further changes to the classification and with Haemagogus in the phylogenies of Besansky of Aedini that more accurately reflect natural & Fahey (1997) and Kumar et al. (1998). It is inter- affinities. esting to note that Wesson et cii. (1992) suggested that Aedes sensu auctoruin should be split into two genera, one including Oc. (Pro.) triseriatus as Ac. MATERIAL AND METHODS (Pro.) triseriatus] along with species of Haemagogus and Psorophora. The COI and COIl phylogenies of TAXA EXAMINED Cook et cii. (2005) agreed with respect to species-level The study taxa (Appendix 1; see also for their groupings, but because sequences for neither gene authorship and geographical distribution) include an were available for all the species analysed, it was ingroup of 74 species. This group comprises species difficult to draw firm conclusions regarding the of Finiaya and Protomacieaya that were placed interspecies relationships. Support for supraspecific within the polyphyletic assemblage of Ochierotatus clades was either very low or meaningless in cases basal to Aedes in figure 4 of Reinert et al. (2004), where higher-lever taxa were represented by several i.e. species included in the clades between Oc. (Fin.) samples of a single species. In general, dade support, chrysolineatus and Oc. (Fin.) poiciiius (now Finiaya assessed by bootstrap proportions, was slightly poiciiia), and incertae sedis species listed in their higher for the COIl phylogeny. Whereas the mono- appendix 4. The ingroup also includes species that phyly of Di. tayiori + Di. furcifer ( as Ac. tayiori and were classified as Finiaya prior to Zavortink (1972). Ae. furcifer) was weakly supported in the COT The outgroup taxa include both aedine and non- phylogeny, it was very strongly supported (bootstrap aedine species. The aedine outgroup includes repre- value = 99) in the COIl phylogeny. Likewise, the sentatives of all generic-level taxa arrayed in the monophyly of St. iuteocephaia + (St. aibopicta + basal polyphyletic assemblage of Ochierotatus in fig- St. aegypti) (as species of Aedes) received low support ure 4 of Reinert et al. (2004), as well as species in the COI phylogeny and modest support (bootstrap selected from major clades within the distal poly- value = 53) in the maximum-likelihood tree of COIl phyletic assemblage of Aedes. The non-aedine out- sequences. Eight species of Ochierotatus included in group comprises the four outgroup taxa used in the the analysis of COT data (COIl data were not avail- analyses of Reinert et al. (2004), i.e. Culex quinque- able) were placed in a sister-group relationship to Oc. fascialus, Cuiiseta inornata, Mansonia titiiians and (Rusticotdus) rusticus (as Ac. rusticus), which was sis- Orthopodomyia signifera. ter to Ac. cinereus, but all of the internal branches We initially intended to include several additional were only weakly supported. The three species of species of Finiaya in the study but this was not pos- Halaedes (not included in the COI phylogeny) formed sible because specimens were unavailable, several life a strongly supported dade in the COIl phylogeny stages were unknown, and/or published descriptions (bootstrap value = 99) comprising ashworthi + (ward- and illustrations were inadequate. Taxa examined angensis + austraiis) (as Aedes species). Overall, the (Appendix 1) include type species of most genus-level results of Cook et al. (2005) are not inconsistent with names (abbreviations for genera and subgenera fol- those of Reinert et al. (2004). low Reinert, 2001a; Reinert et al., 2004). However, we

Journal compilation © 2006 The Linnean Society of London, Zoological ,Journal of the Linnean Society, 2006, 148, 1-101 No claim to original US government works. 4 J. F. REINERT ETAL. did not include type species if they had several miss- CHARACTER DESCRIPTIONS ing life stages or inadequate specimens for detailed study. The data (Appendix 2) comprise 232 characters from Generally, we examined 3-6 specimens (range 1-20) eggs (3), fourth-instar larvae (74), pupae (31), females of each life stage and structure for each species. How- (71), males (11), female genitalia (16), male genitalia ever, for some life stages of a few species no material (25) and habitat of immature stages (1). Characters was available and literature descriptions were either from Reinert et al. (2004) are included in parentheses inadequate or lacking this information, e.g. pupae of following current character numbers. Data for the characters were derived from direct observations Oc. echinus, Oc. pulchri venter and Oc. rubrithorax, and eggs of many species. We also studied individually except where life stages were not available or struc- reared, pin-mounted adults with associated slide- tures were missing from available specimens. We mounted fourth-instar larval and pupal exuviae where coded some such missing data from literature sources available. A phase contrast or differential interference (e.g. characteristics of most eggs) or collection data contrast microscope with 400x magnification is needed sheets. The remaining missing data were denoted by a to observe the very slender distal parts of many setae States in continuous characters were determined for the measurement of total length. Lesser magnifi- either by clear gaps in the observed counts or mea- cation normally results in incomplete measurements, surements (e.g. character 28), or by reference to as noted in many published illustrations of larval and observed intraspecific variation (e.g. character 38). pupal stages. Pinned adults were examined with a Characters that could not be scored owing to the binocular stereomicroscope utilizing cold white light absence of homologous structures (dependent charac- delivered by a fibre-optic illumination system. An ters) are indicated by a dash (—), e.g. character 102 in adjustable examination stage with biaxial rotation Mansonia. All multistate characters were treated as unordered. Polymorphic characters were explicitly capability (see figure 125.3 in Russell et al., 1963) allowed observation and lighting of specimens at any coded as exhibiting only those states observed. angle. Adult characters were derived from females unless A few characters used in the analysis of Reinert otherwise noted. Males of many species often have fewer setae and scales than females, and would skew et al. (2004) are excluded from the present study because all the study taxa exhibit the same character the coding of some character states. We have reinter- state. Some characters were modified by adding or preted the homologies of several structures from those deleting character states or altering the range of con- used in some published works. tinuous variables to score the study taxa more accu- Anatomical nomenclature and chaetotaxy follow rately. A number of new characters are included that Harbach & Knight (1980, 1982) except for terms proposed by Reinert (1990, 1999b, 2000b, 2002a). Many were not used by Reinert et al. (2004). Character states corrected in the present study from those incor- structures of characters used in the present study are illustrated in these references. However, references to rectly entered in Appendix 2 of Reinert et al. (2004) (in character states illustrated in other publications are parentheses) are: Oc. (Brh.)greenii = 25 (19), 67 (48), provided in the explanations of the characters listed 68 (49); Oc. (Mac.) tremulus = 81(59), 114 (88), 183 below. (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 col- EGGS lections: National Museum of Natural History, 1(1). Deposition: (0) laid singly; (1) laid in a raft; (2) Smithsonian Institution, Washington, DC; The Na- laid in a mass. Numerous authors, e.g. Howard, Dyar tural History Museum, London, UK; Department of & Knab (1913, 1917), Edwards (1932, 1941), Ross Zoology and Entomology, University of Queensland, (1947), Carpenter & LaCasse (1955), Belkin (1962), Brisbane, Australia; Australian National Insect Col- Ross & Horsfall (1965), Dobrotworsky (1965), Bohart lection, CSIRO Division of Entomology, Canberra, & Washino (1978) and Clements (1999), routinely indi- Australia; Florida State Collection of Arthropods, cated that the eggs of genera Aedes and Ochierotatus Division of Plant Industry, Gainesville, Florida; (as genus Aedes and previous subgenera) are laid sin- Laboratoire/Cellule Entomologie, EID Méditerranée, gly (state 0). However, egg deposition and morphology Montpellier, France; Bohart Museum, University of (also see characters 2 and 3) of numerous species of California, Davis, California; National Institute of Aedini are unknown (see Appendix 2). For examples of Malariology, Parasitology and Entomology, Hanoi, state (1) see Marshall (1938: figs 10, 11), Forattini Vietnam; Florida Medical Entomology Laboratory, (1965a: figs 9-11) and Dahl (1988: fig. 10.1), and for University of Florida, Vero Beach, Florida; and the state (2) see Forattini (1965b: fig. 18) and Clements first author. (1999: figs 40.27, 40.29).

Journal compilation © 2006 The Linnean Society of London, Zoological Journal oft/ic Linnean Society, 2006, 148, 1-101 No claim to original US government works. PHYLOGENY AND CLASSIFICATION OF FINLAYA 5

2(2, in part). Shape: (0) anterior end flattened, poste- 6(new). Antenna, spicules: (0) absent; (1) few; (2) rior end rounded; (1) both ends tapered; (2) anterior numerous. Some aedine species have antennae with- end elongate and narrow. Various aedine eggs exhib- out spicules (state 0), e.g. Ahraedes (see Zavortink, iting state (1) are illustrated in numerous articles, e.g. 1972: fig. 34, Ab. papago), Aztecaedes, Finlaya, Gym- Howard et al. (1913), Ross & Horsfall (1965), Kalpage nometopa, Halaedes, Kompia and Stegomyia. Other & Brust (1968), Mattingly (1970), Moriya, Yabe & species have a few, normally small and scattered Harada (1973), Matsuo, Lien & Yoshida (1974a, b), spicules (state 1), e.g. Downsiomyia, Haemagogus, Hinton (1981), Linley & Chadee (1990, 1991), Chadee Howczrdina, Kenknightia (see Reinert, 1990: fig. 24, & Bennett (1990), Linley, Geary & Russell (1991a, b), Ke. dissimilis) and Zavortinkius. Many species have Linley & Craig (1993, 1994) and Forattini (1996). numerous, normally well-developed spicules covering However, Reinert (2005) pointed out that only about all or most of the antenna (state 2), e.g. Aedes (see 16% of the species of Aedini have morphological Tanaka et al., 1979: fig. 134, Ae. yamadai), Mucidus, descriptions and/or illustrations of the egg stage pub- Psorophora, Tanakaius and Oc. (Rusticoidus). lished, and many of these are incompletely described and/or illustrated. The outgroup species are illus- 7(7). Seta 1-A, length: (0) short, :^ 3.0 times antennal trated as follows: Cs. inornata (Mattingly, 1972: width at point of attachment; (1) longer, ^! 3.1 times fig. 2a) (state 0), Cx. quinquefasciatus (Forattini, 1996: antennal width at point of attachment. The usual con- figs 6-13A, 7-33C) (state 0), Ma. titillans (Linley, dition of seta 1-S is moderately long to long (state 1) in Linley & Lounibos, 1986: fig. 1; Linley, 1989: fig. 2c,e) most Aedini, e.g. Oc. (Protomacleaya) (see figures in Zavortink, 1972) and Zavortinkius, but this seta is (state 2) and Or signifera (Howard et al., 1913: short in some taxa (state 0), e.g. Opifex (Opi.) fuscus fig. 669; Pratt & Kidwell, 1969: fig. 7) (state 0). Belkin (1962) believed the eggs of Fl. kochi described by and most Stegomyia (see Belkin, 1962: figs 208, 314, Buxton & Hopkins (1927) were probably those of Op. (Opi.) fuscus and St. aegypti). Fl. oceanica, whereas Lee et al. (1982) noted that the 8(6). Seta 1-A, development: (0) single or 2-branched; eggs of Ft. kochi described by Buxton & Hopkins (1) ^ 3 branches. Many species of Aedini have seta 1-A (1925, 1927) were probably Fl. oceanica, but could be, single (state 0) (see Belkin, 1962: fig. 231, Oc. (Fin.) in part, those of Fl. samoana. albilabris) whereas others are multiple-branched (state 1) (see Belkin, 1962: fig. 234, Oc. (Fin.) argy- 3(3). Outer chorion: (0) pattern absent or weakly devel- ronotum). However, this seta is single or two-branched oped; (1) pattern well developed; (2) with spiny appear- (state 0) in some species, and it is also two-branched in ance, each cell with elongate anteriorly inclined some specimens of a few species in which seta 1-A is tubercle. See the literature cited for characters 1 and 2. multiple-branched, which are scored (0,1). 9(8). Seta 2,3-A, position: (0) insertion apical or nearly LARVAE (FOURTH-INSTARS) apical; (1) insertion noticeably subapical. Setae 2-A 4(4). Lahiogula: (0) short, length

The position of seta 4-C appears to be constant in cies ofAedini is interpreted as seta 13-C (see fig. 277 of generic-level taxa for which the taxonomy is stable. Mu. alternans in Belkin, 1962; labelled as 12-C). However, both states occur in some of the larger subgenera, but in different species groups. See Belkin 20(15). Seta 13-C, development: (0) single; (1) (1962) for illustrations of state (0) (his figs 218, 277, branched. Seta 13-C is normally single (state 0) in Ge. longiforceps and Mu. alternans) and state (1) (his many of the species examined. However, it is branched fig. 211, Ha. australis). See character 15 for comment. (state 1) in a number of other taxa, e.g. Aedes, Geoskusea, Kenknightia, Levua, Mucidus, Psorophora, 12(11, in part). Seta 4-C, length: (0) short; (1) moder- Rhinoskusea and Zavortinkius. ate; (2) long; (3) very long. The lengths of setae 4-7-C are compared to the median length of the dorsal 21(new). Seta 14-C, development: (0) single; (1) apodeme (DAp), preferably in exuviae. The setae are branched. Seta 14-C is often single (state 0) (see considered short (:^ 0.19 length of DAp), moderate Tanaka et al., 1979: fig. 101, Ta. savoryi) and may be (0.21-0.38 length of DAp), long (0.40-0.81 length of slender or stiff and stout. The seta is branched (state DAp) or very long (^! 0.90 length of DAp). 1) (see Tanaka et al., 1979: fig. 102, Oc. (Fin.) seoulensis) and may be slender or stellate in some species. 13(new). Seta 5-C, length: (0) short; (1) moderate; (2) long; (3) very long. See character 12 for explanation of 22(16). Seta 19-C: (0) absent; (1) present. The occur- setal lengths. rence of seta 18-C and the presence or absence of seta 19-C ventrally on the cervical membrane of larvae 14(new). Seta 5-C, development: (0) single; (1) belonging to 29 genera, 72 subgenera and 331 species branched. Larval seta 5-C is single (state 0), e.g. of Culicidae was reported by Hochman & Reinert Abraedes, Aztecaedes, Finlaya, Howardina and (1974). See illustrations in Hochman & Reinert (1974) Zavorunkius. Other taxa have the seta branched for examples and a discussion of the occurrence of seta (state 1), e.g. Aedes, Downsiomyia, Geoskusea, Levua 19-C in the large and heterogeneous genus Ochlerota- and Tanakaius. tus sensu auctorum, and its taxonomic importance and 15(12). Seta 6-C, position: (0) insertion anterior to seta possible phylogenetic significance. 7-C; (1) insertion at same level or posterior to seta 7-C. 23(17). Ventromedian cervical scierite: (0) absent; (1) The positions of setae 6-C and 7-C are determined on present. A small solid or fragmented sclerite is present head capsules that are horizontal with the thorax and on the ventromedian area of the cervix in many aedine abdomen, normally exuviae. If the head is in a pro- larvae. It appears to be absent in Ko. purpureipes, Le. gnathous orientation and turned down anteriorly, geoskusea, Oc. (Cha.) wattensis, Oc. (Fin.) macfar- these and other setae may appear to be positioned dif- lanei, Oc. (Mac.) tremulus, Psorophora, Rhinoskusea, ferently. This also applies to characters 11 and 17. Zavortinkius and the non-aedine outgroup species. See illustrations in Tanaka et al. (1979: figs 99, 101, Reinert (1976a) provided a review and illustrations of Ta. togoi and Ta. savoryi) for state (0) and their this sclerite in Culicidae. fig. 128 (Ge. baisasi) for state (1). 24(18). Setae 1-3-P: (0) not attached to a common setal 16(new). Seta 6-C, length: (0) short; (1) moderate; (2) support plate; (1) two or three of these setae attached long; (3) very long. See character 12 for explanation of to a common setal support plate. See Zavortink (1972: setal lengths. figs 18, 28, Oc. (Pro.) kompi and Oc. (Pro.) knabi) for 17(13). Seta 7-C, position: (0) insertion anterior to seta examples of state (0) and his figs 9, 12, Oc. (Pro.) tris- 5-C; (1) insertion at approximately same level as seta eriatus and Oc. (Pro.) zoosophus, for state (1). 5-C; (2) insertion posterior to seta 5-C. See Belkin 25(19). Seta 1-P, length: :!^ length of seta 2-P; (1)> (1962: figs 237, 239, 241, etc., (0) Finlaya, as Kochi length of seta 2-P. Seta 1-P is normally longer (state 1) Group) for examples of state (0), Zavortink (1972: than seta 2-P in Aedini, but it is shorter than or equal fig. 34, Ab. papago) for state (1) and Tanaka et al. in length to 2-P (state 0) in Finlaya (as Kochi Group) (1979: figs 99, 101, Ta. togoi and Th. savoryi) for state (see Belkin, 1962: fig. 239, Fl. hurnetti), Hw. walkeri, (2). See character 15 for comment. Sc. arboricola, Oc. (Och.) muelleri and some 18(new). Seta 7-C, length: (0) short; (1) moderate; (2) Oc. (Finlaya). long. See character 12 for explanation of setal lengths. 26(21). Seta 5-P, length: (0) :5 length of seta 6-P; (1)> 19(14). Seta 12-C, position: (0) insertion mesal to or at length of seta 6-P. The length of seta 5-P in relation to same level as seta 13-C; (1) insertion lateral to seta 13- that of seta 6-P is relatively consistent for most generic- C. Seta 13-C is longer and normally borne lateral to level taxa that are well defined. See Belkin (1962) for seta 12-C in Culicidae. For this reason, the longer seta illustrations of state (0) (his fig. 243, Fl. franclemonti) in a mesal position relative to the other in some spe- and state (1) (his fig. 231, Oc. (Fin.) albilabris).

27(22). Seta 5-P, development: (0) single; (1) branched. an example of single seta 2-T (state 0) and Belkin Seta 5-P is single (state 0) in several taxa, e.g. Aedes, (1962: fig. 211, Ha. australis) for an example of Geoskusea, Hg. (Conopostegus), Isoaedes , Levua, Muci- branched 2-T (state 1). dus and Rhinoskusea. In other taxa, this seta is Seta 4-T, development: branched (state 1), e.g. Abraedes, A.ztecaedes, Gym- 35(new). (0) single; (1) ^! 2 ^! 3 branches, stellate. Stel- nometopa and Kompia. branches, not stellate; (2) late setae have three to numerous, moderately stout to 28(23). Seta 8-P, length: (0) !!^ 1.2 times length of seta 4- stout, stiff branches that are truncate or bluntly P; (1) ^ 1.8 times length of seta 4-P. Seta 8-P is less than pointed (not attenuate) and radiate in different direc- or equal to 1.2 times the length of seta 4-P (state 0) in tions from or near the base (starburst-shaped) (see most taxa examined (see Tanaka et al., 1979: fig. 99, Belkin, 1962: fig. 237, Fl. bougainvillensis). Branches Ta. togoi). It is greater than or equal to 1.8 times the of stellate setae often are of different lengths. In some length of seta 4-P in Aedes, Oc. (Rusticoidus), Oc. (Fin.) species, e.g.Az. ramirezi and Oc. (Pro.) zoosophus, this alhotaeniatus, Oc. (Och.) fluviatilis, Ps. (Jan. ) ferox, Ps. and some other setae approach the stellate condition (Gra.) jamaicensis (see Belkin, Heinemann & Page, except the branches are somewhat narrower and tend 1970: figs 60,67) and the outgroup species Cs. inornata. to be in one plane (fan-shaped). These are scored (1). 29(new). Seta 8-P, development: (0) single or occasion- 36(28). Seta 34, development: (0) single; (1) branched. ally 2-branched; (1) multiple-branched (occasionally 3- The development of seta 3-I is consistent for generic- branched). A few taxa have seta 8-P single or occasion- level taxa (except some of the large ones), with few ally 2-branched (state 0) (see Belkin, 1962: figs 211, exceptions. Some specimens of a few species show an 277, Ha. australis and Mu. alternans). Most taxa have overlap of the two states. seta 8-P multiple-branched, but it is occasionally only 3-branched (state 1) (see Belkin, 1962: figs 239, 316, 37(29). Seta 74, length: (0) <0.45 length of seta 64; ^! Fl. burnetti and Sc. albolineata). (1) 0.55 length of seta 6-I. Seta 7-I is normally moderately long to long (state 1) (^! 0.55 length of seta 30(new). Seta 1-M, length: (0) :!^, 2.5 times length of seta 6-I, see Belkin, 1962: fig. 274, Le. geoskusea) in Aedini, 2-M; (1) ^! 3.5 times length of seta 2-M. See Tanaka but it is shorter in some species (state 0) (:^ 0.45 length et al. (1979: figs 99, 111, Ta. togoi and Oc. (Fin.) oreo- of seta 6-I, see Belkin, 1962: fig. 208, Op. (Opi.)fuscus). philus) for examples of state (0) and their fig. 104, Oc. 38(30). Seta 74, development: (0) single to 3-branched; (Brh.) alektorovi (= kobayashii), for state (1). (1) ^ 4 branches. Seta 7-I is single or occasionally with 31(new). Seta 1-M, development: (0) single (rarely 2- 2 or 3 branches (state 0) in most Aedini (see Belkin, branched); (1) ^! 3 branches. Illustrations of seta 1-M 1962: fig. 227, Oc. (Fin.) notoscriptus). Seta 7-I has single (state 0) in Rhinoskusea are found in Reinert more than three branches (state 1) in Mucidus (see (1976b). Examples of seta 1-M with three or more Belkin, 1962: fig. 277, Mu. alternans). This seta has branches (state 1) are found in Oc. (Protomacleaya) overlapping states in Oc. (Fin.) keefei and Op. (Opi.) (see figures in Zavortink, 1972). fuscus, and is scored (0,1). 32(25). Seta 4.M, development: (0) single; (1) branched. 39(31). Seta 124: (0) absent; (1) present. Seta 12-I is Many generic-level taxa of Aedini examined in the absent (state 0) in Ahraedes, Aedes, Aztecaedes, Gym- present study have seta 4-M branched (state 1) (see nometopa, Howardina, Isoaedes, Kompia, Scutomyia, Schick, 1970: figs 16, 22, Oc. (Pro.) terrens and Oc. Stegomyia, Oc. (Cha.) wattensis, Oc. (Och.) muelleri, (Pro.) berlini). See illustrations in Zavortink (1972: Oc. (Och.) varipalpus and Oc. (Pro.) zoosophus. Rein- figs 32, 34, 37, 40, Ko. purpureipes, Ab. papago, Az. ert (2000a) discussed the presence and absence of seta ramirezi and Gy. mediovittata) for examples of state (0). 12-I in generic-level taxa previously included in Aedes and Ochierot at us. Most taxa examined in the present Seta 7-M, length: < ^! 33(26). (0) length of seta 5-M; (1) study have seta 12-I (state 1) (see Tanaka et al., 1979: length of seta 5-M. Seta 7-M is normally shorter than fig. 86, Oc. communis). seta 5-M (see Tanaka et al., 1979: fig. 98, Oc. (Fin.) hatorii) (state 0) in most Aedini; however, it is longer 40(new). Seta 1-11, development: (0) single; (1) ^! 2 than seta 5-M (state 1) in Geoskusea, Kompia, Tana- branches, not stellate; (2) ^! 3 branches, stellate. Illus- kaius (see Tanaka et al., 1979: fig. 99, Ta. togoi), Ha. trations in Belkin (1962) show the various states of australis, Oc. (Cha.) wattensis, Oc. (Fin.) albocinctus, seta 1-II, i.e. state (0) in fig. 234 (Oc. (Fin.) argyrono- Oc. (Fin.) gilli, Oc. (Mac.) tremulus, Ps. (Jan.) ferox tum), state (1) in fig. 231 (Oc. (Fin.) albilabris) and and the outgroup species Cx. quinquefasciatus. state (2) in fig. 237 (Fl. bougainvillensis). 34(27). Seta 2-T, development: (0) single; (1) branched. 41(new). Seta 241, development: (0) single; (1) See Zavortink (1972: fig. 44, Oc. (Och.) muelleri) for branched. Examples of seta 2-I1 development are illus-

trated in Belkin (1962: fig. 218, Ge. longiforceps) for the length of seta 5-V in his Section II. These mea- state (0) and his fig. 237 (Fl. bougainvillensis) for state surements are slightly modified here. See Appendix 2 (1). for other taxa. 42(new). Seta 5-11, development: (0) single; (1) ^! 2 49(new). Seta 1-Vu, length: (0) short; (1) moderately branches, not stellate; (2) ^! 3 branches, stellate. Seta 5- long; (2) long to very long. The length of seta 1-VII is II is branched in most Aedini, either not stellate (state compared to the length of segment X. Segment X 1) (see Belkin, 1962: figs 227-229, Oc. (Fin.) notoscrip- length is the middorsal length measured along a tus) or stellate (state 2) (see Belkin, 1962: fig. 237, straight line parallel to the longitudinal axis from the Fl. bougainvillensis). This seta is single in a few taxa posterior margin of the saddle (minus marginal spi- (state 0) (see Reinert, 1979: fig. 5, Ia. cavaticus). cules) anteriorly to the first annulation in the membrane cephalad of the saddle. Fourth-instar larvae, not 43(new). Seta 6-I1, length: (0) < length of seta 6-111; (1) exuviae, should be used for measurements of segment ^: length of seta 6-I11. See Tanaka et al. (1979: figs 102, X. Seta 1-VII is considered short (state 0) if it is less 104, Oc. (Fin.) seoulensis and Oc. (Brh.) alektorovi (= than or equal to 0.42 times the length of segment X, kobayashii)) for state (0) and their figs 98, 99, Oc. moderate (state 1) if it is 0.48-0.85 times the length of (Fin.) hatorii and Th. togoi, for state (1). segment X and long to very long (state 2) if it is greater 44(32). Seta 6-11, development: (0) single; (1) branched. than or equal to 0.94 times the length of segment X. Seta 6-I1 is normally branched in Aedini, but it is sin- 50(new). Seta 2-VII, position: (0) insertion near seta 1- gle in Levua, Mucidus and Oc. (Molpemyia) of the VII; (1) insertion far anterior to seta 1-Vu. Seta 2-Vu ingroup, and also in other aedines, e.g. Alanstonea, is inserted near seta 1-Vu (state 0) in most species Eretmapodites, subgenus Harbachius of Verrallina examined (see Tanaka et al., 1979: figs 95, 102, Ar. (Arm.) breinli. and Oc. (Fin.) japonicus and Oc. (Fin.) ,seoulensis). See 45(33). Seta 7-Il, development: (0) similar to seta 7-I; (1) Berlin (1969: figs 18, 44, Hw. walkeri and Hw. sexlin- different than seta 7-I. The length, thickness and eata; Belkin, 1962: figs 237, 251, Fl. bougainvillensis, branching of seta 741 are normally consistent in Fl. hollingsheadi) for examples of seta 2-Vu! inserted generic-level taxa of Aedini but in some of the larger far anterior to seta 1-Vu (state 1). Seta 2-VII is nor- taxa this character is consistent only at the species- mally stellate and located on the anterior half of seg- group level. Length of seta 741 is often slightly less ment VII in state (1). than that of seta 7-I. See Belkin (1962: fig. 208, Op. 51(36). Seta 2-Vu, development: (0) single; (1) (Opi.) fuscus) for an example of state (0) and his fig. 218 branched. Seta 2-VII is normally single and short (Ge. longiforceps) for state (1). Seta 7-I1 is short, slen- (state 0) in most Aedini (see Belkin, 1962: figs 218, der and 2-branched in Oc. (Och.)fluviatilis and differs 274, Ge. longiforceps and Le. geoskusea). This seta has from seta 74, which is long, stout, aciculate and single two or more branches (state 1) in a few taxa, e.g. (see fig. 53 in Zavortink, 1972), and is scored (1). Downsiomyia, Finlaya (see Belkin, 1962: figs 237, 239, 46(34). Seta 8-Il, development: (0) single; (1) branched. 241, Fl. bouganivillensis, Fl. burnetti and Fl. fijiensis), See Zavortink (1972: figs 32, 34, 40, Ko. purpureipes, Haemagogus, Howardina and Scutomyia. Ab. papago and Gy. mediovittata) for examples of sin- 52(new). Seta 3-Vu, position: (0) insertion anterior to gle seta 8-I1 (state 0) and his figs 4, 7, 9 (Oc. (Pro.) seta 1-VII; (1) insertion at same level or posterior to hendersoni, Oc. (Pro.) brelandi and Oc. (Pro.) triser- seta 1-VII. Seta 3-VII is inserted anterior to seta 1-VII iatus) for examples of state (1). (state 0) in most Aedini (see Belkin, 1962: figs 218, 47(new). Seta 6-I11, development: (0) single; (1) 243, Ge. longiforceps and Fl. franclemonti). This seta branched. Seta 6-111 is single (state 0) is a few taxa, is inserted at the same level or posterior to seta 1-VII e.g. Aedes, Isoaedes, Mucidus, Ge. longiforceps, Oc. (state 1) in a few taxa, e.g. Kenknightia, Levua, Opifex, (Fin.) candidoscutellum, Oc. (Fin.) rubrithorax, Tanakaius and Ha. australis (see Belkin, 1962: Oc. (Mol.) pecuniosus, Oc. (Bruceharrisonius), Oc. figs 208, 211, Op. (Opi.) fuscus and Ha. australis). (Ochlerotatus) (several species) and the outgroup 53(new). Seta 3-Vu, development: (0) single; (1) species Ma. titillans and Or signifera. This seta is branched. See Zavortink (1972: figs 32, 34, 37, 40, Ko. branched in most taxa examined. purpureipes, Ab. papago, Az. ramirezi and Gy. mediovittata) for examples of state (0) and Reinert (1990: 48(35). Seta 3-V, length: (0) :!^ 1.55 times length of seta figs 24-32, Kenknightia) for examples of state (1). 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- 54(37, in part). Seta 3-Vu, length: (0) short; (1) mod- V in species of Section I ofOchlerotatus (as defined by erate; (2) long to very long. See character 49 for the Reinert, 2000a) whereas it is shorter than 1.5 times method used to measure the length of this seta. The

Journal compilation © 2006 The Linnean Society of London, Zoological Journal of the Linnean Society, 2006, 148, 1-101 No claim to original US government works. PHYLOGENY AND CLASSIFICATION OF FINLAYA 9 relatively long seta 3-VII in the outgroup species Ma. 0) in most Aedini. This seta is branched (state 1) in titillans is moderated by the very long dorsal length of many of the species that have seta 2-VIII also segment X resulting in a moderate length (state 1), branched (see Belkin et al., 1970: figs 56, 60, Ps. (Pso.) but this condition does not apply to any of the aedine ciliata and Ps. (Jan.) ferox). species examined. 62(43). Comb: (0) few to several scales in one or two 55(new). Seta 10-Vu, development: (0) single; (1) irregular rows; (1) numerous scales in a patch. The branched. Seta lO-Vil is single (state 0) in many aedine development of the comb is consistent for most generic- taxa, e.g. Downsiomyia, Haemagogus (see figures in level taxa; however, in some large taxa it is normally Arnell, 1973), Scutomyia and Tanakaius. This seta is consistent at the species-group level. See examples in branched (state 1) in other taxa, e.g. Finlaya, Opifex Belkin (1962: fig. 316, Sc. albolineata) for state (0) and and Ha. australis (see Belkin, 1962: figs 208, 211). his fig. 243 (Fl. franclemonti) for state (1). 56(38). Seta 12-Vu, position: (0) insertion anterior to 63(44). Comb plate: (0) absent; (1) present. A comb seta 13-VII; (1) insertion at approximately same level plate is normally present in Psorophora (see Belkin as seta 13-VII; (2) insertion posterior to seta 13-Vu. et al., 1970: fig. 67, Ps. (Gra.)jamaicensis) and usually Seta 12-WI is inserted anterior to seta 13-VII (state 0) absent in most other Aedini. It occurs in Stegomyia in a few taxa, e.g. Hg. (Haemagogus) (see figures in (eight species), Sc. arhoricola (most specimens), Do. Arnell, 1973), most Howardina, Kompia, Scutomyia dorseyi, Hg. (Hag.) capricornii, Hg. (Hag.) janthino- and St. africana. It is inserted at approximately the mys, Oc. (Cha.) elchoensis and Oc. (Mac.) tremulus same level as seta 13-VII (state 1) in a few taxa, e.g. (Reinert et al., 2004). Abraedes, Gymnometopa and Hg. (Conopostegus) (see figures in Zavortink, 1972). Most aedine taxa have 64(45). Siphon, acus: (0) absent; (1) present. An seta 12-VII posterior to seta 13-WI (state 2) (see Rein- attached or detached acus located at the base of the ert, 1976b: figs 12, 13, Rhinoskusea). siphon (in line with the pecten) is found in many aedine taxa, e.g. Downsiomyia, Finlaya, and species of 57(39). Seta 12-Vu, development: (0) single; (1) Ochlerotatus and Oc. (Ochlerotatus) examined in this branched. Seta 12-VIII is single (state 0) in many study (see figures ofOchlerotatus in Arnell & Nielsen, Aedini, e.g. Downsiomyia, Finlaya, Howardina (see 1972; Arnell, 1976). This structure is absent in other figures in Berlin, 1969) and Stegomyia. It is branched taxa, e.g. Abraedes, Gymnometopa, Halaedes, Stego- (state 1) in other taxa, e.g. Aedes, Levua, Opifex and myia, Zavortinkius, Oc. (Cha.) wattensis, Oc. (Mac.) Zavortinkius (see Reinert, 1999b: fig. 14). tremulus and Oc. (Molpemyia) (see Reinert, 1993: fig. 8, Oc. (Mol.) pecuniosus). 58(40). Setae 1,2-VIII, insertion: (0) not attached to common setal support plate; (1) both attached to com- 65(46). Pecten: (0) absent; (1) present, spines evenly mon setal support plate. Setae 1,2-VIII are not spaced; (2) present, distal one or more spines more attached to a common setal support plate (state 0) in widely spaced. Most aedine taxa have a pecten with most aedine taxa. Both setae are attached to a common the spines evenly spaced (state 1) (see Belkin, 1962: setal support plate (state 1) in Mucidus (see Belkin, figs 227-229, Oc. (Fin.) notoscriptus). A few taxa have 1962: figs 277, 279, Mu. alternans and Mu. painei). a pecten with the distal one or more spines more 59(new). Seta 1-VIII, length: (0) short; (1) moderate; widely spaced (state 2) (see Tanaka et al., 1979: (2) long. Seta 1-VIII is considered short (state 0) if it is figs 131, 132,Ae. esoensis). The pecten is absent (state less than or equal to 0.38 times the length of seta 2- 0) in the outgroup taxa Ma. titillans and Or signifera. VIII, moderate (state 1) if it is 0.50-0.95 times the 66(47). Seta 1-S, development: (0) one seta at base of length of seta 2-VIII and long (state 2) if it is greater siphon; (1) one seta some distance distally from base of than or equal to 1.10 times the length of seta 2-VIII. siphon; (2) two or more setae some distance from base See Belkin (1962) for examples of state (0) (fig. 211, of siphon. Seta 1-S inserted ventroposteriorly some dis- Ha. australis), state (1) (fig. 234, Oc. (Fin.) argyrono- tance from the base of the siphon, and without acces- tum) and state (2) (fig. 231, Oc. (Fin.) albilabris). sory setae (state 1) (see Tanaka et al., 1979: fig. 101, 60(41). Seta 2-VIII, development: (0) single; (1) Ta. savoryi), is the usual condition in Aedini. Aedes (see branched. Seta 2-VIII is single (state 0) in most Aedini. Tanaka et al., 1979: fig. 131,Ae. esoensis) and Oc.(Rus- It is branched (state 1) in a few taxa, e.g. Opifex (see ticoidus) possess accessory setae in addition to the non- Belkin, 1962: fig. 208, Op. (Opi.) fuscus), Psorophora, basal seta 1-S (state 2). Reinert (1999a, 2000a) dis- of Zavortinkius and Oc. (Bruceharrisonius). cussed the presence accessory setae in these two taxa and noted their occurrence in Oc. (Och.) crinifer, Oc. 61(42). Seta 4-VIII, development: (0) single; (1) (Och.) hexodontus, Oc. (Och.) hokkaidensis and Oc. branched. Seta 4-VIII, like seta 2-VIII, is single (state (Och.) punctor (see Tanaka et al., 1979). Accessory

Journal compilation © 2006 The Linnean Society of London, Zoological Journal of the Linnean Society, 2006, 148, 1-101 No claim to original US government works. 10 J. F. REINERT ETAL. setae on the siphon (state 2) are typical of genus Culex 72(new). Saddle, moderate to well-developed spicules whereas seta 1-S at the base of the siphon (state 0) is on posterior margin: (0) absent; (1) present. See Belkin characteristic of many species of Culiseta Felt. (1962: fig. 314, St. aegypti) for an example of the saddle without well-developed spicules on the posterior 67(48). Seta 6.S, development: (0) absent; (1) short; (2) margin and his fig. 237 (Fl. hougainvillensis) for an long. Setae 6-S and 9-S are absent (state 0) from the example of well-developed spicules on this area. highly modified siphon of Ma. titillans (see Belkin et al., 1970: fig. 50). See illustrations in Belkin (1962: 73(52). Seta 1-X, position: (0) inserted on saddle; (1) fig. 227, Oc. (Fin.) notoscriptus) for state (1) and his inserted ventral to saddle. See Belkin (1962: fig. 227, Oc. (Fin.) notoscriptus) for an example of state (0) and fig. 237 (Fl. bougainvillensis) for state (2). lu his fig. 211 (Ha. australis) for an example of state (1). 68(49). Seta 8-S, development: (0) short; (1) long. Seta Seta 1-X is inserted on the ventral or ventroposterior 8-S is normally short (state 0) in Aedini. It is notably margin of the saddle in Ge. baisasi, Oc. (Mac.) trem- long (state 1) in genus Finlaya (see illustrations in ulus and Oc. (Och.) muelleri, and is scored (0). Belkin, 1962; as Kochi Group), Oc. (Fin.) papuensis 74(53). Seta 3-X, development: (0) single; (1) branched. and the outgroup species Ma. titillans. Seta 3-X is normally single in Aedini (see Zavortink, 69(new). Seta 9-S, development: (0) absent; (1) short, 1972: fig. 9, Oc. (Pro.) triseriatus) but it is branched in slender, nearly straight or slightly curved; (2) stouter, Az. ramirezi, Gy. mediovittata, Ko. purpureipes, Hw. hooklike. Most Aedini have seta 9-S relatively short sexlineata (see Zavortink, 1972: figs 32, 37, 40 for the and nearly straight or gently curved (state 1) (see Bel- first three species) and the outgroup species Cs. inor- kin, 1962: fig. 234, Oc. (Fin.) argyronotum). Some nata and Ma. titillans (see Belkin et al., 1970: fig. 50). taxa have seta 9-S short and hooklike (state 2) (see 75(54). Scleroüzation supporting seta 4-X (ventral Belkin, 1962: fig. 274, Le. geoskusea). See explanation bush): (0) absent; (1) grid with only transverse bars; for Ma. titillans (state 0) under character 67. (2) grid with both lateral and transverse bars; (3) boss. Reinert (2002a) conducted a comparative analysis of 70(51). Sclerotization of segment X: (0) dorsal saddle the ventral brush and its attachment to segment X in incomplete ventrally; (1) saddle completely encircles Aedini. Sclerotization supporting the ventral brush segment X. All species of Aedini have either a small or was divided into four types and 18 subtypes. The large dorsal saddle or sclerotization that completely ancestral form of the ventral brush was hypothesized encircles segment X. A saddle completely encircling to have consisted of numerous, multiple-branched segment Xis found in Oc. (Ochierotatus) (some species setae with short, stout stems attached to a well-devel- groups), other Ochlerotatus (see illustrations in Arnell, oped grid consisting of both transverse and lateral 1976; Scapularis Group), Psorophora, the non-aedine grid bars. This brush also had numerous multiple- outgroup species, and some other aedirie outgroup branched, precratal setae with short stems inserted on species, e.g. Edwardsaedes imprimens and Huaedes the ventral midline of a complete saddle. The derived wauensis. Reinert (2002a) provided illustrations of dif- condition was described as being a brush consisting of ferent developmental states of this character in Aedini. a few, simple, single setae on a poorly developed grid 71(50). Saddle, acus: (0) absent; (1) present. Most spe- with only transverse grid bars. This brush lacked pre- cies examined in this study do not have a saddle acus cratal setae, and the saddle was incomplete ventrally. Reinert further proposed that the condition in Chris- (state 0). However, within genus Ochlerotatus an acus tophersiomyia of Aedes) is present (state 1) in Oc. communis (see Wood, Dang (as a subgenus with 6-8 moderately long, 2-7-branched setae with short stems & Ellis, 1979: p1. 20), Oc. infirmatus and Oc. scapu- inserted in an irregular row without both lateral and laris, Oc. (Rus.) refiki and Oc. (Rus.) rusticus. It is also present in Oc. (Fin.) albilabris, Oc. (Fin.) notoscrip. transverse grid bars could represent the most derived state because of the occurrence of a similar condition tus, Oc. (Psk.) bancroftianus and Oc. (Psk.) postspi- in most sabethine mosquitoes (see discussion in Har- raculosus. The acus is attached to the base of the saddle in these species except in Oc. (Fin.) albilabris bach & Kitching, 1998). The ventral brush in sabethines, however, is normally represented by a single and Oc. (Fin.) notoscriptus in which the acus is a small oval sclerite located in a similar position but pair of setae (species of subgenus Sabethinus of Sabe- thes detached from the base of the saddle. Orthopodoinyia have an auxiliary pair of setae 4-X). The well- developed boss of Oc. (Mol.) pecuniosus signifera has a narrow, basal, sclerotized band that is (state 3) is shown in Reinert (1993: fig. 8) and that of Mu. laniger separated from the saddle. Because of its structure and location, it is not considered to be homologous is illustrated in Tyson (1970: fig. 17). with the acus in Aedini. An acus is present in the out- 76(55). Precratallpreboss setae (i.e. two or more setae group species Cs. inornata. anterior to grid or boss): (0) absent; (1) present. Setae

Journal compilation © 2006 The Linnean Society of London, Zoological Journal of the Linnean Society, 2006, 148, 1-101 No claim to original US government works. PHYLOGENY AND CLASSIFICATION OF FINLAYA 11 of the ventral brush are normally paired but an occa- 80(58). Trumpet, tracheoid area: (0) absent; (1) present sional specimen may possess an odd number of setae. at base, weakly developed; (2) present distal to base, Some species, e.g. Hg. (Hag.) splendens, have a single well developed. The tracheoid area of the trumpet is unpaired preboss seta whereas other species have a well developed (state 2) in Finlaya, Mucidus, single unpaired precratal seta (these are scored as 0). Rhinoskusea and the non-aedine outgroup species Cx. Psorophora have numerous precratal setae that are quinquefasciatus and Ma. titillans (see Belkin et al., inserted in the ventral margin of the saddle (see Bel- 1970: figs 33, 51). The tracheoid area in most generic- kin et al., 1970: figs 56, 60, 67). level taxa of Aedini is weakly developed at the base of the trumpet (state 1), but it is absent (state 0) in out- Seta 4a-X of ventral brush, length: 77(56). (0) short; (1) group taxa, e.g. Gymnometopa, Op. (Opi.) fuscus, Pso- long. Reinert (2002a), following Belkin (1962) and rophora (most species), Oc. (Fin.) albotaentatus, other Knight & Laffoon (1971), defined seta 4-X as Any seta aedine taxa, e.g. Alanstonea, Armige res , Fredwardsius, of the ventral brush, the variable number of paired or Huaedes, Leptosornatomyia, Skusea, and Ae. (Adm.) unpaired setae arising from the ventral longitudinal alboscutellatus, as well as the non-aedine outgroup midline of abdominal segment X. The most caudal species Cs. inornata and Or signifera. (posterior) seta is designated as seta 4a-X, the next cephalad (anterior) seta as 4b-X, and so on. Seta 4a-X 81(59). Seta I-CT, development: (0) similar to seta 3- is normally long, rarely moderately long, and similar CT; (1) very strongly developed, considerably longer in length to seta 4c-X in Aedini. A hraedes , Aztecaedes, than seta 3-CT; (2) weakly developed, considerably Kompia, Oc. (Och.) muelleri and Oc. (Fin.) biocella- shorter than seta 3-CT. With few exceptions, the length tus have a short seta 4a-X (for first four taxa see and development of seta 1-CT is constant within well- Zavortink, 1972: figs 32, 34, 37, 44). defined generic-level taxa. See illustrations in Belkin (1962) for examples of state (0) (fig. 233, Oc. (Fin.) Seta 4a-X of ventral brush, development: 78(new). (0) argyronotum), state (1) (fig. 242, Fl. franclemonti) and single to 3-branched (occasionally 4-branched); (1) ^! 5- state (2) (fig. 266, Oc. (Och.) edgari). branched. See Belkin (1962: fig. 314, St. aegypti) for an example of state (0) and his fig. 316 (Sc. albolin- 82(60). Seta 5-CT, length: (0) :^ 1.2 times length of seta eata) for an example of state (1). 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- PUPAE CT (state 1) in other taxa, e.g. Kenknightia (see Rein- 79(57). Cephalothorax, with clear unpigme,ited spots: ert, 1990: figs 15-23). (0) absent; (1) present. Clear unpigmented areas of the Seta 7-CT, length: (0) !^ length of seta 6-CT; (1) cephalothorax (scutum and metanotum) (state 1), and 83(61). 1.2-5.0 times length of seta 6-CT; (2) ^ 6.0 times length also the abdominal terga in some species, of pupal exu- of seta 6-CT. The length of seta 7-CT is shorter than or viae are characteristic of Finlaya (see illustrations in equal to the length of seta 6-CT (state 0) in Stegomyia Belkin, 1962; as Kochi Group). Belkin (1962) reported (see Belkin, 1962: fig. 313, St. aegypti; Huang, 1990: that in living pupae of these mosquitoes, which breed africana). For examples of state (1), see Bel- almost entirely in water in plant axils, the unpig- fig. 10, St. kin (1962: figs 207,210, Op. (Opi.)fuscus and Ha. aus- mented areas are iridescent blue and are very similar to those found in pupae of many New World sabe- tralis) and for state (2) see Zavortink (1972: figs 60, 62, Hg. (Con.) leucocelaenus and Hg. (Con.) clarki). thines and species of Cx. (Microculex), which also inhabit water held in leaf axils. Mattingly (1969) also 84(62). Seta 11-CT, development: (0) single; (1) reported that Some South American sabethines and branched. See Belkin (1962: fig. 226, Oc. (Fin.) noto- culicine pupae, occurring in bromeliad axils, are pro- scriptus) fhr example of state (0) and his fig. 266 (Oc. vided with luminescent integumentary patches which (Och.) edgari) for state (1). A few species occasionally persist for some time after emergence of the adult. have seta 11-CT 2-forked distally and are scored (0). Berlins (1969) illustrations of the pupal metanotum of Seta 13-CT (0) absent; (1) present. Among Hw. inaequalis and Hw. stenei show an unpigmented 85(63). Ab. papago lateral spot. Because no specimens of these two spe- three paratypes of examined, a well- developed, 2-branched seta is present on both sides of cies were available for examination, it is uncertain if one exuviae, an alveolus is present on both sides of these spots are similar to those reported above. Alan- another, and the seta and alveolus are absent on the stonea brevitibia and Belkinius aurotaeniatus have similar clear unpigmented spots. A small clear spot third. Reinert (1980, 2000a) provided a review of the occurrence of seta 13-CT in Culicidae and Aedini. dorsal to the trumpet base was observed in the two available specimens ofOc. (Fin.) loi, hence this char- 86(new). Seta 34, length: (0) < length of seta 64; acter is scored (1) for this species. (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. (state 0) (see Belkin, 1962: fig. 233, Oc. (Fin.) argy- Oc. scapularis (see Arnell, 1976: fig. 36). For examples ronotum) in most Aedini. See Belkin (1962: fig. 207, of state (1), in which seta 3-I is longer than seta 6- Op. (Opi.)fuscus) for an example of state (1). This seta I, see Schick (1970: figs 15, 31, Oc. (Pro.) terrens and is normally lateral to seta 4-I1, rarely positioned Oc. (Pro.) homoeopus). mesally, in Oc. (Pro.) hendersoni. - 87(new). Seta 34, development: (0) single, rarely split 94(new). Seta 5-I1, length: (0) :^ length of seta 3-I1; apically; (1) branched. Seta 3-I is often single (rarely (1)> length of seta 3-I1. Most taxa examined have seta with apex split) (state 0) in numerous taxa, e.g. 541 shorter than or equal to the length of seta 3-I1 Abraedes, Aedes, Aztecaedes, Downsiomyia (see (state 0), e.g. Haemagogus (see Arnell, 1973: figs 34, Tanaka, 2002: fig. 8, Do. nipponica), Finlaya and Oc. 39). Examples of taxa with seta 5-1I longer than seta (Protomacleaya). See Belkin (1962: figs 210, 217, Ha. 341 are Oc. (Fin.) albilabris and usually Oc. (Fin.) australis and Ge. longiforceps) for examples of state (1). notoscriptus (see Belkin, 1962: figs 226, 230). 88(64). Seta 64, length: (0) !^ length of seta 74; 95(71). Seta 6-I1, length: (0) !^ length of seta 7-1I; (1) (1)> length of seta 74. The length of seta 6-I is shorter > length of seta 741. Seta 641 is normally longer than than or equal to the length of seta 7-I (state 0) in some seta 741 (state 1) in Aedini (see Belkin, 1962: fig. 230, taxa, e.g. Op. (Opi.) fuscus and Ha. australis (see Be!- Oc. (Fin.) albilabris), however, it is shorter (state 0) kin, 1962: figs 207, 210). Examples of taxa with seta (see Belkin, 1962: fig. 207, Op. (Opi.) fuscus) in some 6-I longer than seta 7-I (state 1) are Ps. (Jan.) ferox generic-level taxa and species groups (see Appendix 2). (see Belkin et al., 1970: fig. 59) and Za. longipalpis (see Reinert, 1999b: fig. 11). 96(69). Seta 3-Ill, development: (0) single; (1) branched. Most taxa examined have seta 3-I11 single, 89(65). Seta 2-I1, position: (0) insertion mesal to or at long and stout (state 0) (see Belkin, 1962: fig. 230, Oc. same level as seta 141; (1) insertion lateral to seta 1- (Fin.) albilabris). Seta 3-I11 is branched and normally IT. Seta 24I is normally lateral to seta 141 (state 1) in slender (state 1) in several taxa, e.g. Isoaedes, Kenk- Aedini, however, it is mesal to or at the same level nightia, Opifex and Rhinoskusea (see Reinert, 1976b: as seta 141 (state 0) in Finlaya, Oc. (Fin.) chungi figs 9-11, for Rhinoskusea). and Oc. (Mol.) pecuniosus (see Reinert, 1993: fig. 7). 97(72). Seta 6-I11, development: (0) single; (1) 90(66). Seta 2-I1, length: (0) < length of seta 14I; branched. The development of seta 6-I11 is constant for (1) ^! length of seta 1-I!. Seta 2-I1 is normally shorter generic-level taxa, with few exceptions. However, in than seta 1-1I in Aedini, however, it is as long or longer some specimens of some species, e.g. Ke. harbachi, and often stouter than seta 141 (state 1) in some taxa, Oc. (Fin.) hanksi and Oc. (Pro.) berlini, this seta usu- e.g. Rhinoskusea, Scutomyia and Ha. australis. ally exhibits one state but occasionally exhibits the other state, e.g. normally single but sometimes 2- 91(67). Seta 3-I1, position: (0) insertion mesal to or at same level as seta 2-1I; (1) insertion lateral to seta 2- branched, these are scored (0,1). II. The position of seta 341 in relation to seta 2-I1 is 98(74). Seta 5-V, length: (0) < medial length of tergum constant for most generic-level taxa, with few excep- VI; (1) ^! medial length of tergum VT. Seta 5-V is often tions. However, in Oc. (Finlaya) and Oc. (Protoma- single (rarely 2-branched), very long, stout and longer cleaya) this character appears to be constant at the than the tergum of the following segment, but in some species-group level. Examples of state (0) are found in taxa it is branched, short to moderately long, slender Aztecaedes and Kompia (see Zavortink, 1972: figs 31, and shorter than tergum VI. See Belkin (1962) for a 36), and state (1) in Abraedes and Zavortinkius (see comparison of the two character states (fig. 313, Zavortink, 1972: fig. 33, Ab. papago; Reinert, 1999b: St. aegypti, state 0; fig. 207, Op. (Opi.) fuscus, state 1). fig. 11,Za. longipalpis). 99(75). Seta 2-VI, position: (0) insertion mesal to or at 92(68). Seta 3-I1, length: (0) !^ length of seta 6-I1; same level as seta 1-VI; (1) insertion lateral to seta 1- (1)> length of seta 6-I1. State (0) i g found in a number VI. The position of seta 2-VT, in relation to seta 1-VI, is of aedine taxa, e.g. Aedes, Geoskusea, lsoaedes and normally consistent within most well-defined generic- Levua. Numerous taxa have seta 34I longer than seta level taxa. See examples in Belkin (1962: fig. 233, Oc. 641 (state 1), e.g. Abraedes, Aztecaedes, Gymnometopa (Fin.) argyronotum, state 0; fig. 210, Ha. australis, and Kenknightia. Seta 341 is normally slender when it state 1). is short to moderately long and stout when it is long. 100(76). Seta 3-VI, position: (0) insertion mesal to or 93(70). Seta 5-1I, position: (0) insertion lateral to or at at same level as seta 1-VT; (1) insertion lateral to seta same level as seta 44I; (1) insertion mesal to seta 4-I1. 1-VT. Seta 3-VI is normally lateral to seta 1-VT (state Seta 541 is longer and normally lateral to seta 441 1) in Aedini. However, it is positioned at the same

Journal compilation © 2006 The Linnean Society of London, Zoological ,Journal of the L,innean Society, 2006, 148, 1-101 No claim to original US government works. PHYLOGENY AND CLASSIFICATION OF FINLAYA 13

level or mesal to seta 1-VI (state 0) in Finlaya, Down- 106(81, in part). Paddle; apical margin: (0) sharply or siomyia, Oc. (Cha.) wattensis, Oc. (Fin.) albilabris, broadly rounded or flat; (1) with moderate to deep Oc. (Fin.) argenteitarsis, Oc. (Fin.) assamensis, Oc. emargination at or near apex of midrib. Most Aedini (Fin.) loi, Oc. (Fin.) subalbitarsis, Oc. (Mac.) tremu- have the apical margin of the paddle rounded, but lus, Op. (Opi.) fuscus (see Belkin, 1962: fig. 207) and some have this area flattened or with a slight emar- the non-aedine outgroup species Cs. inornata and gination, e.g. Oc. (Och.) deserticola and Oc. (Och.) Cx. quinquefasciatus. varipalpus, and are scored (0). A few taxa have a deeper emargination on the apical margin (state 1), 101(73). Seta 6-Vu, position: (0) insertion anterior to e.g. Abraedes, A.ztecaedes, Kompia, Fl. kochi, Ha. aus- seta 9-VII; (1) insertion posterior to or at some level as go tralis, Hw. walkeri, Oc. (Och.) atropalpus and Oc. seta 9-VII. Seta 6-VII is inserted posterior (occasion- (Och.) epactius (see Zavortink, 1972: fig. 46). ally at the same level) to seta 9-Vu (state 1) in most Aedini, e.g. Oc. (Protomacleaya) (see illustrations in 107(new). Seta 1-Pa, length: (0) short to moderately Schick, 1970 and Zavortink, 1972). Seta 641 is ante- long, 0.33 length of paddle; (1) long, ^! 0.40 length of rior to seta 9-VU (state 0) in some species examined, paddle. Downsiomyia and Finlaya are examples of state e.g. Psorophora (see Belkin et al., 1970: figs 57, 59, 66), (0) (see illustrations of Finlaya , as Kochi Group, in Bel- Oc. (Bruceharrisonius), Oc. (Fin.) gilli, Oc. (Fin.) loi kin, 1962). Levua geoskusea and Oc. (Brh.)greenit are and Za. fulgens. examples of state (1) (see Reinert, 2003: fig. 2E). 102(77). Seta 9-Vu, length: (0) !^ length of seta 6-VII; (1) 108(82, in part). Seta 1-Pa, development: (0) absent; (1) > length of seta 6-VU. Seta 9-VII is normally longer and present, single, rarely 2-branched or 2-forked; (2) often stouter than seta 6-VII (state 1) in Aedini, but it present, ^! 3 branches. Seta 1-Pa is absent (state 0) in is occasionally shorter than or equal to the length of Ma. titillans (see Belkin et al., 1970: fig. 51). See Rein- seta 6-VII (state 0). Seta 6-VII does not occur in pupae ert (1990: figs 15-23, Kenknightia) for state (1) and of Ma. titillans, nor in Ma. dyari and Ma. flaveola, Reinert (1976b: figs 9-11, Rhinoskusea) for state (2). therefore this character could not be scored for this spe- Some species show an overlap between the two char- cies and is entered as missing (—) in the data matrix. acter states, e.g. Do. leonis (2 or 3 branches), Ps. (Jan.) ferox (2 or 3 branches) and Cs. inornata (2 or 3 103(78). Seta 9-VIII, development: (0) single or 2- branches), and are scored (1,2). However, most speci- branched; (1) _> 3 branches. Many aedine taxa have mens of these species have three or more branches. seta 9-VIII multiple-branched, often aciculate and often strongly developed (see Belkin, 1962: fig. 233, 109(83). Seta 2-Pa: (0) absent; (1) present. Absence of Oc. (Fin.) argyronotum). See Reinert (1993: fig. 7, Oc. seta 2-Pa (state 0) is the usual condition in Aedini, how- Grab- (Mol.) pecuniosus) for example of state (0). ever, seta 2-Pa is present (state 1) in subgenera hamia (except Ps. infinis) and Psorophora of 104(79). Paddle, midrib: (0) weakly developed, not Psorophora (see Belkin et al., 1970: figs 57, 64-66), reaching apex of paddle; (1) well developed, extending Ta. savoryi (rarely absent on one side of a few speci- to or near apex of paddle. Most generic-level taxa of mens) (see Tanaka, 2002: fig. 3) and the outgroup spe- Aedini have the midrib well developed and extending cies Cx. quinquefasciatus. One of six pupal exuviae of to or near the apex of the paddle (state 1). The midrib, Oc. (Och.)fluviatilis examined has the apex of one pad- however, is weakly developed to nearly indistinct in dle with two setae and the other with two setae and a some taxa, e.g. Udaya and Zeugnomyia. The midrib is setal alveolus, which was considered an anomaly. shortened and often restricted to about the basal 0.60 of the paddle (state 0) in other taxa, e.g. Abraedes, ADULTS (FEMALES EXCEPT WHERE OTHERWISE NOTED) Aztecaedes, Gymnometopa, Kompia, as well as 110(84). Erect scales of head: (0) absent; (1) restricted deserticola, Oc. muelleri, Oc. sierrensis and Oc. van- to occiput; (2) on occiput and vertex. See Harbach & palpus (see Arnell & Nielsen, 1972: fig. 8) of subgenus Kitching(1998) for examples of state (1) (their fig. 6B) Ochlerotatus, these are scored (0). and state (2) (their fig. 6A). Erect scales in Aedini are either restricted to the occiput or borne on both the Paddle, fringe of hairlike spicules: (0) absent; 105(80). occiput and vertex. In some species, several semierect, (1) present. Reinert (2000a) listed the generic-level narrow, forked scales similar to erect scales on the taxa of Aedini with a fringe of hairlike spicules on the occiput occur on the vertex (often posterior to the ocu- pupal paddle. These are present or absent in generic- lar setae), and these are scored (2). Erect scales are level taxa, with few exceptions. Most species of Stego- absent in Op. (Opi.) fuscus, however, numerous short, myia have well-developed hairlike spicules on the pad- curved, dark setae occur on the occiput and vertex. dle margin, but there are notable exceptions, e.g. St. aegypti and the Dendrophilus Group (see illustra- 111(85). Decumbent scales of vertex: (0) broad; (1) nar- tions in Huang, 1997). row; (2) both broad and narrow. Harbach & Knight

(1980) illustrated examples of scales that are narrow interocular space occur in Ahraedes, Downsiomyia, and broad. In species with both broad and narrow Gymnometopa, Haemagogus, Kenknightia, Kompia, scales on the vertex (state 2), the narrow scales are Stegomyia, Zavortinkius, Ps. (Jan.) ferox and five spe- usually on the median area and may be restricted to a cies of Oc. (Finlaya). Opifex (Opi.) fuscus has only a double row along the coronal suture (see Tanaka et al., few scattered small broad scales on this area. Both 1979: fig. 228, Oc. (Fin.) oreophilus). In Scutomyia, a narrow and broad scales (state 3) occur on the interoc- patch of narrow scales occurs on the anterior median ular space ofOc. (Cha.) wattensis. area (state 2). Examples of state (0) are shown in Tanaka et al. (1979: fig. 222, Oc. (Fin.) seoulensis) and 116(89). Interocular space, setae: (0) absent; (1) state (1) in their fig. 221 (Ta. savoryi). present. The interocular space is defined here as the area between the compound eyes extending dorsad 112(86). Ocular line, width: (0) narrow; (1) broad. The from the postfrontal suture to a point level with the dor- ocular line is interpreted here as the area anterior to sal margins of the eyes. Setae are normally present on the ocular setae and posterior to the dorsal margin of this area in Aedini and the outgroup species. They are the eye. Examples of a narrow ocular line (state 0) are absent in Gymnometopa and Kompia. They are also found in Tanaka el al. (1979: figs 220, 222, Th. togoi absent in the aedine genera Ayurakitia and Udaya. and Oc. (Fin.) seoulensis) and a broad one (state 1) in Reinert (1990: fig. 3, Ke. harbachi). The ocular line of 117(91). Antennal pedicel, mesal surface: (0) with few Cs. inornata is moderate in width, and is here scored as to numerous scattered scales (not overlapping to only (0). See discussion of scale types under character 111. slightly overlapping and not silvery) and/or setae; (1) with patch of broad, overlapping, silvery scales; (2) 113(87, in part). Ocular scales: (0) narrow; (1) broad; bare. State (0) occurs in most Aedini and the outgroup (2) both narrow and broad. Most species have either species. State (1) occurs in Ahraedes,Aztecaedes, Gym- narrow or broad scales on the ocular line but a few nometopa and Kompia, Oc. (Fin.) albilabris and Oc. have both narrow and broad scales (state 2), e.g. (Fin.) notoscriptus, as well as Albuginosus, Bothaella, Howardina, Mu. laniger and Oc. communis. See dis- Scutomyia, Stegomyia and Zeugnomyia. In state (0), cussion of scale types under character 111. the mesal surface of the pedicel has few to numerous, 114(88). Eyes, immediately above antennal pedicels: (0) small to large, scattered to slightly overlapping, pale contiguous; (1) narrowly to moderately separated; (2) (white, cream-coloured, golden brown, etc.) or dark broadly to very broadly separated. The distance sepa- scales and/or setae (see Tanaka et al., 1979: fig. 222, rating the eyes is determined on the area immediately Oc. (Fin.) seoulensis), whereas in state (1) a moder- above the antennal pedicels and is measured in number ately large to large patch of tightly overlapping, of eye facets (diameter). Species with eyes that touch or silvery scales is present (see Tanaka et al., 1979: are separated by less than or equal to one eye facet are fig. 234, St. aegypti). The mesal surface of Oc. (Och.) deserticola considered contiguous and are scored (0), e.g. Oc. (Fin.) has a moderate-sized patch of broad, over- assamensis, those separated by 2-4 (rarely 5) facets are lapping, white scales above and a few scattered, small, scored (1), e.g. Oc. (Fin.) geniculatus, and those sep- white scales below, and is scored (0). State (2) occurs in arated by 6 or more facets are scored (2), e.g. Op. (Opi.) two Afrotropical species examined, Oc. (Fin.) luteo- fuscus. Scales protruding from the median, anterior striatus and Oc. (Fin.) pulchrithorax. area of the vertex or upper part of the interocular space 118(92). Antennal pedicel, lateral surface: (0) bare; (1) may obscure this area, and in some cases, e.g. Howar- with scales. The outer surface of the antenna] pedicel dma, the scales must be removed to determine the dis- is bare (state 0) (see Tanaka et al., 1979: fig. 228, Oc. tance between the eyes. Gutsevich (1974a, b, 1975a, b) (Fin.) oreophilus) in most Aedini. Scales are present used the diameter of eye facets compared to the distance on this area (state 1) in Oc. (Rusticoidus), Ochlerotatus between the eyes as a means of distinguishing several (some species) (see Tanaka et al., 1979: fig. 212, genera and subgenera, however, he used a different cri- Oc. dorsalis) and Oc. (Ochlerotatus) (some species), terion than the number of facets used here. Stegomyia and the outgroup species Ma. titillans. 115(90). Interocular space, scales: (0) absent; (1) nar- 119(93). Apical flagellomeres (males): (0) both apical row; (2) broad; (3) both narrow and broad. In Geosku- fiagellomeres disproportionately long compared with sea, Levua and Rhinoskusea, the interocular space is other fiagellomeres; (1) these fiagellomeres not dispro- reduced to a very small triangle and is apparently portionately long in comparison with others. Antennae without scales (state 0). In these subgenera, the eyes with the two distal fiagellomeres approximately equal are contiguous for nearly the entire mesal length, in length and much longer than the proximal flagello- resulting in a very small interocular space. Most taxa meres is the usual condition in Aedini (state 0) (see examined in the present study have narrow scales on Tanaka et al., 1979: fig. 228, On. (Fin.) oreophilus). this area (state 1). Broad scales (state 2) on the However, the two apical flagellomeres are approxi-

Journal compilation © 2006 The Linnean Society of London, Zoological Journal of the Linnean Society, 2006, 148, 1-101 No claim to original US government works. PHYLOGENY AND CLASSIFICATION OF FINLAYA 15 mately the same length as the other flagellomeres in pomere 4 (males): (0) absent or few; (1) numerous. both subgenera of Opifex (see Belkin, 1968: figs 2, 19). Many aedines have numerous long setae on the lateroventral or ventral margins distally on palpomere 3 120(94). Antenna, development of flagellar whorls and the entire length of palpomere 4 (state 1) (see (males): (0) few short setae, dispersed more or less Tanaka et al., 1979: figs 213, 215, Oc. (Och.) excru- around flagellomeres; (1) several moderately long to cians and Oc. (Och.) sticticus), whereas other species long setae, directed more or less dorsally and ven- have no or few setae on the distal part of palpomere 3 trally; (2) numerous long setae, normally directed dor- and few to several on palpomere 4 (state 0) (see sally and ventrally. Opifex is notable in having only Tanaka et al., 1979: figs 220, 221, Ta. togoi and a few short setae on each flagellomere (state 0) Th. sauoryi). Species with numerous long setae on the (see Belkin, 1968: figs 2, 19, Op. (Opi.) fuscus and distal part of palpomere 3 often also have the distal Op. (Not.) chaihamicus). Levua, Rhinoskusea and part of the palpomere somewhat swollen. Ge. longiforcep.s have several setae that are moderately long to long and normally directed dorsally and 125(new). Proboscis, pale scales: (0) absent; (1) ventrally (state 1) (see Reinert, 1976b: fig. 2, Rh. Ion- present. The proboscis may be entirely dark-scaled girostris). Antennal whorls comprising numerous long (state 0) (see Russell, 1996: 81, Oc. (Fin.) rubrithorax) setae normally directed dorsally and ventrally (state or have pale scales intermixed with or forming bands 2) (see Tanaka et al., 1979: fig. 222, Oc. (Fin.) seou- or patches in the dark-scaled areas (state 1) (see lensis) are usually present in Aedini. Russell, 1996: 61, 73, Fl. kochi and Oc. (Fin.) notoscriptus). 121(new). Maxillary palpus,pale scales: (0) absent; (1) present. Maxillary palpi may be entirely dark-scaled 126(96). Antepronota: (0) approximated; (1) widely (state 0) or have pale scales intermixed with or form- separated. The antepronota are nearly touching dor- ing bands or patches in the dark-scaled areas (state 1). sally (state 0) in Hg. (Hag.) equinus and Hg. (Hag.) splenden.s, and in some other taxa studied previously 122(95, in part). Maxillary palpomeres (males): (0) five, (Reinert et al., 2004), whereas they are widely sepa- palpomeres 2 and 3 fusedlankylosed; (1) three, fourth rated (state 1) in most Aedini. absent or vestigial. Palpomeres 2 and 3 exhibiting state (0) have a paler, somewhat distinct ring where they are 127(97, in part). Anterior acrostichal setae: (0) absent; fusedlankylosed whereas no such area is apparent (1) present. Anterior acrostichal setae are defined as when palpomeres exhibit state (1). Very short palpi are those on the acrostichal area cephalad of the scutal found in males of the following Aedini: Aedes, Ae. (Can- angle whereas those caudad of this point are the pos- craedes), Belkinius, Bothaella, Christophersiomyia, terior acrostichal setae. Setae on the anterior promon- Geoskusea (most species), Haemagogus (few species), tory (the broad median area of the mesonotum at the Heizmannia, Huaedes, Levua, Leptosomatomyia, anterior end of the acrostichal area that projects more Opifex (Nothoskusea), Paraedes, Rhinoskusea, Verral- or less cephalad over the cervix) should not be con- lina and Zeugnomyia (Reinert, 2000a). There is no way fused with the presence or absence of setae on the of knowing whether palpomere 2 of males with short anterior end of the acrostichal area. Arnell (1973: 8) palpi actually consists of palpomeres 2 and 3 that are reported the presence of minute dorsocentral and completely and unrecognizably fused. The number of acrostichal setae (visible only in slide preparations) in palpomeres is consistent for generic-level taxa, with Haemagogus. These setae were not seen in pinned few exceptions, e.g. Ge. baisasi and Ge. longiforceps. specimens of Hg. equinus and Hg. splendens examined, and are scored as (0). 123(new). Maxillary palpus, length (males): (0) Posterior acrostichal setae: short, :^ 0.25 length of proboscis; (1) moderate, 0.48- 128(97, in part). (0) absent; 0.80 length of proboscis; (2) long, ^! 0.84 length of pro- (1) present. See discussion under character 127. boscis. The maxillary palpus is noticeably short in 129(98, in part). Anterior dorsocentral setae: (0) males of some aedine taxa (state 0), e.g. Levua and absent; (1) present. The dorsocentral area is defined as Rhinoskusea (see Reinert, 1976b: fig. 2, Rh. longiros- the longitudinal area on either side of the acrostichal tris), moderately long (state 1) in some, e.g. Scutomyia area extending caudally from the antedorsocentral albolineata and Sc. arhoricola (see Huang, 1979: fig. 6) area (see Reinert, 1999b) to the prescutellar area. and long, often as long as or longer than the proboscis Setae on the area cephalad of the juncture of the pre- (state 2) in others (see Tanaka et al., 1979: figs 212, scutal suture are referred to as anterior dorsocentral 213, 222, Oc. dorsalis, Oc. (Och.) excrucians and Oc. setae and those caudad as posterior dorsocentral (Fin.) seoulensis). setae. Also see discussion under character 127. 124(new). Maxillary palpus, long lateroventral setae 130(98, in part). Posterior dorsocentral setae: (0) absent; on distal part of palpomere 3 and entire length of pal- (1) present. See discussion under character 129.

131(new). Acrostichal and dorsocentral areas, anterior dark-scaled (state 0) as in Ge. baisasi (see Tanaka 0.30-0.70 covered with pale scales: (0) absent; (1) et al., 1979: fig. 239), have pale scales in a large patch present. Females of some species have the combined or be completely pale-scaled (state 1) as in Oc. (Fin.) acrostichal and dorsocentral areas dark-scaled (state seoulensis and Do. nishikawai (see Tanaka et al., 1979: 0), e.g. Ge. baisasi (see Tanaka et al., 1979: fig. 239) and figs 222, 227) or bear small pale patches or narrow Ke. dissimilis (see Reinert, 1990: fig. 1) whereas other pale-scaled lines on the lateral and/or mesal and/or species have this area, and often the entire anterior posterior margins (state 2) (see Tanaka et al., 1979: 0.30-0.70 (acrostichal, dorsocentral and scutal fossal fig. 221, Ta. savoryi). Pale-scaled areas in states (1) areas) of the scutum, covered with pale (normally white and (2) contrast (normally white or silvery) with back- or silvery) scales (state 1) (see Barraud, 1923: figs 13, ground scales that are normally dark but may be a 17-19, Oc. (Fin.) deccanus, Oc. (Fin.) cacharanus, Oc. darker shade or colour of pale scales in some species, (Fin.) lophoventralis and Oc. (Fin.) cogilli). e.g. Oc. (Fin.) pulchri venter and Oc. (Fin.) tubbutiensis. The scutal fossa of Oc. communis normally has a 132(new). Anterior acrostichal area, with narrow, pale-scaled margin with contrasting darker scales on pale-scaled stripe: (0) absent; (1) present. When the median area (see Carpenter & LaCasse, 1955: p1. present (state 1), the pale-scaled stripe normally con- 54), however, some specimens have the scutal fossa sists of narrow scales; but these are broad in some spe- nearly covered with pale scales and only a few darker cies, e.g. Oc. (Mol.) pecuniosus (see Reinert, 1993: scales medially and is therefore scored (1,2). Some fig. 3), Za. fulgens and Za. longipalpis (see Reinert, species, e.g. Ps. (Jan.) ferox (see Carpenter & LaCasse, 1999b: fig. 1). The pale-scaled stripe is normally nar- 1955: p1. 35), have the scutal fossa covered with inter- row as in Oc. (Fin.) oreophilus but may be moderate mixed broad pale and dark scales in no defined pat- in width and slightly separated by the acrostichal tern, scored here as state (2). setae as in Ta. togoi (see Tanaka et al., 1979: figs 228, 220, respectively). In St. aegypti, a narrow white- 137(99). Prescutellar area, median and/or posterior scaled stripe occurs on either side of the scutum on parts: (0) bare; (1) with scales and/or setae. The the outer edge of the acrostichal area at the interface median part of the prescutellar area is bare (state 0) in with the dorsocentral area (see Tanaka et al., 1979: most aedine species. This area in Oc. (Fin.) jugraensis fig. 234). Belkin et al. (1970) referred to this stripe as (see Knight, 1968: fig. 1), Oc. (Fin.) rizali and Oc. the outer acrostichal line. It is included here as an (Fm.) sherki has a median, longitudinal, narrow, pale- acrostichal stripe (state 1). In female paratypes ofOc. scaled stripe, and the scales mesad of the prescutellar (Fin.) candidoscutellum, the acrostichal, dorsocentral, setae are dark. Some other taxa have state (1), e.g. scutal fossal, prescutellar and supraalar areas have Aztecaedes, Hg. (Haemagogus), Mucidus, Oc. (Rusti- indistinct, narrow, stripes of whitish scales on a back- coidus), Oc. (Ochlerotatus) (few species), Psorophora ground of golden-brown scales. These are scored as (some), Rhinoskusea , Hw. walkeri and Za. longipalpis. present (1). See discussion under character 127. Finlaya have scales on the anterior area and/or in transverse bands on the posterior area. 133(new). Posterior acrostichal area, with narrow, pale-scaled stripe: (0) absent; (1) present. See discus- 138(100). Prescutellar setae: (0) absent; (1) present. sion under character 127. Prescutellar setae are normally present in Aedini, but are absent in Hg. (Haemagogus) as well as in Alan- 134(new). Anterior dorsocentral area, with narrow, stonea brevitibia, Pseudarmigeres argenteoventralis pale-scaled stripe: (0) absent; (1) present. A stripe nor- dunni, Hz. (Heizmannia) and Hz. (Mattinglyia) acha- mally extends over the central part of the dorsocentral etae (sometimes present in the last species). area (see Tanaka et al., 1979: fig. 217, Oc. (Fin.) japonicus), however, it may be on the inner dorsocen- 139(new). Prescutellar area, pale scales on outer mar- tral area (see Berlin, 1969: figs 8, 9) as in Ab. papago gin inesal to setae: (0) absent; (1) present. Many aedine and Hw. walkeri. See discussion of character 132 for species have a narrow to moderately wide stripe of Oc. (Fin.) candidoscutellum. Also see discussion pale scales mesal to the prescutellar setae (state 1) under character 129. (see Tanaka et al., 1979: figs 217, 219, 220, Oc. (Fin.) japonicus, Oc. (Fin.) hatorii and Th. togoi) but other 135(new). Posterior dorsocentral area, with narrow, species have dark scales on this area (state 0) (see pale-scaled stripe: (0) absent; (1) present. See discus- Tanaka et al., 1979: fig. 229, Oc. (Fin.) watasei). sion under character 129. 140(new). Antealar area, anterior part, scales: (0) 136(new). Scutal fossa, scales: (0) dark; (1) contrasting absent; (1) present, dark; (2) present, pale. This char- pale scales in large patch; (2) contrasting pale-scaled acter refers to the part of the antealar area anterior to lines or small patches on lateral and/or mesal and/or the forward edge of the paratergite. This area may posterior margins. The scutal fossa may be entirely lack scales (state 0) (see Zavortink, 1972: fig. 35, Az.

Journal compilation © 2006 The Linnean Society of London, Zoological Journal of the Linnean Society, 2006, 148, 1-101 No claim to original US government works. PHYLOGENY AND CLASSIFICATION OF FINLAYA 17 ramirezi) or bear dark scales (state 1) (see Tanaka 147(106). Postpronotal scales: (0) absent; (1) present. et al., 1979: fig. 239, Ge. baisasi) or pale scales (state Broad, narrow or both broad and narrow scales are 2) (see Tanaka et al., 1979: fig. 222, Oc. (Fin.) normally present on the postpronotum. They are seoulensis). absent from this structure in Rhinoskusea (see Rein- ert, 1976b: fig. 2, Rh. longirostris), Hw. walkeri, Oc. Supraalar area, scales: 141(new). (0) all dark; (1) with (Fin.) candidoscutellum and Ps. (Pso.) ciliata. longitudinal patch or stripe of pale scales; (2) with transverse patch of pale scales. The supraalar area 148(107). Prespiracular setae: (0) absent; (1) present. may be entirely dark-scaled (state 0) (see Tanaka Setae are normally absent from the prespiracular area et al., 1979: fig. 227, Do. nishikawai), bear a longitu- in aedines, however they are present in Psorophora dinal pale-scaled patch (state 1) (see Tanaka et al., and the outgroup species Cs. inornata. Oc. (Fin.) japonicus) or have a trans- 1979: fig. 217, 149(108). Postspiracular setae: (0) absent; (1) present. verse pale-scaled patch (state 2) (see Tanaka et al., Postspiracular setae are present in most Aedini but are 1979: fig. 229, Oc. (Fin.) watasei; Barraud, 1923: absent in some taxa (see Reinert et al., 2004). They dcc- figs 11-13, 17, Oc. (Fin.) gubernatoris, Oc. (Fin.) are absent in Ko. purpureipes and the outgroup species canus and Oc. (Fin.) cacharanus). Cs. inornata, Cx. quinquefa.sciatus and Or signifera. Scutellum, scales on midlohe: (0) narrow; (1) 142(101). 150(109). Postspiracular scales: (0) absent; (1) present. broad; (2) both narrow and broad. Scales on the mid- Presence (state 1) (see Tanaka ci al., 1979: fig. 220, lobe (as well as lateral lobes) of the scutellum may be Th. togoi) or absence (state 0) (see Arnell, 1976: fig. 35, ci al., all narrow (state 0) (see Tanaka 1979: Oc .scapularis) of scales on the postspiracular area is fig. 219, Oc. (Fin.) hatorii), all broad (state 1) (see normally constant for generic- and species-group taxa 1)0. nishikawai) or both Tanaka et al., 1979: fig. 227, in Aedini. narrow and broad (state 2) (see Knight, 1968: fig. 1, Oc. (Fin.) jugraensis). Mucidus alternans and Mu. 151(110). Hypostigmal scales: (0) absent; (1) present. langer have the lobes bearing several narrow and a Scales on the hypostigmal area should not be confused few moderately broad, decumbent, falcate scales and with those on the dorsal part of the subspiracular area numerous long, twisted, erect scales (scored 0). Long, when two patches are present on the latter structure, twisted, erect scales are also present on other body one immediately dorsal to the other. Scales are nor- areas, e.g. head, thorax, abdomen, etc. (see illustra- mally absent on the hypostigmal area of Aedini (state tions in Tyson, 1970). See discussion of scale types 0) (see Zavortink, 1972: fig. 5, Oc. (Pro.) brelandi). A under character 111. small patch of scales is present in some species (state 1) (see Zavortink, 1972: fig. 35, Az. ramirezi). 143(102). Scutellum, scales on lateral lobes: (0) narrow; (1) broad; (2) both narrow and broad. See discus- 152(111). Subspiracular area: (0) bare; (1) with scales sion under characters 111 and 142. and/or setae. Presence or absence of scales on the subspiracular area is normally constant for genera and 144(103). Paratergal scales: (0) absent; (1) present. species groups of Aedini, with few exceptions. Scales Generic-level taxa normally either have paratergal and setae are present on this area, e.g. Oc. (Och.) scales or lack them, with few exceptions. The varipalpus (see Arnell & Nielsen, 1972: fig. 7) and Ps. Oc. (Fin.) paratergite may be bare (state 0) as in (Pso.) ciliata. Also see discussion under character 151. japonicusjaponicus and Ge. baisasi (see Tanaka ci al., 1979: figs 217, 239) or have several to numerous scales 153(112). Upper proepisternal setae: (0) 1-4; (1) 5-19; (state 1) as in Oc. (Fin.) seoulensis and St. aegypti (2) ^20. Some specimens of a few species examined (see Tanaka ci al., 1979: figs 222, 234). have a slight overlap (usually of one seta) of the states, e.g. Oc. (Fin.) assamensis, Oc. (Fin.) chrysolineatus, Parascutellar scales: (0) absent; (1) present. 145(104). Ta. savoryi, Ge. longiforceps, Ia. cavaticus and Ko. Parascutellar scales in Aedini were first noted in spe- purpureipes, whereas most specimens of these species cies of Oc. (Rusticoidus) (Reinert, 1999a) and a few have setal numbers of a single character state. species of Oc. (Ochlerotatus). Upper proepisternum, scales: (0) absent; (1) Antepronotal scales: (0) absent; (1) present. 154(113). 146(105). present. Aedine species normally have scales on the Scales are present on the antepronotum (state 1) (see upper proepisternum (state 1) (see Tanaka et al., Arnell, 1973: fig. 38, Hg. (Hag.) splendens) in the 1979: fig. 218, Oc. (Fin.) japonicus). These scales are majority of species examined in the present study and absent in Geoskusea, Levua and Rhinoskusea may be narrow, broad or both narrow and broad. (see Reinert, 1976b: fig. 2, Rh. longirostris). Scales are absent (state 0) in Geoskusea (see Tanaka ci al., 1979: fig. 239, Ge. baisasi), Levua, Rhinoskusea, 155(114). Lower proepisternum, scales: (0) absent; (1) Oc. (Fin.) candidoscutellum and Ps. (Pso.) ciliata. present. When scales are present on the lower proepi-

Journal compilation © 2006 The Linnean Society of London, Zoological Journal of the Linnean Society, 2006, 148, 1-101 No claim to original US government works. 18 J. F. REINERT ET AL. sternum they usually cover all or much of the surface 162(121). Mesepimeron, fine setae on ventral area: (0) and are normally broad and silvery or white. Scales absent; (1) present. Setae are absent in all taxa exam- are present in the following taxa of the present study: ined in the present study except Geoskusea (see Mat- Abraedes, Haemagogus, Kompia, Scutomyia, Stego- tingly, 1959: fig. 2C, Ge. kabaenensis). See Reinert myia, Zavorttnkius, Oc. communis, Oc. infirmatus, Oc. et al. (2004) for other aedine taxa with state (1). scapularis, Oc. (Och.) muelleri, Oc. (Pro.) burgeri, Oc. (Pro.) terrens, Oc. (Rusticoidus) (2 species) and Ps. 163(122). Metameron: (0) bare; (1) with scales; (2) with (Jan.) ferox. setae. The metameron is bare (state 0) in many Aedini. Scales are present (state 1) in Az. ramirezi (absent in 156(115). Upper mesokatepisternal setae: (0) absent; some specimens), Oc. (Rusticoidus), Oc. (Ochlerotatus) (1) present. One or more upper mesokatepi sternal (several species), Oc. (Fin.) echinus, Oc. (Mol.) pecu- setae are normally present in Aedini. Reinert et al. niosus (absent in some specimens), Oc. (Pro.) burgeri, (2004) noted taxa without these setae. These setae are Ps. (Jan.) ferox (absent in some specimens) and the out- absent in Hg. (Haemagogus), Mucidus (present or group species Cs. inornata. Species of Geoskusea have absent), Scutomyia and Oc. (Fin.) harperi. short, fine setae on the metameron (state 2) (see 157(116). Prealar setae: 05 20; (1) ^! 21. A few speci- Mattingly, 1959: fig. 2C, Ge. kabaenensis). mens of some species exhibit a slight overlap of one or 164(125). Alula, marginal scales: (0) absent; (1) nar- two setae between the two states, and are scored (0, 1). row; (2) broad. The presence of narrow scales on the margin 158(118). Upper prealar area, scales: (0) absent; (1) of the alula (state 1) is the usual condition in present. Scales on the prealar area may be inter- Aedini. Mattingly (1957, 1958, 1959) pointed out the spersed among the setae on the prealar knob or value of the varied development of scales on the alula present on the lower part of the knob (this character) (also see character 165). Broad scales on the margin and contiguous with the patch of scales on the lower (state 2) are found in Finlaya, Gymnoinetopa and Oc. prealar area (character 159, see Arnell, 1976: fig. 35, (Fin.) subalbitarsis. Scales are absent (state 0) in Hg. Oc. scapularis). (Hag.) splendens and Ps. (Gra.) jamaicensis. See Har- bach & Kitching (1998) for illustrations of state (1) 159(117). Lower prealar area, scales: (0) absent; (1) (their fig. 12E) and state (2) (their fig. 12D). See dis- present. When scales are present (state 1) on the cussion of scale types under character 111. lower prealar area (below knob) they form a small to Alula, dorsal moderately broad or broad moderately large patch (see Tanaka et al., 1979: 165(126). scales: figs 217, 220, 222, Oc. (Fin.) japonicus, Th. togoi and (0) absent; (1) present. See discussion of this Oc. (Fin.) seoulensis). Lower prealar scales are character in Reinert et al. (2004). Gymnometopa mediovittata is the only species in the present study absent in some species (state 0) (see Tanaka et al., that exhibits state (1). 1979: fig. 239, Ge. baisasi). See discussion under character 158. 166(127). Remigium, dorsal setae: (0) absent; (1) 160(119). Mesepimeron, scales: (0) absent; (1) present. present, distally; (2) present, proximally. The dorsal Scales are normally present (state 1) (see Tanaka remigial setae inserted distally (state 1) (see Tanaka et al., Oc. (Fin.) hatorii) et al., 1979: figs 217, 218, Oc. (Fin.) japonicus and 1979: fig. 219, usually extend Oc. (Fin.) koreicus) in one or two patches on the noticeably beyond the remigial scales, but are short mesepimeron in Aedini. However, they are absent in and hidden beneath the scales in some species. Marks (1962: fig. lc, Oc. (Cha.) humeralis Levua and Rhinoskusea (see Reinert, 1976b: fig. 2, Rh. (Edwards)) illus- longirostris). See Reinert et al. (2004) for other Aedini trated the normally long, stout, blunt-tipped setae without scales on the mesepimeron. (state 2). See Tanaka et al. (1979: fig. 220, Ta. togoi) for an illustration of state (0). 161(120). Lower anterior mesepimeral setae: (0) absent; (1) present. Most taxa examined in the present 167(128). Remigium, ventral setae: (0) absent; (1) study have no setae on the lower anterior area of the present. Setae are normally absent (state 0) from the mesepimeron. These setae are, however, present in ventral surface of the remigium in Aedini. They are Ha. australis, Ia. cavaticus, Mucidus (some species), present (state 1) on both males and females in the two Opifex Oc. (Rusticoidus), Oc. communis, Op. (Opi.) fuscus, monobasic subgenera of (see Belkin, 1968: fig. 2, Op. (Opi.) fuscus), and are also present in the Psorophora, Th. togoi and the outgroup species Cs. outgroup species Cs. inornata. inornata, Cx. quinquefasciatus and Ma. titillans. See Reinert et al. (2004) for a discussion of this character 168(new). Costa, scales: (0) all dark; (1) one pale-scaled in other Aedini and illustrations in Harbach & Kitch- patch at or near base; (2) ^ 3 pale-scaled patches; (3) ing (1998: fig. 11H, setae absent, state (0); and hG, pale and dark scales intermixed for all or most of setae present, state (1)). length, not forming defined pattern. Most aedine spe-

Journal compilation © 2006 The Linnean Society of London, Zoological Journal of the Linnean Society, 2006, 148, 1-101 No claim to original US government works. PHYLOGENY AND CLASSIFICATION OF FINLAYA 19 cies have the costa entirely dark-scaled (state 0) (see present in Oc. (Fin.) alboannulatus, but these are Tanaka et al., 1979: fig. 217, Oc. (Fin.) japonicus) or incomplete on one or more of the femora, and this spe- have a small to large patch of pale scales at the base of cies is scored (0). et al., 1979: fig. 222, Oc. the costa (state 1) (see Tanaka Midfemur, narrow, median, pale-scaled seoulensis). A few species, e.g. Oc. (Fin.) banksi 174(new). (Fin.) stripe on anterior surface from base to or near apex: (0) and Oc. (Fin.) pseudotaeniatus, have a small dark- scaled patch at the base of the costa and a white-scaled absent; (1) present. Most aedine species do not have a median, pale-scaled stripe on the anterior surface patch near the base on the anterior surface (scored as of the midfemur (state 0) (see Tanaka et al., 1979: state 1). Three or more white-scaled patches on the This costa (state 2) are typical of genus Finlaya (see Russell, fig. 223, Oc. (Br/i.) alektorovi (= kobayashii)). feature is present (state 1) in Oc. (Fin.) albilabris, 1996: 61, Fl. kochi). Pale and dark scales are intermixed along the entire length of the costa and do not Oc. (Fin.) banksi, Oc. (Fin.) elsiae, Oc. (Fin.) hatorii, Oc. (Fin.) macfarlanei, Oc. (Fin.) notoscriptus and Oc. form a defined pattern (state 3) in some species, e.g. Ps. (Fin.) pseudotaeniatus St. aegypti (Gra.)jamaicensis, Ps. (Pso.) ciliata and the outgroup of the ingroup, and et al., 1979: fig. 234) and Gy. mediovittata species Cs. inornata, Ma. titillans and Or signifera. (see Tanaka of the aedine outgroup taxa. In the last two species, this 169(129). Anal vein: (0) ending approximately at level stripe is sometimes incomplete in some specimens but of intersection of mcu and cubitus; (1) ending notice- is scored (1). Other species have a pale-scaled stripe on ably distad of this point. Termination of the anal vein the ventral margin of the anterior surface (scored 0). beyond the intersection of the mediocubital crossvein pale scales dorsally and/or and the cubitus, with the distal portion normally 175(new). Hindfemur, nearly straight or gently curved, is the usual condition anteriorly at apex: (0) absent; (1) present. The apices of in Aedini. Termination near the intersection of mcu the hindfemora in some species of Aedini have dark and the cubitus, with the distal part sharply curved, scales on the dorsal and anterior areas (state 0), e.g. Oc. (Fin.) seoulensis occurs in Rhinoskusea, Belkinius, Zeugnomyia and Oc. (Fin.) crossi and (see exam- et al., 1979: figs 222, 226, 227, Oc. Ae. (Cancraedes). ples in Tanaka (Fin.) seoulensis, Do. nipponica and Do. nishikawai). 170(131). Anteprocoxal scales: (0) absent; (1) present. A few species possess a subapical pale-scaled area Scales are normally absent from the anteprocoxal with a very narrow band or fringe of dark scales at membrane of Aedini. They are, however, found in sub- the apex, e.g. Oc. (Fin.) chrysolineatus, Oc. (Fin.) genus Rusticoidus of Ochlerotatus (present study) and embuensis, Oc. (Fin.) ingrami and Oc. (Fin.) japoni- some other species (see Reinert et al., 2004). cusjaponicus (see Tanaka et al., 1979: fig. 217). These species are scored (0). Many species of Aedini have a 171(132). Postprocoxal scales: (0) absent; (1) present. narrow to broad band of pale scales at the apex of Most taxa examined have the postprocoxal area bare the hindfemur (state 1), e.g. Ta. savoryi and Ta. togoi (state 0) (see Zavortink, 1972: fig. 5, Oc. (Pro.) (see Tanaka et al., 1979: fig. 220). brelandi). Scales are present (state 1) on the postprocoxal membrane in Abraedes, Gymnometopa 176(new). Hindtibia: (0) with dark scales only; (1) with (see Zavortink, 1972: fig. 38, Gy. mediovittata), Hg. pale-scaled areas. The hindtibia may be entirely dark- (Haemagogus), Oc. (Rusticoidus), Oc. (Och.) flu viati- scaled (state 0), or dark-scaled with pale scales that lis, Oc. (Och.) muelleri, Oc. (Och.) varipalpus and are intermixed, forming an anterior and/or posterior some other Aedini (see Reinert et al., 2004). stripe, developed as a narrow or broad basal, median or apical spot(s) or band(s) (see Banez & Jueco, 1966: Base of hindcoxa: 172(133). (0) well below dorsal mar- p1. 1, Fl. poicilia; Russell, 1996: 61, Fl. hoc/i), or gin of mesomeron; (1) more or less in line with or entirely pale-scaled (state 1). slightly above dorsal margin of mesomeron. See the discussion of this character in Reinert et al. (2004). 177(new). Hindtarsomere 1, basal pale scales or band: State (1) occurs in Hg. (Haemagogus), Scutomyia and (0) absent; (1) present. The basal part of hindtarso- Oc. (Cha.) wattensis in the present study. mere 1 may be dark-scaled (state 0) (see Tanaka et al., 1979: fig. 226, Do. nipponica) or have pale scales form- 173(134). Fore-, mid- and hindfemora, all with ing a narrow to wide band (see Tanaka et al., 1979: narrow, subapical, pale-scaled band: (0) absent; (1) fig. 222, Oc. (Fin.) seoulensis) or small spot on the dor- present. In Aedini, the fore-, mid- and hindfemora nor- sal, ventral or posterior surface (state 1). mally do not have a complete subapical band (state 0). All three femora with a distinct subapical pale-scaled 178(new). Hindtarsomere 1, one or more median pale- band (state 1) occur in some aedine taxa, e.g. Finlaya, scaled bands: (0) absent; (1) present. The median area Fredwardsius (see Mattingly, 1965: fig. 5A), Huaedes of hindtarsomere 1 may be entirely dark-scaled (state and Ps. (Grabhamia). Subapical pale-scaled bands are 0) or have one or more pale-scaled bands (rarely

reduced to dorsal pale-scaled spots) (state 1) as in gues (state 0) occur in males of Op. (Opi.) fuscus, Le. Finlaya (see Banez & Jueco, 1966: p1. 1, Fl. poicilia; geoskusea, Sc. albolineata and St. aegypti. Some spe- Russell, 1996: 61, Fl. kochi). cies, e.g. Ab. papago, Hg. (Hag.) splendens, Oc. (Cha.) wattensis tremulus, 179(new). Hindtarsomere 1, apical pale scales or band: and Oc. (Mac.) exhibit state (1). (0) absent; (1) present. The apical part of hindtarso- Most species examined have both midungues toothed mere 1 may be dark-scaled (state 0) (see Zavortink, (state 2). 1972: fig. 42, Oc. (Och.) muelleri) or have pale scales 186(new). Larger midunguis, development (males): (0) forming a narrow to wide band or small spot on simple; (1) with 1 tooth; (2) with 2 teeth. Males of only the dorsal, ventral or posterior surface (state 1) a few taxa examined have the larger midunguis sim- (see Zavortink, 1972: fig. 48, Oc. (Och.) atropalpus). ple (state 0), e.g. Hg. (Con.) leucotaeniatus, Le. geoskusea, Op. (Opi.) fuscus, Scuto,nyia 180(new). Hindtarsomere 2, basal pale scales or band: and Stegomyia. (0) absent; (1) present. See discussion under character The larger midunguis with a tooth (state 1) occurs in 177. several species, e.g. Abraedes, Aedes, Geoskusea, Isoaedes, Kenknightia, Oc. (Mac.) tremulus, Op. (Not.) 181(new). Hindtarsomere 2, apical pale scales or band: chathamicus and Zavortinkius. Numerous other taxa (0) absent; (1) present. See discussion under character have the larger unguis with two teeth (state 2), e.g. 179. Aztecaedes, Downsiomyia, Finlaya, Psorophora and Tanakaius. 182(136). Foreungues: (0) both simple; (1) one simple, one toothed; (2) both toothed. Only a few taxa examined 187(139). Hindungues (males): (0) both simple; (1) have females with both foreungues simple (state 0), e.g. both toothed. Males of most species examined have Abraedes, Geoskusea, Howardina (see Berlin, 1969: both hindungues simple (state 0) but some have both fig. 16, Hw. walkeri), Levua, Rhinoskusea, Scutomyia toothed (state 1), e.g. Ha. australis, Mu. alternans, and the outgroup species Cs. inornata,Ma. titillans and Mu. laniger, Op. (Not.) chathamicus, Ps. (Jan.) ferox, Or signifera. State (1) with one unguis simple and the Ps. (Pso.) ciliata, Tanakaius and Zavortinkius. other unguis toothed occurs in the outgroup species Cx. quinquefasciatus. Most aedine taxa have both foreun- 188(140). Hindungues: (0) both simple; (1) both gues toothed (state 2) (see Tanaka et al., 1979: figs 219, toothed. Females of most taxa examined have both 220, Oc. (Fin.) hatorii and To. togoi). hindungues simple (state 0), but those of a few taxa have both toothed (state 1), e.g. Aedes, Halaedes, Mu. 183(137). Foreungues (males): (0) both simple; (1) one alternans, Mu. laniger, Opifex, Tanakaius, Zavor- simple, one toothed; (2) both toothed. The male fore- tinkius and subgenus Rusticoidus of Ochlerot at us. ungues are both simple (state 0) and large in Op. (Opi.) fuscus (see Belkin, 1968: figs 2, 18). One unguis 189(141). Laterotergite, scales: (0) absent; (1) present. is simple and the other toothed (state 1) in specimens Scales are absent (state 0) in Geoskusea, Oc. (Fin.) examined of Abraedes, Aedes, Hg. (Hag.) splendens, candidoscutellum, Oc. (Och.) atropalpus, Oc. (Och.) Hw.sexlineata, Levua, Oc. (Cha.) wattensis, Oc. epactius, Oc. infirmatus, Oc. scapularis, Opifex, Ps. (Och.) muelleri, Rhinoskusea, Scutomyia, Stegomyia (Pso.) ciliata and the four outgroup species. Most and the outgroup species Ma. titillans and Or sig- Aedini have numerous, or at least a few, scales on the nifera. The other species examined have both foreun- laterotergite of abdominal segment I (state 1). gues toothed (state 2). 190(142). Abdominal terga, lateral setae (males): (0) 184(new). Larger foreunguis, development (males): (0) few, relatively short; (1) numerous, short to moder- simple; (1) with 1 tooth; (2) with 2 teeth. The larger ately long; (2) numerous, long, with curved apices. foreunguis of males may be simple (state 0) as in The lateral setae on the abdominal terga of males are Op. (Opi.) fuscus (see Belkin, 1968: figs 2, 18) and few in number and relatively short (state 0) in some St. africana, have a single tooth (state 1) as in taxa, e.g. Geoskusea, Hg. (Conopostegus), Hg. (Hag.) Abraedes, Aedes, Gymnometopa (see Zavortink, 1972: splendens, Howardina, Isoaedes, Kenknightia, Oc. (Bruceharrisonius), Op. (Opi.) fuscus, Rhinoskusea, fig. 38, Gy. mediovittata), Haemagogus, Isoaedes, Ken- knightia, Rhinoskusea, Scutomyia and others, or have Scutomyia, St. aegypti, Tanakaius and Zavortinkius. two teeth (state 2) (see Zavortink, 1972: fig. 35, Az. These setae are numerous and relatively short to mod- Downsiomyia, ramirezi), one small tooth usually at or near the base erately long (state 1) in other taxa, e.g. and a second larger tooth near midlength as in many Finlaya, Oc. (Mac.) tremulus and Oc. (Mol.) pecunio- of the species examined. sus. The setae are numerous and long with apices curved (state 2) in numerous taxa, e.g. Aedes, Mu. 185(138). Midungues (males): (0) both simple; (1) one alternans, Mu. laniger, Oc. (Protomacleaya) (most simple, one toothed; (2) both toothed. Simple midun- species) and Oc. (Ochlerotatus) (many species).

191(143). Abdominal segment VII, shape: (0) laterally 195(147, in part). Tergum VIII, scales: (0) absent, compressed; (1) dorsoventrally flattened; (2) cylindri- occasionally with 1-3 adventitious scales; (1) numer- cal. Abdominal segment VII (and often segment VIII) ous. Most aedines examined in the present study have of dead, dried females has a distinctive shape when numerous scales on tergum VIII (14 or more scales, viewed transversely, i.e. laterally compressed, dors- some species with scales nearly covering the tergum), oventrally flattened or cylindrical. The cylindrical especially on the distal part (state 1). Some species shape occurs in Psorophora and the outgroup species have no scales (state 0), e.g. Geoskusea, Levua, Oc. Ma. titillans. See Tanaka et al. (1979: fig. 229, communis, Oc. infirmatus, Oc. scapularis, Oc. (Och.) Oc. (Fin.) watasei) for state (0) and their fig. 212 deserticola, Oc. (Pseudoskusea), Psorophora, Rhino. (Oc. dorsalis) for state (1). skusea 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 FEMALE GENITALIA specimen with 4 or 5) scales scattered over the tergum, and are scored (0). The series of papers by Reinert (2000b, c, d, e, f, 2001b, c, d, e, 2002b, c, d, e) on the comparative anatomy of 196(148). Sternum VIII, position of seta 2-S: (0) inser- Aedini should be consulted for illustrations and tion noticeably posterior to seta 1-S; (1) insertion lat- descriptions of female genital structures. erally at or near same level as seta 1-S. A number of taxa have numerous long setae along the median area 192(144, in part). Intersegmental membrane between that do not occur in a distinct pattern. Since these segments VII and VIII: (0) short to intermediate; (1) setae are mainly arranged longitudinally, they are long to very long. Taxa with intersegmental mem- scored as state (0). branes separating segments VII and VIII that allow 50% or less of tergum VIII to be retracted into segment 197(149, in part). Sternum VIII, scales: (0) absent, VII are considered short to intermediate whereas occasionally with 1-3 adventitious scales; (1) several those allowing 65% or more retraction are scored long to numerous scales, often covering much of surface. to very long. The very long intersegmental membrane Some species of Aedini have no scales on sternum VIII; in Psorophora allows for nearly complete retraction of however, some specimens occasionally have 1-3 scales segment VIII into segment VII, whereas taxa with scattered over the surface (state 0) (see Reinert, short intersegmental membranes do not allow retrac- 2002e: fig. 3, Oc. (Fin.) geniculatus). Ochierotatus tion or only a slight retraction of segment VIII. (Fin.) macfarlanei has no scales or rarely one or two present, but the sternum has numerous short Ian- Tergum VIII, development: 193(145, in part). (0) ceolate setae interspersed with slender setae (scored mostly membranous, with lateral rod-shaped struc- 0). Most aedine species have several (10 or more tures; (1) entirely sclerotized (rarely with only small scales) (see Reinert, 2002e: fig. 5, Oc. (Fin.) guberna- median nonsclerotized area), without lateral rod- tons) or numerous scales covering most of the surface shaped structures. Psorophora have tergum and ster- of sternum VIII (see Reinert, 2002e: figs 4, 6, Oc. num VIII nearly completely membranous except for (Brh.) greenii and Fl. kochi) (state 1). small scattered, scierotized islands usually hearing setae and a rod-shaped structure laterally on each 198(150, in part). Tergum IX: (0) wide, width ^ 2.0 side (state 0) (see Reinert, 2000c: figs 1-3). Most times length; (1) narrower, width !^ 1.8 times length. Aedini have tergum and sternum VIII entirely selero- Few aedines have tergum IX width greater than or tized (state 1) (see figures in Reinert, 2000f, 2001c, equal to 2.0 the length, e.g. Oc. (Fin.) hanksi, Oc. 20020. Tergum VIII of the outgroup species Ma. titil- (Fin.) embuensis, Oc. (Fin.) pseudotaeniatus, Oc. lans has distal and lateral sclerotized areas and a (Och.) atropalpus, Oc. (Och.) epactius and Oc. (Och.) large basomesal membranous area (scored 1). Ster- fluviatilis. This also applies to the four outgroup spe- num VIII is scierotized in this species. cies. The three species of Oc. (Ochlerotatus) listed here also have tergum IX divided into two small scler- 194(146). Tergum VIII, selae: (0) on distal 0.60 or less; ites that are separated by membrane (see character (1) on distal 0.70 or more. Some aedine taxa have most 199). The outgroup taxa, e.g. Cx. quinquefasciatus, of the surface (^ 0.70) of tergum VIII covered with sev- titillans (see Gerry, 1932: figs 28, 40) and Cs. inor- Aedes, Geoskusea, Ma. eral to numerous setae (state 1), e.g. nata (see Rees & Onishi, 1951: fig. 35) have a wide and Halaedes, Kompia, Levua, Oc. (Bruceharrisonius), narrow tergum IX. Oc. (Ochierotatus) (most species), Oc. (Pseudosku- sea), Oc. (Rusticoidus), Op. (Nothoskusea), Ps. (Jan- 199(new). Tergum IX, development: (0) single scierite, thinosoma), St. aegypti and Ta. togoi. Other groups usually with small to deep apical emargination; (1) two have few to several setae that are restricted to the dis- scierites connected by membrane. Most aedine species tal 0.60 or less of the tergum (state 0). have tergum IX as a single selerite (state 0). This scler-

ite may be entirely scierotized or have a narrow mesal Ge. baisasi. Culex quinquefasciatus has a well- connection between the lateral parts, and it may be developed, horseshoe-shaped, lower vaginal sclerite darkly pigmented or have the mesal area slightly paler. located medially on the lower vaginal wall between Two lateral sclerites connected by membrane (state 1) the inner margins of the lower vaginal lip. are found in Ke. dissimilis, Ke. harbachi, Oc. (Och.) atropalpus, Oc. (Och.) epactius, Oc. (Och.) fluviatilis, 204(155, in part). Insula: (0) tonguelike; (1) liplike. Op. (Not.) chathamicus and Op. (Opi.) fuscus. Most taxa examined in the present study have a liplike insula (state 1) that is noticeably wider than long. Taxa 200(151). Tergum IX, setae: (0) absent; (1) present dis- examined that have a tonguelike (ungulate) insula tally; (2) present, distally and on much of medial area. (state 0) include Aedes, Haemagogus, Isoaedes, Psoro- Tergum IX with no setae (state 0) (see Reinert, 2002e: phora, Scutomyia and Stegomyia. See Reinert et al. fig. 9, Oc. (Fin.) papuensis) occurs in a few species. (2004) for a discussion of this character in tribe Aedini. Most aedine species have one or more setae distally on Insular setae: (0) tergum IX (state 1) (see Reinert, 2002e: figs 2, 5, 9, Oc. 205(156). absent; (1) present in lat- (Finlaya) species). Psorophora and Az. ramirezi have eral patches; (2) present in medial patch. The majority setae on the distal area that extend along the mesal of taxa examined in the present study have lateral area (state 2) (see Reinert, 2000c: figs 1-3, Psorophora). patches of setae on the insula (state 1). Aedes, Isoaedes, Scutomyia and Stegomyia have no setae 20 1(152, in part). Postgenital lobe, position of ventral (state 0) but instead may have small tuberculi (each setae: (0) distal; (1) lateral; (2) on entire surface. After with or without a tiny spicule). Haemagogus and the examining a larger series, including new specimens, of outgroup taxa Cs. inornata and Cx. quinquefasciatus the outgroup species, this character was reevaluated; have setae in a median patch (state 2). hence, Cs. Lnornata (see Rees & Onishi, 1951: fig. 35), Cx. quinquefasciatus (see Sirivanakarn, 1976: fig. 14), 206(new). Cercus, scales: (0) absent; (1) present. Most Ma. titillans (see Ronderos & Bachmann, 1963: fig. 27) species have no scales on the cerci (0) (see Appendix 2). and Or. signifera are now scored (1). The presence of Some specimens of a few species have one adventitious numerous (usually 10 or more) setae in elongate, lat- scale on one cercus, and are scored (0). eral patches on the ventral surface of the postgenital 207(157). Spermathecal capsule(s): (0) one; (1) three. lobe of these species differs from the species of Aedini Downsiomyia (except two species) and Rhinoskusea examined (see Reinert, 2002e: figs 3, 12, 34, 38, Oc. have a single large spermathecal capsule (state 0). The (Fin.) geniculatus, Ke. dissimilis, Oc. (Pro.) triseria- other species studied have one large and two smaller tus and Oc. (Rus.) refiki), which have setae located dis- spermathecal capsules (state 1). Reinert et al. (2004) tally on the postgenital lobe, scored 0, except Op. provide information on this character in Aedini. (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 MALE GENITALIA apical emargination forming two lobes, each bearing a 208(new). Tergum IX, posterior margin: (0) with pair few to several setae distally (see Reinert, 2002e: of strongly scierotized, darkly pigmented, short lobes, fig. 14, Le. geoskusea), and these are scored (0). often narrow and acute; (1) with pair of poorly sclerotized, lightly to moderately pigmented, broad, gently 202(153). Upper vaginal sclerite: (0) absent; (1) present. An upper vaginal scierite is present in most rounded or flattened lobes. See Zavortink (1972: figures ofOc. (Protomacleaya)) et al. of the species examined, including most species of and Tanaka (1979: Oc. (Finlaya). The sclerite may be small and restricted fig. 91, Oc. (Och.) hakusanensis) for state (0) and Op. (Opi.) fuscus to the inner margin of the upper vaginal lip, or extend Belkin (1962: figs 207, 233, and Or. (Fin.) argyronotum) as a well-developed plate mesally on the upper vaginal for state (1). Also, see the discussion under character 209. wall. This scierite is absent, e.g. Aztecaedes, Hg. (Conopostegus), Kompia, Oc. (Finlaya) (few species), 209(158). Tergum IX, setae: (0) absent; (1) all slender; Oc. (Ochierotatus) (most species), Oc. (Rusticoidus), Oc. (2) some or all stout. Species with strongly sclerotized, communis, Oc. infirmatus, Oc. scapularis, Psorophora, short lobes normally bear stout and often flattened Rhinoskusea and Tanakaius. The upper vaginal scier- setae. Those with weakly sclerotized, broad, gently ite is present in the outgroup species Cs. inornata, Cx. rounded lobes normally bear longer, slender setae, and quinquefasciatus, Ma. titillans and Or. signifera. if the lobes are flattened and often sloping the setae are often absent (see discussion for character 208). 203(154). Lower vaginal sclerite: (0) absent; (1) present. The lower vaginal scierite is normally absent 210(159). Sternum IX, length: (0) short; (1) moderately in Aedini. A poorly developed scierite on the inner long; (2) long. Sternum IX is moderately long (state 1) margin of the lower vaginal lip is present in in the majority of aedine species. It is short (state 0)

Journal compilation © 2006 The Linnean Society of London, Zoological Journal of the Linnean Society, 2006, 148, 1-101 No claim to original US government works. PHYLOGENY AND CLASSIFICATION OF FINLAYA 23 in a few species, e.g. Aedes, Oc. (Fin.) albocinctus, Oc. 217(161). Gonocoxite, lateral setae: (0) mostly short; (1) (Fin.) banksi and Oc. (Fin.) pseudotaeniatus, and rel- mostly long. Lateral setae are considered short if they atively long (state 2) in Ge. longiforceps, Rhinoskusea are less than 0.80 of the width of the gonocoxite dorsal (see Reinert, 1976b: figs 6-8) and Oc. (Och.) sierrensis. surface at the point of attachment whereas they are long if they are longer than, and normally noticeably 211(160). Sternum IX, setae: (0) absent; (1) present. longer than, the width. In most Aedini, the lateral sur- Most taxa have setae present on sternum IX (state 1). face of the gonocoxite has numerous long, stout setae Setae are absent (state 0), e.g. Ia. cavaticus, Op. (Opi.) (state 1) (see Belkin, 1962: figs 276, 278, Mu. altern- fuscus, St. aegypti, St. africana and the outgroup spe- ans and Mu. painei). Most of the lateral setae of the cies Cx. quinquefasciatus. Setae are present or absent gonocoxite in Op. (Opi.) fuscus are short (state 0) (scored 0,1) in specimens of the outgroup species (see Belkin, 1962: fig. 207). Ma. titillans and Or signifera. Gonocoxite, ventral surface with specialized 212(new). Gonocoxite, dorsomesal apical lobe: (0) 218(new). absent; (1) present. The dorsomesal apical lobe is scales on mesal margin: (0) absent; (1) present. The absent (state 0) in most taxa examined in the present gonocoxites of Aedini usually lack specialized scales on study. The lobe is present (state 1) in Oc. cominunis, the mesal margin of the ventral surface (state 0) (see Le. geoskusea, Mu. Oc. infirinatus and Oc. scapularis (see Arnell, 1976: Belkin, 1962: figs 273, 276, 315, Sc. albolineata). Well-developed, long, fig. 30, Oc. infirmatus, lobe well developed; fig. 36, Oc. alternans and Downsiomyia, scapularis, lobe moderately developed), Oc. (Rusticoi- specialized scales (state 1) are found in Finlaya Finlaya, as dus) and Rhinoskusea. The outgroup species Cx. quin- (see figures in Belkin, 1962 of Kochi Group), Haemagogus, Oc. (Fin.) albotaeniatus, quefasciatus has a well-developed lobe bearing specialized setae mesally on the dorsal surface of the Oc. (Fin.) biocellatus, Oc. (Fin.) harperi, Oc. (Fin.) gonocoxite, but the lobe is subapical. This is scored (0). pulchri venter and Tanakaius. 213(new). Gonocoxite, dorsomesal basal lobe: (0) 219(164). Gonostylus, attachment to gonocoxite: (0) absent; (1) present. Geoskusea lack a basomesal lobe subapical; (1) apical. The dorsal surface of the gono- on the dorsal surface of the gonocoxite (state 0), but coxite extends noticeably beyond the attachment of most species (all except Ge. longiforceps) have a the gonostylus in Op. (Opi.) fuscus (see Belkin, 1962: densely setose lobe on this surface at approximately fig. 207) whereas the ventral surface extends notice- midlength of the mesal margin (see Belkin, 1962: ably beyond the attachment of the gonostylus in the figs 214, 215, Ge. becki, Ge. daggyi and Ge. perryi; Africanus Group of Stegomyia (see Huang, 1990: Tanaka et al., 1979: fig. 127, Ge. baisasi). fig. 23A, B, St. opok and St. pseudoafricana) and genus Aedes (see Tanaka et al., 1979: fig. 133, 214(163). Gonocoxite, scales: (0) absent; (1) present. Ae. esoensis and Ac. yamadai), and these taxa are Most species of Aedini have scales on the gonocoxite scored (0). In Op. (Not.) chathamicus, the dorsal sur- (state 1) (see Belkin, 1962: figs 210, 226). A few species face of the gonocoxite extends slightly beyond the have no scales on the gonocoxite (state 0), e.g. Op. attachment of the gonostylus, but is scored (1). (Opi.) fuscus (see Belkin, 1962: fig. 207), Ps. (Pso.) ciliata, as well as the outgroup species Cs. inornata and 220(166, in part). Gonostylus, development: (0) moder- Cx. quinquefasciatus. ately long to long, relatively narrow throughout length but distal part narrower and usually somewhat 215(162). Gonocoxite, mesal surface: (0) entirely mem- curved mesally; (1) same as state 0 but distal part branous; (1) partly or entirely sclerotized. The mesal somewhat expanded; (2) short to moderately long, surface of the gonocoxite is commonly membranous basal area broader, bifurcate; (3) proximal and distal (state 0) in the species of Aedini examined, however, parts narrow, middle part noticeably expanded. The this area is partially or completely scierotized (state 1) gonostylus is normally moderately long to long, rela- in Psorophora, Rhinoskusea and the outgroup species tively narrow throughout its length and the distal part inornata, Cx. quinquefasciatus and Or. signifera. Cs. is somewhat narrower and usually slightly curved 216(new). Gonocoxite, seta(e) on hasomesal area of mesad (state 0) in most aedines examined in the dorsal surface: (0) all slender; (1) one or more stout or present study (see Belkin, 1962: figs 226, 235, 276, Oc. flattened; (2) absent. The basomesal area of the (Fin.) notoscriptus, Fl. samoana and Mu. alternans). gonocoxite dorsal surface bears only slender setae in Rhinoskusea (see Reinert, 1976b: fig. 6, Rh. longiros- state (0) (see illustrations of Oc. (Protomacleaya) in tris) and most Stegomyia have the proximal part of the Zavortink, 1972). This area in state (1) has one or more gonostylus relatively narrow and the distal part of the setae stout and/or flattened (see Wood et al., expanded (state 1). State (2) occurs in a few species, 1979: p1. 20, Oc. communis). In Ge. baisasi this area is e.g. Aedes, Op. (Opi.) fuscus (see Belkin, 1962: fig. 207) bare (see Tanaka et al., 1979: fig. 127) and is scored (2). and the outgroup species Ma. tilillans. Psorophora

have the proximal and distal parts relatively narrow Species that do not have a gonostylar claw (see previous and the middle part noticeably expanded (see Belkin character), e.g. Aedes, are scored as missing H. et al., 1970: figs 57, 59, 66, Ps. (Pso.) ciliata, Ps. (Jan.) ferox and Ps. (Gra.) jamaicensis), state (3). 224(new). Gonostylus/gonocoxite index: (0) <0.40; (1) ^! 0.42. The gonostylus/gonocoxite index was 221(new). Gonostylus, subapical seta(e): (0) absent; (1) defined by Reinert (1990) as the ratio determined by one or more present. The gonostylus of most aedines dividing the gonostylus length by the gonocoxite studied have one or a few normally short, fine subap- length. Length of the gonocoxite (Reinert, 1990) is the ical setae (state 1) (see Tanaka et al., 1979: fig. 97, Oc. length measured along a straight line from the tip of (Fin.) japonicus), but these setae are stouter (state 1) the dorsal attachment of gonocoxite (dga of Knight & in some species (see Barraud, 1934: fig. 37d, e, Oc. Harrison, 1988) to the apex. See discussion under (Fin.) elsiae and Oc. (Fin.) shortti). Subapical setae character 223 for measurement of gonostylus length. are absent (state 0) in some taxa (see Belkin, 1962: 225(168, in part). Claspette, development: (0) basal fig. 315, Sc. albolineata), but Scutomyia possess a sin- setose plaque variously developed (setae simple, spe- gle long, stout spiniform inserted some distance from cialized or both); (1) single short to long, slender, the apex (see discussion for character 222). columnar lobe; (2) two lobes, one long and columnar. 222(167, in part). Gonostylar claw: (0) absent; (1) The claspettes and the basal mesa] lobes are homolo- present, one narrow, short to long spiniform, insertion gous structures in Aedini and can be identified as such at or near apex; (2) present, one striated, short, spat- by their connection basomesally to a spiculate, more- ulate spiniform, insertion at or near apex; (3) present, or-less narrow, somewhat troughlike aedeagal guide one stout, short, clawlike spiniform, insertion at or (see Reinert, 1999b, 2000a). Reinert (2000a) noted near apex; (4) present, one stout, long spiniform, that this structure is apparently nearly lost in species insertion some distance from apex; (5) present two or of Geoskusea and Oc. (Pseudoskusea) because it more narrow, short to moderately long spiniforms, appears to be reduced to a spiculate aedeagal guide insertion distally; (6) present, two stout, short, side by and a short thin strip extending laterally onto the side, clawlike spiniforms, insertion at or near apex. basosternal area of the mesal surface of the gonocox- Aedes lack a spiniform (state 0) (see Tanaka et al., ite, and bears one or a few short, thin, inconspicuous 1979: fig. 133, Ae. esoensis and Ae. yamadai). Most setae. He also reported that the claspette is without a aedines examined exhibit state (1) (see Belkin, 1962: filament in Rhinoskusea, and is bifurcate in Rh. Ion- figs 210, 254, Ha. australis and Fl. neogeorgiana). A girostris and Rh. wardi. In Oc. (Bruceharrisonius), single, short, striated, spatulate claw (state 2) occurs the claspette comprises a median, ventral, columnar in Rhinoskusea (see Reinert, 1976b: fig. 6, Rh. longi- stem with a terminal, large, flattened filament, and a rostris) and the outgroup species Or. signifera. A sin- narrow, lateral arm extending from the base of the gle, dark, stout, clawlike spiniform (state 3) occurs in stem that bears few to several setae, some of which are Op. (Opi.) fuscus (see Belkin, 1962: fig. 207), Psoro- flattened (scored 2) (see Reinert, 2003: fig. 2A, Oc. phora and the outgroup species Ma. titillans. Scuto- (Brh.) greenii). The filament is absent in Op. (Not.) myia (see figures in Huang, 1979) have a single, stout, chathamicus and the stem is developed as a bluntly long spiniform inserted some distance from the apex rounded, lobelike process that is densely covered with (state 4). Two or more narrow, short to moderately short, thin setae (see Belkin, 1968: fig. 20). In long spiniforms inserted on the distal part of the gono- Halaedes, the claspette appears to be developed as an stylus (state 5) occur in Oc. (Fin.) elsiae, Oc. (Fin.) oblong plaque bearing a number of curved, lanceolate macfarlanei and related species (see Barraud, 1934: setae, and the dorsal lateral part is fused with the fig. 37). Two dark, short, clawlike spiniforms inserted basomesal portion of the tergal surface of the gonocox- side by side near the apex (state 6) occur in Le. geosk- ite (see Belkin, 1962: fig. 210). Arne]] (1973) pointed usea (see Belkin, 1962: fig. 273) and the outgroup spe- out that in Hg. (Conopostegus) and two species of sub- cies Cs. inornata. Ochierotatus (Fin.) biocellatus has a genus Haemagogus, i.e. Hg. anastasionis and Hg. moderately long, slender gonostylar claw with the dis- chrysochlorus, the claspette filament is inserted in a tal part forked into two attenuate branches (scored 1). distinct alveolus on the apex of the claspette stem. However, in the other species of Haemagogus the alve- 223(new). Gonostylar claw, length: (0) short, 0.35 olus is either incomplete or entirely absent at the base gonostylus length; (1) moderately long to long, 0.39 of the filament. Species of Haemagogus are scored (1). gonostylus length. Reinert (1990) defined the gonosty- Psorophora have the claspette variously developed lar claw length as the absolute length measured from from a short plaque, e.g. Ps. (Gra.) jamaicensis, to a the base to the apex and the gonostylus length as the long, flattened stem with distal part expanded, e.g. Ps. absolute length of gonostylus, minus gonostylar claw, (Jan.) ferox and Ps. (Pso.) ciliata, that projects dorso- measured from the tip ofthe basal apodeme to the apex. caudally and bears a few to several specialized setae

Journal compilation © 2006 The Linnean Society of London, Zoological Journal of the Linnean Society, 2006, 148, 1-101 No claim to original US government works. PHYLOGENY AND CLASSIFICATION OF FINLAYA 25 with flattened, fringed or simple apices, or several 230(171). Proctiger, sternal arm: (0) absent; (1) slender, simple setae, and none or one or more flat- present. The sternal arm of the proctiger is absent tened, broadened and strongly contorted filaments (state 0) in all species evaluated except for the follow- (see Belkin et al., 1970: figs 57, 59, 66). These species ing (state 1): Ia. cavaticus, St. aegypti, St. africana of Psorophora are scored as (0). In state (2), the slen- and the outgroup species Cx. quinquefasciatus. der lobe is normally distinctly columnar and may be 231(172). Proctiger, cereal setae: (0) absent; (1) short to long. The claspette is absent in the outgroup present. Most aedine species examined have cereal species Cx. quinquefasciatus, and is scored (—). In this setae present on the proctiger (state 1) (see Belkin, species the lobe with modified setae that arises from 1962: fig. 233, Oc. (Fin.) argyronotum). Opifex (Opi.) the mesa] area on the dorsal surface of the gonocoxite fuscus is unusual in that both minute and short cereal is not homologous with the claspette in Aedini, which setae are present on the proctiger (see Belkin, 1962: arises from the ventromedian area near the base of fig. 207). Cereal setae are absent (state 0) in Aedes, the gonocoxite and is connected mesally with its mate Isoaedes (see Reinert, 1979: fig. 3, Ia. cavattcus), by an aedeagal guide. Scutomyia and Stegomyia. 226(new). Claspette filament(s), development: (0) ^: 2 setae; (1) one slender or stout seta; (2) one stout, curved BIOLOGY spiniform, elliptical or circular in cross section; (3) one 232(new). Immature habitat: (0) freshwater ground moderately wide to wide, normally elongate, flattened pools; (1) brackish-water ground and rock pools; structure. In Oc. (Rusticoidus), the claspette filament is (2) freshwater rock pools; (3) freshwater containers relatively short but transversely annulate (see Guts- (phytotelmata, small artificial containers, rock holes); evich, Monchadskii & Shtakelberg, 1974: figs 133, 135, (4) crab holes. The female selects the habitat in which Oc. (Rus.) rusticus and Oc. (Rus.) refiki), and is scored the immature stages develop when she deposits her (3). This character is scored as missing (—) for Cx. quin- eggs. The primary habitat of each study taxon is listed quefasciatus. See discussion under character 225. in the data matrix (Appendix 2). Atypical habitats are 227(new). Claspette filament, distinct transverse stria- not included. Belkin (1962) classified mosquitoes into tions: (0) absent; (1) present. In most species examined two principal groups based on the habitat require- in the present study the claspette filament does not ments of the immature stages, i.e. species that inhabit have transverse striations (state 0). The claspette ground-water habitats and those that inhabit plant- filament of Finlaya is long, foliform and with a container habitats. He considered the former as membranelike sheath or flap that is annulated trans- undoubtedly ancestral and the later derived. Reinert versely on approximately the middle third whereas (2002a) indicated that aedine species with a well- the filament of several species of Oc. (Finlaya) have developed ventral brush (setae 4-X) normally inhabit the sheath absent but with several transverse stria- ground-water habitats, and the strong development of tions near midlength. These are scored (1). The fila- the brush seems to be the ancestral condition. By con- ment in Oc. (Rusticoidus) has transverse striations trast, species with a weakly developed ventral brush but it is shorter, broader and thicker (scored 1) inhabit plant containers, and the weaker brush seems (see discussion under character 226). This character is to be the more derived condition. scored (—) for Cx. quinquefasciatus (see discussion under character 225). PHYLOGENETIC ANALYSIS Aedeagus: (0) single tubelike, scooplike or 228(170). Reinert et al. (2004) conducted phylogenetic analyses troughlike structure; (1) comprising two lateral plates using both equal and implied weighting, applied to (aedeagal scierites). State (0) occurs in all species adult data and immature stage data separately, as examined in the study except in Aedes, Isoaedes, Scu- well as the data set as a whole. They concluded that tomyia, Stegomyia Cs. mar- and the outgroup species down-weighting the more homoplastic characters by nata and Cx. quinquefasciatus, which have two lateral the application of implied weights was preferable, and et al. plates (state 1). Reinert (2000a) and Reinert that this procedure should be applied to the entire (2004) discussed this character and its importance in data set in a total evidence approach. We therefore separating generic-level taxa of Aedini. analysed the present data using only implied weights 229(new). Aed.eagus, width: (0) widest in distal third; as implemented by PIWE version 3.0 (for Windows) (1) widest in middle third; (2) widest in proximal third. (Goloboff, 1997), with the default value of the concav- The various states of the aedeagal width are illus- ity constant, K = 3. We performed the analysis by heu- trated in Zavortink (1972), i.e. state (0) fig. 39, Gy. ristic search, using 5000 replications (mult5000) and mediovittata, state (1) fig. 43, Oc. (Och.) muelleri and 10 starting cladograms per replicate (holdl10). We state (2) fig. 15, Dc. (Pro.) hurgeri. then checked that PIWE had found all most parsimo-

Journal compilation © 2006 The Linnean Society of London, Zoological ,Journal of the Linnean Society, 2006, 148, 1-101 No claim to original US government works. 26 J. F. REINERT ET AL. nious (fittest) cladograms (MPCs) by searching for ofT comprises Halaedes + Opifex, followed by successively less fit cladograms using the commands Himalaius gen. nov. +Bruceharrisonius stat. nov. (all sub n (where n is the decrease in fit, in steps of 0.1) new taxa are described in Appendix 3). At the next and find (to search for all cladograms of best fit - n), level, a group separates that comprises Oc. (Fin.) can- up to a maximum of 100 000 cladograms. We then didoscutellum + Oc. (Fin.) keefri in a sister-group rela- applied the best command to this set of 100000 cla- tionship with Vansomerenis gen. nov. + (Zavortinkius + dograms to confirm that the included set of MPCs was Kenknightia). This is followed by a dade comprising the same as that with which we started. Agreement in six species of Oc. (Finlaya), and then Oc. (Pro.) grouping within a set of MPCs was summarized using zavortinki, which is sister to the remaining taxa. These a strict consensus tree (SCT). We assessed dade are divided into two large clades. In the first of these, support using Bremer support (Bremer, 1994) and Oc. (Pro.) knabi is sister to the remaining taxa, which relative Bremer support (Goloboff & Farris, 2001). comprise three clades in an unresolved trichotomy Illustrated cladograrns were prepared as described by (Fig. 113). There are two small clades, the first compris- Reinert et al. (2004) except they were first cropped, ing Oc. (Fin.) loi + Macleaya stat. nov. (including sub- resized and positioned on pages with Coreiflraw 10 genus Chaetocruiomyia stat. nov.), and the second before being imported into Microsoft Word 2000 for (Jihlienius gen. nov. + Molpemyia stat. nov.) + Phago- final editing. myia stat. nov. In the third, largest dade, Rampamyia gen. nov. is sister to the remaining taxa, which are RESULTS AND DISCUSSION divided into two almost equal subclades. The first comprises Dobrotworskyius gen. nov. + (Georgecraigius Analysis of the entire data set under implied weights gen. nov. + Patmarksia gen. nov.), of which George- produced four MPCs (Fit = 836.8), the SCT of which craigius includes subgenus Horsfallius subgen. nov. (including Bremer and relative Bremer supports for The second subclade comprises Hulecoeteomyia stat. each dade) is shown in Figure 1. The characters that nov. as the sister group of (Gilesius gen. nov. + Tana- support the branches of this SCT are shown in Figure kaius) + Collessius gen. nov. In the second large dade 2. It should be emphasized that because this is a strict (Figs 1C, 2C), there are two terminal clades comprising consensus tree, character mappings on branches groups of genera: Dahliana gen. nov., Howardina and immediately subtending and arising from polytomies Finlaya + (Downsiomyia + Haemagogus); and (Jczrnel- are not necessarily as parsimonious as they are on the lius gen. nov. + (Abraedes + (Aztecaedes + (Gymnome. MPCs from which the SCT was derived. In particular, topa + Kompia)))) + ((Isoaedes + Aedes) + (Scutomyia + apparent parallel gains in clades arising from polyto- Stegomyia)). These genera and groups of genera are all mies may be synapomorphies for subsets of these interrelated through a set of relationships that also clades on some (but not all) MPCs. Thus, caution is includes ten species ofOc. (Protomacleaya) (shaded in required not to over-interpret these mappings. Fig. 1C) and three species of Oc. (Finlaya), i.e. oreo- Aedini is recovered as monophyletic and the rela- philus, harperi and alhotaeniatus. tionships among the non-aedine outgroup taxa are the The ambiguity among the four MPCs is relatively same as reported by Reinert et al. (2004): Culiseta + minor and concerns just four parts of the cladograms, (Culex + (Orthopodomyia + (Mansonia + Aedini))). The two of which are linked. The first involves the rela- first dade to branch off the main stem is Psorophora. tionships among the terminal three species of Jarnel- Next is a group comprising Mucidus in a sister-group lius, which are either varipalpus + (deserticola + relationship with a dade that includes two subclades, sierrensis) or sierrensis + (deserticola + varipalpus). one consisting of Geoskusea, Levua and Rhinoskusea The second area of ambiguity concerns the relation- (with the last two in a sister-group relationship) and ships of Howardina, which can be placed as the sister the other comprising Pseudoskusea stat. nov. and group of either Dahliana or the dade comprising Ochierotatus (which includes the two subgenera Ochie- Oc. (Fin.) harperi, Oc. (Fin.) albotaeniatus, Finlaya, rotatus and Rusticoidus, of which the former is para- Downsiomyia and Haemagogus. The third area of phyletic relative to the latter). The next dade to branch ambiguity involves the relationships ofOc. (Fin.) loi +

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 dade 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, Go!- lessius gen. nov.; 16, Dahliana gen. nov.; 17, Jarnellius gen. nov. Species ofOc. (Protomacleaya) are shaded.

,Journal compilation © 2006 The Linnean Society of London, Zoological ,Journal of the Linnean Society, 2006, 148, 1-101 No claim to original US government works. PHYLOGENY AND CLASSIFICATION OF FINLAYA 27

Cs. (Cus.) inornata Cx. (Cux.) quinquefasciatus Or. signifera Ma. (Man.) titilians

>04 >04 Ps. (Jan.) ferox >° >100 >0.4 Ps. (Gra.) jamaicensis >100 Ps. (Pso.) ciliata >0.4 >OA Mu. altemans >100 >100 Mu. laniger 0.2 Ge. baisasi

>04 6 Ge. ion giforceps >100 >100 >04 >04 Le. geoskusea >100 >100 >0,4 Rh. longirostris 0.3 >100 Rh. wardi 18 Oc. (Psk.) bancroftianus >100 Oc. (Psk.) postspiracu!osus I® > 18 Oc. (Och.) infirmatus >100 Oc. (Och.) scapularis >0,4 >100 >04 Oc. (Och.) communis >100 >100 >0.4 Oc. (Rus.) refiki >100 Oc. (Rus.) rusticus

>0.4 Ha. austrails 100 Op. (Not.) chathamicus >100 Op. (Opi.) fuscus Oc. (Fin.) gull >100 Oc. (Brh.) alektorovi Oc. (Brh.) greenil ] ® Oc. (Fin.) candidoscutellum 1 EII Oc. (Fin.) keefei Oc. (Fin.) luteostriatus 1 >100 Oc. (Fin) pulchrithorax i >04 Za. fulgens 100 Za. Ion gipalpis >100 Ke. dissimilis >100 Ke. harbachi Oc. (Fin.) biocellatus 073 Oc. (Fin.) crossi 04 0.3 Oc. (Fin.) embuensis 40or >04 24 Oc. (Fin.) ingrami 01 Oc. (Fin.) albocinctus Oc. (Fin.) seoulensis

28 J. F. REINERT ET AL.

Oc(Pro)zavortihkij 1#1

Oc. (Fin.) Ia! 14 Oc. (Cha.) wattensis >100 Oc. (Mac.) tremulus Oc. (Fin.) chungi >100 - Oc. (MoL) pecuniosus

Oc. (Fin.) assamensis 1 4,- Oc. (Fin.) gubematoris >100 31 b- Oc. (Fin.) lophoventralis "- Oc. (Fin.) prominens Oc. (Fin.) albilabris 9 >100 Oc. (Fin) notoscriptus

03 Oc. (Fin.) alboannulatus 18 0.2 Oc. (Fin.) rubrithorax @ Oc. (Fin.) tubbutiensis >04 Oc. (Och.) fluviatilis >100 04 >100 04 Oc. (Och.) atropalpus ® >100 >04 Oc. (Och.) epactius >100 Oc. (Fin.) argyronotum

0.3 >100 Oc. (Fin.) papuensis 0.4 > Oc. (Fin.) argenteitarsis >100 >100 Oc. (Fin.) subalbitarsis Oc. (Fin.) j. japonicus Oc. (Fin.) jugaraensis 04 13 9 0,3 Oc. (Fin.) chrysolineatus 31 Oc. (Fin.) sherki >100 Oc. (Fin.) pulchriventer >100 > Ta. savory! >100 0.2 Ta. to gal Oc. (Fin.) hator!! >0.4 >04 Oc. (Fin.) els!ae >100 >100 Oc. (Fin.) macfarlanej Oc. (Fin.) banks! >100 Oc. (Fin.) pseudotaeniatus In

PHYLOGENY AND CLASSIFICATION OF FINLAYA 29 C 02 IOc. (Pro.) berIini 81 0.1 Oc. (Pro.) gaIindoi 8 Oc. (Pro.) terrens] - Oc. (Fin.) oreophilus Oc. (Pro.) burger!

9 If,— Oc. (Pro.) kompi If Oc. (Pro.) brelandi 0.2 Oc. (Pro.) hendersoni]

1 1 Oc. (Pro.) triseriatusl 8 >0.4 Oc. (Fin.) echinus 16 03 (ElIlQc.Fin.ygenicuIatus 0 H >100 Hw. walked 0.3 Oc. (Fin.) harped

04 Oc. (Fin.) albotaeniatus 8 >04 Fl. kochi .2 >100 Fl. poicilia 0.3 >04 Do. leonis 10 >100 Do. nivea >0.4 >0.4 Hg. (Hag.) equinus >100 >04 >100 Hg. (Hag.) splendens >100 01 h,e Hg. (Con.) Ieucocelaenus Hg. (Con.) Ieucotaeniatus —Oc. (Pro.) zoosophus Oc. (Och.) muelleri 0,4 Oc. (Och.) deserticola 7 >0.4 >100 Oc. (Och.) sierrensis © >0.4 14 >100 Oc. (Och.) varipalpus i Ab. papago >0.4 >100 0.4 Az. ramirezi 29 >0.4 Gy. 0.4 mediovittata >100 16 Ko. purpureipes

>0.4 Ia. cavaticus >100 >0.4 Ae. cinereus >100 >0.4 Ae. esoensis >100 >0.4 Sc. albolineata >100 >0.4 Sc. arboricola >100 >0.4 St. aegypti >100 St. africana

Macleaya and the four species of Phagomyia. When remains unchanged at 0.1. The more inclusive dade the former dade is placed as the sister group of(Jihlie- found by Reinert et al. (2004), comprising Oc. (Pro.) nius + Molpemyia) + Phagomyia, then the relation- triseriatus, Oc. (Fin.) geniculatus, Downsiomyia and ships among the species of Phagomyia are assamensis Haemagogus, is also recovered in the current study + (lop hovent ralis + (gubernatoris + prominens)). but now also includes Finlaya and Howardina. Previ- Alternatively, Oc. (Fin.) loi + Macleaya can be placed ously, Finlaya was placed as the sister group of Molpe- as the sister group of the larger dade comprising myia pecuniosus comb. nov. In the present study, this Rampamyia, Dobrotworskyius, Georgecraigius, Pat- latter taxon is recovered as the sister group of Jihlie- marksia, Hulecoeteomyia, Gilesius, Tanakaius and nius and included in the other major distal dade from Collessius, whence the relationships among the spe- that which contains Finlaya. Reinert et al. (2004) cies of Phagomyia are (gubernatoris + lop hoventralis) placed Howardina as the sister group to a dade con- + (assamensis + prominens). sisting of Abraedes, Aztecaedes, Gymnometopa, Kompia and Oc. (Och.) muelleri. This latter group is also recovered here, although its internal relation- COMPARISON WITH REINERT ETAL. (2004) ships are different. Furthermore, it is placed as the An exact comparison of the relationships portrayed in sister group to a dade consisting of representatives of Figures 1 and 2 with those reported by Reinert et al. the Aedes genus-group (i.e. Aedes in the previous (2004: Fig. 4) is not possible because numerous taxa broad sense; Section B of Belkin, 1962), namely analysed in the latter study are not included in the Isoaedes, Aedes, Stegomyia and Scutomyia. These four present analysis, and vice versa. It is nevertheless genera (seven species) represent the much larger dade revealing to compare the two patterns of relationships. of 63 species in Reinert et al. (2004), but are, never- First, and most importantly, all those taxa that Reinert theless, recovered as a monophyletic unit. Their inter- et al. (2004) treated as genera, for which two or more nal relationships are slightly different, however, being species are included in the current analysis, are recov- (Isoaedes + Aedes) + (Stegomyia + Scutomyia) instead ered here as monophyletic groups. Thus, the mono- of Isoaedes + (Aedes + (Stegomyia + Scutomyia)), but phyly of the following genera is corroborated: Aedes, this may be an artefact of the reduction in the number Downsiomyia, Geoskusea, Haemagogus, Kenknightia, of taxa sampled from this group. Mucidus, Ochierotatus, Opifex, Psorophora, Rhinosku- Another dade found by Reinert et al. (2004) com- sea, Scutomyia, Stegomyia and Zavortinkius. No genus prised (Oc. (Fin.) chrysolineatus + Oc. (Fin.) macfar- proposed by Reinert et al. failed this test. lanei) + (Oc. (Och.) atropalpus + (Tanakaius + Second, a number of more inclusive groups, compris- (Halaedes + Opifex))). This group has been split up in ing two or more genera, are also common to both sets the present analysis. The species atropalpus, chryso- of results, often with identical or very similar patterns lineatus and macfarlanei are here placed, respectively, of interrelationships among their members, and which in Georgecraigius, Hulecoeteomyia and Collessius. All may eventually form the basis of a higher-level classi- three, and Tanakaius, are still found together within a fication of the tribe. The dade Geoskusea + (Levua + single large dade (Figs 113, 2B), but the addition of Rhinoskusea) is consistently recovered. A group com- numerous species of Finlaya to this dade has prising Downsiomyia + Haemagogus is also found in changed their interrelationships into Georgecraigius + both studies, although in the present study the inter- (Hulecoeteomyia + (Tanakaius + Collessius)). Halaedes relationships among the four species of Haemagogus and Opifex are still sister groups but are now placed are slightly different in that both subgenera, Haema- much nearer to the base of the cladogram (Figs 1A, gogus and Conopostegus, are now recovered as mono- 2A). The two other species included in the study of phyletic groups. Previously the latter subgenus was Reinert et al. (2004), namely Macleaya (Chaetocrui- paraphyletic, although the dade Hg. (Con.) leucocelae- omyia) wattensis comb. nov. and Macleaya (Macleaya) nus + Hg. (Haemagogus) was supported by only a sin- tremulus comb. nov. form a sister-group pair in the gle character (71:0) and had very low branch support same large dade that includes Collessius, George- (Bremer support = 0.1). In the present study, four craigius, Hulecoeteomyia and Tanakaius. This pair, homoplastic characters (83:2, 107:1, 132:1, 213:1) now together with Finlaya and Molpemyia, was placed in support Hg. (Conopostegus), but the Bremer support the MPCs of Reinert et al. (2004) as a paraphyletic

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.

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C.) 34 J. F. REINERT ET AL. series between the dade consisting of Oc. (Pro.) group is that portion of the total group that is not triseriatus + Mc. (Fin.) geniculatus + (Downsiomyia + included within the crown group and is thus paraphy- Haemagogus)) and that comprising Gymnometopa - letic. In the present study, only a paraphyletic group Aztecaedes and the Aedes genus-group. However, such as Oc. (Pro.) brelandi, Oc. (Pro.) hendersoni and support for these groupings was low (0.1-0.2) and so Oc. (Pro.) triseriatus strictly meets the criteria for changes following the addition of a large number of being a stem group, in this case the dade comprising related taxa are not entirely unexpected. Oc. (Pro.) brelandi - Kompia purpureipes (Fig. 1C); The two large clades of the present study (Figs 113, C, Oc. (Protomacleaya) in its entirety is polyphyletic. It 213, C) were also found in the MPCs of Reinert et al. is apparent, however, that many of the groups that con- (2004: fig. 4), although with the differences in internal sist of a species of Oc. (Protomacleaya) and its larger relationships described above. Between these and the sister group are poorly diagnosed on short branches, non-aedine outgroup taxa (which also show the same and also that those species ofOc. (Protomacleaya) are branching pattern as found previously) is a series of poorly characterized in comparison with their sister clades that Reinert et al. (2004) resolved into two groups (Fig. 2). Thus, we agree with Zavortink (1972) monophyletic groups placed in a trichotomous relation- that the species ofOc. (Protomacleaya) do, in general, ship with the large dade. The first of these clades (with resemble each other phenetically, by virtue of what updated nomenclature) was Zavortinkius + (Kenk- they lack. However, such a paraphyletic/polyphyletic nightia + (Bruceharrisonius + Psorophora)), and the assemblage is in no meaningful evolutionary sense second was ((Geoskusea + (Levua + Rhinoskusea)) + natural. Ultimately, such groups should be split up (Mucidus + (Pseudoskusea + Ochlerotatus)). This latter and classified into their constituent monophyletic dade is recovered in the present analysis (Figs 1A, 2A), parts. As an interim solution, the generic and subge- although Mucidus has moved to a position as the sister neric names of such assemblages should be placed in group of the two remaining clades. Of the others, quotation marks to indicate their non-monophyletic Zavortinkius is now sister to Kenknightia in a dade status (the single quotation marks are used here that also includes Oc. (Fin) candidoscatellum, Oc. merely as a matter of journal style). Therefore, knabi, (Fin.) keefei and Vansomerenis. Bruceharrisonius is for example, should be Ochlerotatus (Protomacleaya) placed with a newly included species, Himalaius gilli knahi or Ochierotatus (Protomacleaya) knabi (abbre- comb. nov. in another dade, and Psorophora is a sep- viated as Oc. (Pro.) knabi). This convention is adopted arate lineage that is the sister group of the remaining herein for all non-monophyletic taxa that must be Aedini. Thus, these four clades no longer form two retained for the time being. monophyletic groups but are arranged in a para- Pseudoskusea is a small, homogeneous group phyletic series (Figs 1A, 2A). Overall, however, despite comprising four species (bancroftianus, culiciformis, the removal and inclusion of a large number of taxa, multiplex, postspiraculosus) in the Australasian the results of Reinert et al. (2004) and those of the Region. Theobald (1907) established Pseudoskusea as present study show remarkable congruence. a monobasic genus for Skusea multiplex Theobald. Edwards (1932) classified Pseudoskusea as a subgenus Aedes, CLASSIFICATION OF EVALUATED TAXA of where it remained until Reinert (2000a) transferred it to genus Ochlerotatus when he formally When Zavortink (1972) removed Protomacleaya from elevated this taxon from subgeneric rank within Aedes. Aedes (Finlaya), he stated: Although the subgenus is The placement of Pseudoskusea as the sister group to quite diverse, is defined mainly on negative characters, Ochlerotatus s.s. in the preferred cladogram of Reinert and many of the included species depart from the "nor- et al. (2004: fig. 4) is corroborated by the results of the mal" in 1 or more characteristics of 1 or more stages, it present analysis (Figs 1A, 2A). Whereas Reinert et al. is, I believe, natural because of the basically similar (2004) were obliged to retain Pseudoskusea in the non- male genitalia of virtually all species. However, the monophyletic Ochlerotatus sensu auctorum because species ofOc. (Protomacleaya) included in the present its sister-group relationship with Ochlerotatus was not analysis (shaded in Fig. 1) are arrayed in a pattern sim- weighting-independent (undoubtedly because it was ilar to what palaeontologists call a stem group, in con- represented only by a single species), we propose to trast to a terminal, monophyletic crown group. A restore this taxon to its original generic status based on crown group is an extant monophyletic taxon that also the present results. Support for the sister-group includes extinct phena with all the diagnostic relationship between Pseudoskusea and Ochlerotatus attributes of that taxon (or can be shown to have sec- is strong (Bremer support > 0.4; relative Bremer ondarily lost such attributes), whereas the total group support> 100), but the supporting characters are all is an extant monophyletic taxon that contains all homoplastic (Fig. 2A). Likewise, the characters that extinct phena possessing one or more of the diagnostic diagnose Pseudoskusea (10) and Ochlerotatus (9) are attributes of the taxon (Smith, 1994: 94-95). The stem also homoplastic. As is usually the case with generic-

Journal compilation © 2006 The Linnean Society of London, Zoological Journal of the Linnean Society, 2006, 148, 1-101 No claim to original US government works. PHYLOGENY AND CLASSIFICATION OF FINLAYA 35 level groups ofAedini, these genera are polythetic taxa, dence. Lien (1967) stated that crossi is very closely i.e. they are diagnosed not by unique characters but by related to suffusus, the only member of the Suffusus unique combinations of characters. Hence, although Subgroup (Geniculatus Group) in the classification of the individual characters may be independently Knight & Marks (1952). After examining adult and derived elsewhere, they do not occur in the same com- immature stages ofOc. suffusus, we concur with Lien bination in any other group. (1967) and fail to see any unambiguous morphological The sister-group relationship of Himalaius gilli and evidence of a natural relationship between crossi and Bruceharrisonius is strongly supported (Bremer the other species of the biocellatus - seoulensis dade. support > 0.4; relative Bremer support> 100) by five Consequently, it will be necessary to include addi- homoplastic characters (Figs 1A, 2A). However, we tional related species and/or character data in future have chosen to recognize gilli and Bruceharrisonius as studies to resolve the relationships of these species. separate genera because the number of characters that The first dade in the unresolved trichotomy distinguish them is similar to the number that (Figs 113, 2B) is a bit of an enigma. Chaetocruiomyia + separates other generic pairs, e.g. Zavortinkius and Macleaya is placed as sister to Oc. (Fin.) loi. However, Kenknightia. Because Himalaius exhibits clear dis- support for this sister-group relationship is weak tinctions in all life stages, there seems to be no empir- (Bremer support = 0.3; relative Bremer support = 14) ical basis for including gilli and Bruceharrisonius as and the two clades do not appear to be closely related subgenera of a single genus. They are distinct taxa based on an intuitive interpretation of morphological that merit equivalent rank. Consequently, we have data that is not included in the analysis. The pairing of chosen to elevate Bruceharrisonius to generic status Chaetocruiomyia and Macleaya is strongly supported with the concomitant recognition of gilli as a separate by 15 homoplastic characters (Bremer support > 0.4; genus. Reinert (2003) provided a full description and relative Bremer support> 100). These two taxa were illustrations of Bruceharrisonius (as a subgenus of originally described as genera (Theobald, 1910, 1903, Ochlerotatus). Himalaius, which includes simlensis in respectively), and later reduced to subgenera within addition to gilli, is described in Appendix 3. Aedes (Edwards, 1932). Despite the large number of The placement of Vansomerenis as the sister group of morphological features that distinguish them, includ- Zavortinkius + Kenknightia is well supported (Bremer ing a unique character among those that diagnose support> 0.4; relative Bremer support> 100), and the Chaetocruiomyia (115:3, interocular space with both monophyly of the former taxon is strongly supported broad and narrow scales), we propose to recognize (Bremer support > 0.4; relative Bremer support> 100) these lineages as subgenera of genus Macleaya. Taylor (Fig. 1A) by two characters, one of which (117:2, anten- (1929), Marks (1962, 1964), Dobrotworsky (1965) and nal pedicel without scales and/or setae) is unique and Lee et al. (1982, 1984) all reported the close similarity not contradicted (Fig. 2A). Additional morphological and relationship of Chaetocruiomyia and Macleaya in distinctions (not included in the analysis) of this group adult and immature stages as well as biology and dis- of Afrotropical species further corroborate its unique- tribution in the Australasian Region. Additionally, ness and support its recognition as a distinct lineage Reinert (2002e) pointed out the close similarity of within the Ochlerotatus genus-group. We adjudge this female genitalia of the two taxa. In addition to the lineage to be a genus based on the strength of the char- diagnostic combinations of characters arrayed on the acter support and the principle of equivalent rank. The branches in Fig. 2B, salient differences between the diagnostic and differential features that distinguish adults, larvae and pupae of Chaetocruiomyia and Vanso,nerenis are detailed in Appendix 3. Macleaya are listed in Marks (1962). In view of the Although support for the dade consisting of many morphological differences that distinguish these Oc. (Fin.) candidoscutellum + Oc. (Fin.) keefei taxa from Oc. (Fin.) loi, we regard the pairing of Oc. (Fig. 1A) is weak (Bremer support = 0.1; relative Bremer (Fin.) loi with genus Macleaya as an anomaly and support = 3), morphological observations (data not retain it in the non-monophyletic assemblage of Oc. included herein) suggest that this species pair is a nat- (Finlaya) pending further study. ural relationship. Furthermore, morphological data Reinert (1993) resurrected Molpemyia from suggest that this taxon may be related to the Oc. synonymy with Finlaya as a subgeneric category for (Fin.) biocellatus, but there is no evidence in the four three species of Aedes (auridorsum, pecuniosus, par- MPCs to unite these taxa (Figs 1A, 2A). The inclusion pureus), and Reinert (2000a) subsequently transferred ofOc. (Fin.) crossi in the biocellatus - seoulensis dade the subgenus to Ochierotatus. In the study of Reinert raises additional questions. We carefully examined et al. (2004), this taxon, represented only by pecunio- published descriptions and available material of all sus, was placed as sister to Fl. kochi + Fl. poicilia, with known species ofFinlaya and their apparent allies to Macleaya tremula as sister to these three taxa. determine which species might be closely related to Whereas a relationship between Finlaya s.s. and Oc. (Fin.) crossi based on morphological correspon- Macleaya is not evident in the present study, affinities

that may exist between Molpemyia and Finlaya sensu pedicels (character 117:1), which only occurs elsewhere auctorum within the trichotomy (Figs 1B, 2B) are insol- as independent derivations in the Stegomyia + Scuto- uble with the current data set. Although Oc. (Fin.) myia and Abraedes -Kompia clades oftheAedes genus- chungi is strongly supported as sister to Molpemyia group and Ochierotatus genus-group, respectively. (Bremer support> 0.4; relative Bremer support> 100), Dobrotworskyius is proposed for the dade repre- morphological differences not included in the analysis sented by alboannulatus, rubrithorax and tubbutien- indicate that chungi is not closely related to Molpe- sis (Figs 1B, 213). Although this genus is not as lnyia. Ochierotatus (Fin.) chungi and Molpemyia are strongly supported as most other genera of Aedini diagnosed by 16 and 19 homoplastic characters, respec- (Bremer support = 0.3; relative Bremer support = 18), tively (Fig. 2B), which exceed the number of such char- shared morphological characters not included in the acters that diagnose several traditionally accepted analysis indicate that these species form a natural genera, i.e. Armigeres (7), Eretmapodites (15), Haem- assemblage. The genus, which is confined to Australia, agogus (13), Heizmannia (8), Udaya (10) and Zeugno- also includes milsoni, occidentalis, rupestris and sub- myia (9) (Reinert et al., 2004: figs 5, 6). Therefore, the basalis (Appendices 3 and 4). recognition of chungi and Molpemyia as generic-level Georgecraigius is proposed as a genus for three spe- taxa is certainly equipollent. For these reasons, in addi- cies that occur in the Nearctic (atropalpus, epactius), tion to restoring Molpemyia to its original generic sta- Neotropical (epactius, fluviatilis) and Palaearctic tus based on shared diagnostic features observed in the (atropalpus) regions. These species were classified as three included species (see Reinert, 1993), a new genus, species of Aedes (Finlaya) until Zavortink (1972) Jihliensius, is proposed for chungi and two apparently assigned them to Ae. (Ochierotatus). Reinert (2000a) related species (Appendix 3). transferred them to Ochlerotatus (Ochlerotatus), and Phagomyia Theobald is resurrected from synonymy they were subsequently left without generic place- with Ftnlaya for the dade comprising assamensis, ment when Reinert et al. (2004) restricted the limits of gubernatoris, lophoventralis and prominens (Figs 113, Ochierotatus. The monophyly of Georgecraigius is 213). This dade is strongly supported (Bremer strongly supported (Bremer support > 0.4; relative support> 0.4; relative Bremer support> 100) by a Bremer support> 100) by a combination of 11 combination of eight homoplastic characters. Of these, homoplastic characters (Fig. 213). Nine homoplastic six (129:0, 131:1, 133:0, 135:0, 141:2 and 179:1) are characters diagnose flu viatilis and place it in a sister- common to all four MPCs, and the other two (10:1 and group relationship with atropalpus + epactius 166:0) appear only after the MPCs have been com- (Figs 113, 213). The atropalpus + epactius dade is diag- bined to form the SCT (see caveat on over-interpreta- nosed and distinguished from the fluviatilis dade by tion above). Phagomyia seems to be a homogeneous 20 homoplastic characters. It is noteworthy that atro- group that currently includes 16 species (see Appendix palpus and epactius were strongly paired (bootstrap 4). The genus was proposed originally for three nomi- value= 71) in the rDNA RFLP phylogeny of Kumar nal species (Theobald, 1905), including gubernatoris et al. (1998)]. In view of the number of characters that and two species now placed in Ae. (Aedimorphus). distinguish the two clades, we propose to recognize Culex gubernatoris Giles is the type species of Phago- them as separate subgenera. The atropalpus + epac- myia by subsequent designation (Edwards, 1932). tius dade is the nominotypical subgenus because atro- The monophyly of the dade comprising aihilabris + palpus is the type species of Georgecraigius. The notoscriptus is strongly supported by an extensive monobasic flu viatilis dade is named and described as combination of 16 homoplastic characters (Bremer Horsfallius subgen. nov. in Appendix 3. support > 0.4; relative Bremer support> 100) (Figs 113, Genus Patmarksia is established for Australasian 2B), hence the reason we have chosen to establish species that, for the most part, were included in the Pap- genus Rampamyia for this lineage within the Ochie- uensis Subgroup of the Alboannulatus Group of Knight rotatus genus-group. In addition to the two species & Marks (1952). The sister-group relationship of Pat- included in the analysis (Ra. albilabris and Ra. noto- marksia with Georgecragius is strongly supported by scripta), this taxon also includes the allied Ra. quin- seven homoplastic characters (Bremer support > 0.4; quelineata (see Appendix 4). These species occur in the relative Bremer support> 100) (Figs 1B, 2B). Although Australasian Region. Belkin (1962) placed albilabris monophyly of Patmarksia is rather weakly supported and notoscriptus in the Notoscriptus Subgroup of the by a combination of only four homoplastic characters Mediovittatus Group of Finlaya (as a subgenus of (Bremer support = 0.3; relative Bremer support = 34) Aedes) and noted that it probably should be recog- (Figs 113, 2B), it is further diagnosed and distinguished nized as a distinct group. Rampamyia, as defined here, from Georgecragius by a number of distinctive features includes a distinct part of Belkins Notoscriptus Sub- that were not included in the present data set (see group. This group is marked by the presence of a patch description in Appendix 3). Relationships of the four of broad, overlapping, silvery scales on the antennal species of Patmarksia included in the analysis indicate

Journal compilation © 2006 The Linnean Society of London, Zoological ,Journal of the Linnean Society, 2006, 148, 1-101 No claim to original US government works. PHYLOGENY AND CLASSIFICATION OF FINLAYA 37 the possible existence of two species-groups or subgen- strongly supported (Bremer support > 0.4; relative era within the genus, represented by the pairing of Bremer support> 100) (Fig. 1B) by five homoplastic argyronotum + papuensis and argenteitarsis + suhal- characters (Fig. 2B). Because, like Gilesius above, the hitarsis. However, because only five homoplastic char- monophyly of this dade is further supported by mor- acters support the latter dade, further analyses that phological data that are not included in the analysis, include several or all of the 13 species currently we have chosen to recognize it as a generic-level lin- assigned to Patmarksia (see Appendix 4) are required eage, genus Collessius, within the Ochierotatus genus- to corroborate the reality of these groups. group. In addition to the five species included in the Hulecoeteomyia Theobald is resurrected for the analysis, this taxon also includes four allied species dade comprising japonicus - sherki (Figs 113, 213). (Appendix 3). Members of the genus, which occur in Although the basal branch of the dade has only the Oriental Region and the Manchurian Subregion of weak support (Bremer support = 0.2; relative Bremer the Palaearctic, exhibit morphological differences, support = 10), the group appears to be a natural principally in the male and female genitalia, indica- assemblage of species that share a number of distinctive of subgroup affinities that only partially corre- tive morphological features (Fig. 2B), especially the spond with the pattern of relationships shown in the broad basal pale bands on hindtarsomeres 1-3 and the cladograms, i.e. hatorii + ((elsiae + macfarlanei) + development and positions of larval setae 4-7-C. With (hanksi + pseudotaeniatus)). Although the characters the exception of japonicus, all species of Hulecoet- that diagnose the elsiae + macfarlanei branch include eomyia included in the analysis exhibit character 95:0 a unique character (222:5, gonostylar claw with two (seta 641 shorter than seta 7-I1). Within Hulecoet- narrow, short to moderately long spiniforms inserted eomyia, japonicus is sister to the remaining taxa, distally) that is not contradicted (Fig. 213), other ana- which comprise two groups in a pectinate relationship tomical characters suggest that hatorii, and three spe- (Figs 113, 213). Because the relationships within Hule- cies not included in the analysis, are more closely coeteomyia are only weakly supported, it is not prac- allied to this species pair than to banksi and pseudo- tical to propose internal subdivisions until additional taeniatus. In view of the morphological contradictions, character data are available. it is not possible to divide the genus into subgeneric Gilesius pulchri venter comb. nov. shares a number groups; hence, two informal assemblages of species of characters with species of Hulecoeteomyia but the are recognized (Appendix 3) until the relationships of hindtarsomeres of the adults are dark-scaled and all species of Collessius are resolved. pupal seta 6-I1, based on the pupa of Gi. alius, which The monophyly of echinus + geniculatus is strongly Lien (1968b) originally described as a subspecies of supported (Bremer support > 0.4; relative Bremer pulchri venter, is significantly longer than seta 7-I1. support> 100) by a combination of ten homoplastic The sister-group relationship of pulchriventer and characters (Figs 1C, 2C) and otherwise forms a distinct Tanakaius (Figs 113, 213) is strongly supported element within the Eurasian aedine fauna. Edwards (Bremer support> 0.4; relative Bremer support> 100) (1932) included these species along with species from by a single character of the male genitalia (218:1, the Oriental Region and the New World in his Group H gonocoxite with specialized scales on the mesal mar- of Aedes (Finlaya). Although similarities have been gin of the ventral surface) that otherwise occurs in the noted between geniculalus and Oc. (Protomacleaya) distantly related Oc. (Fin.) biocellatus and the dade (e.g. Becker et at., 2003), the studies by Schick (1970) comprising Oc. (Fin.) harperi - Haemagogus. How- and Zavortink (1972) did not include or discuss this ever, the many morphological differences that distin- species in their revisions of species currently placed in guish these taxa indicate that they are not so closely Protomacleaya. In a biochemical systematics study related. Gilesius is diagnosed by a combination of 12 that included geniculatus and Oc. (Pro.) triseriatus, homoplastic characters (Fig. 213), which exceed the Munstermann (1980) reported that When enzyme number of such characters that define four tradition- migration rates of geniculatus and triseriatus were ally accepted genera, i.e. Armigeres (7), Heizmannia compared, only 7 of the 21 enzymes assayed migrated (8), Udaya (10) and Zeugnornyia (9) (Reinert et at., at similar rates, indicating that the two species are not 2004: figs 5, 6). Shared morphological characters not closely related. The results of the present study cor- included in the analysis also support the monophyly of roborate this observation. Based on these observations, the group (Appendix 3); thus, this dade is recognized the strength of character support (Figs 1C, 2C) and as a genus based on the degree of morphological dif- other distinguishing features not included in the anal- ferentiation and the principle of equivalent rank. ysis, generic-level status is proposed for the echinus + Gilesius + Tanakaius shares a weakly supported geniculatus dade. This genus is formally named and sister-group relationship with hatorii + elsiae - pseudo- described as I)ahliana in Appendix 3. taeniatus (Bremer support = 0.2; relative Bremer The pairing ofOc. (Och.) muelleri with deserticola - support = 7), but the monophyly of the latter taxon is varipalpus (in an unresolved relationship) is only

supported (Bremer support = 0.4; relative Bremer mated that Aedini should include about 87 genera. As support = 7) by a combination of six homoplastic char- a result of our present and previous studies, 63 genera acters (Fig. 2C), but it is strongly supported as sister are currently recognized within the tribe. Again, we toAbraedes + (Aztecaedes + (Gymnometopa + Kompta)) emphasize that this is a fundamental desideratum for (Bremer support> 0.4; relative Bremer support> 100) achieving a natural classification of Aedini. Two points (Figs 113, 2B). The three species that comprise the lat- support this statement. First, the results of Reinert ter dade belong to the Varipalpus Group of five tree- et al. (2004) and those of the present study show hole-breeding species that occur in western North remarkable congruence despite the disjunction America (Arnell & Nielsen, 1972). Belkin & McDonald between the number of taxa and morphological char- (1957) excluded muelleri from the Varipalpus Group acters included in the two studies. Second, we would (their varipalpus complex) and considered it an aber- note that the majority of subgenera recognized in the rant member of the pulchritarsis-varipalpus group traditional classification of Aedini (Edwards, 1932; based on differences in the scutal ornamentation of Belkin, 1962) are the same taxa that we recognize as the adults and the absence of teeth on the fore- and polythetically diagnosed genera, the difference being midungues of females. Zavortink (1972) pointed out that we choose to recognize explicitly monophyletic similarities between the adult, larval and pupal groups of generic rank rather than intuitively con- stages and distributions of muelleri and members of structed groups of lower rank within large paraphyl- the Varipalpus Group and placed it in a monobasic etic or polyphyletic (and hence unnatural) genera. group (Muelleri Group) allied with the Pulchritarsis Mitter (1999) appropriately stated: Intelligent and Varipalpus Groups in a more inclusive Varipalpus selection of the sample of species to be used for molec- Section. The results of the present study corroborate a ular study depends on the existence of a sound initial close relationship between muelleri and the Varipal- classification, which will almost always be based on pus Group. In as much as the muelleri - varipalpis lin- morphology. Morphological classifications will also eage is tantamount to other clades that we regard as continue to serve as the initial standard for gauging genera, we have chosen to identify this monophyletic which genes are suitable for solving a given problem. group as genus Jarnellius. Because a combination of Our proposed classification based on a comparative 16 homoplastic characters distinguishes muelleri as morphological study provides such a framework. 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 ACKNOWLEDGEMENTS formally recognized and described in Appendix 3. We gratefully acknowledge Mark Carine (Department of Botany, The Natural History Museum (NHM), Lon- FINAL COMMENTS don, UK) and M. Anice M. Sallum (Nücleo de Pesquisa Taxonômica e Sistemática em Entomologia Médica, This is the second study of a four-phase project that Faculdade de Saüde Püblica, Universidade de Sao aims objectively to assess the phylogenetic relation- Paulo, Brazil), for critically reviewing the manuscript ships and classification of Aedini. As with our previous and providing valuable comments. Gratitude is study (Reinert et al., 2004), the results and proposals expressed to Richard C. Wilkerson, James E. Pecor of the present investigation are likely to be viewed as and Thomas V. Gaffigan (Walter Reed Biosystematics controversial. In response to this, we unreservedly Unit, Smithsonian Institution, Washington, DC, state that we regard our reclassification of the tribe as USA), Theresa M. Howard (NHM), Desmond H. Foley a developing empirical hypothesis, subject to change and Greg Daniels (University of Queensland Insect and revision as more information becomes available. Collection, Department of Zoology and Entomology, As Zavortink (1979) aptly stated, A classification, University of Queensland, Brisbane, Australia), Jac- again like any other hypothesis, has heuristic value, quie Recsei and Don Colless (Australian National so that its very existence should stimulate further Insect Collection, CSIRO Division of Entomology, Can- research that may ultimately bring about its modifi- berra, Australia), Richard F. Darsie, Jr, Florida Med- cation or demise. With this in mind, it is worth reit- ical Entomology Laboratory, Vero Beach, FL, USA), erating that the reclassification of species into smaller Francis Schaffner (Laboratoire/Cellule Entomologie, and more numerous genera that indicate their genetic EID Méditerranée, Montpellier, France), Gary J. Steck [recte genealogical] relationships more accurately is a (Florida State Collection of Arthropods, Division of consequence of beta-level taxonomic study; and in Plant Industry, Gainesville, FL, USA), Vu Due Huong comparison with the classifications of other groups of (National Institute of Malariology, Parasitology and organisms based on beta taxonomy, the total number Entomology, Hanoi, Vietnam) and Thomas J. Zavort- of mosquito genera should be 225 (Zavortink, 1990). In ink (Bohart Museum of Entomology, University of Cal- proportion to the size of the family, we previously esti- ifornia, Davis, CA, USA) for the loan of specimens.

Journal compilation © 2006 The Linnean Society of London, Zoological Journal of the Linnean Society, 2006, 148, 1-01 No claim to original US government works. PHYLOGENY AND CLASSIFICATION OF FINLAYA 39

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Mosquito News 29: 545-548. of the American Entomological Institute (Gainesville) 32:45- Rees DM, Onishi K. 1951. Morphology of the terminalia and 52. internal reproductive organs, and copulation in the mos- Reinert JF. 2000f. Comparative anatomy of the female quito, Culiseta inornata [sic] ( Williston). Proceedings of the genitalia of genera and subgenera in tribe Aedini (Diptera: Entomological Society of Washington 53: 233-246. Culicidae). Part V. Genus Aedes Meigen. Contributions of the Reinert JF. 1976a. A ventromedian cervical scierite of mos- American Entomological Institute (Gainesville) 32: 1-102. quito larvae (Diptera: Culicidae). Mosquito Systematics 8: Reinert JF. 2001a. Revised list of abbreviations for genera 205-208. and subgenera of Culicidae (Diptera) and notes on generic Reinert JF. 1976b. Medical entomology studies - V. The sub- and subgeneric changes. Journal of the American Mosquito genus Rhinoskusea of the genus Aedes (Diptera: Culicidae). Control Association 17: 51-55. Contributions of the American Entomological Institute (Ann Reinert JF. 2001b. Comparative anatomy of the female gen- Arbor) 13: 1-60. italia of genera and subgenera in tribe Aedini (Diptera: Culi- Reinert JF. 1979. A description of Isoaedes, a new subgenus cidae). Part VI. Genus Ayurakitia Thurman. Contributions of of Aedes Meigen, and its type-species, Ac. (Isa.) cavaticus the American Entomological Institute (Gainesville) 32: 1-7. new species (Diptera: Culicidae). Mosquito Systematics 11: Reinert JF. 2001c. Comparative anatomy of the female gen- 144-162. italia of genera and subgenera in tribe Aedini (Diptera: Culi- Reinert JF. 1980. A literature review of pupal seta 13-CT in cidae). Part VII. Genus Opifex Hutton. Contributions of the the family Culicidae (Diptera) and a report of its occurrence American Entomological Institute (Gainesville) 32: 9-15. in Aedes aurotaeniatus Edwards. Mosquito Systematics 12: Reinert JF. 2001d. Comparative anatomy of the female gen- 441-444. italia of genera and subgenera in tribe Aedini (Diptera: Culi- Reinert JF. 1990. Medical entomology studies -XVII. Biosys- cidae). Part VIII. Genus Verrallina Theobald. Contributions of tematics of Kenknightia, a new subgenus of the mosquito the American Entomological Institute (Gainesville) 32:17-39. genus Aedes Meigen from the Oriental Region (Diptera: Culi- Reinert JF. 2001e. Comparative anatomy of the female cidae). Contributions of the American Entomological Insti- genitalia of genera and subgenera in tribe Aedini (Diptera: tute (Gainesville) 26: 1-119. Culicidae). Part IX. Genus Eretmapodites Theobald. Contri- Reinert JF. 1993. Redescription of Molpemyia, and its butions of the American Entomological Institute (Gainesville) revalidation as a subgenus of Aedes (Diptera: Culicidae). 32: 41-50. Mosquito Systematics 25: 41-63. Reinert JF. 2002a. A comparative analysis of the ventral Reinert JF. 1999a. The subgenus Rusticoidus of genus Aedes brush and its attachment to segment X of fourth-stage lar- (Diptera: Culicidae) in Europe and Asia. European Mosquito vae in tribe Aedini (Diptera: Culicidae). Journal of the Amer- Bulletin 4: 1-7. ican Mosquito Control Association 18: 16-25. Reinert JF. 1999b. Descriptions of Zovortinkius, a new sub- Reinert JF. 20021b. Comparative anatomy of the female gen- genus of Aedes, and the eleven included species from the Afro- italia of genera and subgenera in tribe Aedini (Diptera: Culi- tropical Region (Diptcra: Culicidae). Contributions of the cidae). Part X. Genus Heizmannia Ludlow. Contributions of American Entomological Institute (Gainesville) 31: 1-105. the American Entomological institute (Gainesville) 32: 1-15. Reinert JF. 2000a. New classification for the composite genus Reinert JF. 2002c. Comparative anatomy of the female gen- Aedes (Diptera: Culicidae: Aedini), elevation of subgenus italia of genera and subgenera in tribe Aedini (Diptera: Culi- Ochlerotatus to generic rank, reclassification of the other cidae). Part XI. Genus Haemagogus Williston. Contributions subgenera, and notes on certain subgenera and species. of the American Entomological Institute (Gainesville) 32: 17- Journal of the American Mosquito Control Association 16: 30. 175-188. Reinert JF. 2002d. Comparative anatomy of the female gen- Reinert JF. 2000b. Comparative anatomy of the female italia of genera and subgenera in tribe Aedini (Diptera: Culi- genitalia of genera and subgenera in tribe Aedini (Diptera: cidae). Part XII. Genus Armigeres Theobald. Contributions of Culicidae). Part I. Introduction, preparation techniques, and the American Entomological Institute (Gainesville) 32:31-46. anatomical terminology. Contributions of the American Reinert JF. 2002e. Comparative anatomy of the female Entomological Institute (Gainesville) 32: 1-18. genitalia of genera and subgenera in tribe Aedini (Diptera: Reinert JF. 2000c. Comparative anatomy of the female Culicidae). Part XIII. Genus Ochlerotatus Lynch Arrib- genitalia of genera and subgenera in tribe Aedini (Diptera: alzaga. Contributions of the American Entomological Insti- Culicidae). Part II. Genus Psorophora Robineau-Desvoidy. tute (Gainesville) 33: 1-111.

Reinert JF. 2003. Description of Bruceharrisonius, a new sub- Theobald FV. 1903. Two new Australian culicids. The Entom- genus of Ochlerotatus, and a redescription of its type species ologist 36: 154-157. Oc. (Brh)greenii. Journal of the American Mosquito Control Theobald FV. 1905. Genera insectorum. Diptera, fam. Culi- Association 19: 309-322. cidae. Fascicle 26, Belgium. Reinert JF. 2005. List of species described in the egg stage of Theobald FV. 1907. A monograph of the Culicidae or mosqui- tribe Aedini (Diptera: Culicidae) with their literature cita- toes. Mainly compiled from collections received at the British tions. Journal of the American Mosquito Control Association Museum, Vol. IV. London: British Museum (Natural History). 21:252-262. Theobald FV. 1910. A monograph of the Culicidae or mosqui- Reinert JF, Harbach RE, Kitching IJ. 2004. Phylogeny and toes. Mainly compiled from collections received at the British classification of Aedini (Diptera: Culicidae) based on mor- Museum, Vol. V. London: British Museum (Natural History). phological characters of all life stages. Zoological Journal of Tyson WH. 1970. Contributions to the mosquito fauna of the Linnean Society 142: 289-368. Southeast Asia VIII. Genus Aedes, subgenus Mucidus Ronderos BA, Bachmann AO. 1963. Mansoniini [sic] neo- Theobald in Southeast Asia. Contributions of the American tropicales. I. (Diptera-Culicidae) [sic]. Revista de la Sociedad Entomological Institute (Ann Arbor) 6: 28-80. Entomologia Argentina 26: 57-65. Villa AS-C, Rodriguez JAR, Llera JLG. 1985. Estudio de la Ross HH. 1947. The mosquitoes of Illinois (Diptera, Culi- larva de cuarto estado de las especies del subgenero Fznlaya cidae). Bulletin of the Illinois Natural History Survey (Diptera: Culicidae) [sic] de Espana peninsular. Sociedade (Urbana, Illinois) 24: 1-96. Portuguesa de Entomologia Suplemento 1: 2-12. Ross HH, Horsfall WR. 1965. A synopsis of the mosquitoes of Wesson DW, Porter CH, Collins FH. 1992. Sequence and Illinois (Diptera, Culicidae). Illinois Natural History Survey secondary structure comparisons of ITS rDNA in mosquitoes Biological Notes, Number 52: 1-50. (Diptera: Culicidae). Molecular lhylogenetics and Evolution Russell RC. 1996. A colour photo atlas of mosquitoes of south- 1: 253-269. eastern Australia. Sydney: Published by the author. Wood DM, Dang PT, Ellis RA. 1979. The insects and arach- Russell PF, West LS, Manwell RD, Macdonald G. 1963. nids of Canada Part 6. The mosquitoes of Canada, Diptera: Practical malariology, 2nd edn. London: Oxford University Culicidae. Publication 1686. Quebec: Canadian Government Press. Printing Centre. Schick RX. 1970. Mosquito studies (Diptera, Culicidae) XX. Yamaguti 5, LaCasse WJ. 1951. Mosquito fauna of North Amer- The Terrens Group of Aedes (Finlaya). Contributions of the ica Part V - Genus Aedes [sic] (with 116 original plates). Office American Entomological Institute (Ann Arbor) 5: 1-158. of the Surgeon, HQ. Japan Logistical Command, APO 343. Sirivanakarn S. 1976. Medical entomology studies - III. A Zavortink TJ. 1972. Mosquito studies (Diptera, Culicidae) revision of the subgenus Culex in the Oriental Region XXVIII. The New World species formerly placed in Aedes (Diptera: Culicidae). Contributions of the American Entomo- (Finlaya). Contributions of the American Entomological logical Institute (Ann Arbor) 12: 1-272. Institute (Ann Arbor) 8: 1-206. Smith A.B. 1994. Systematics and the fossil record: document- Zavortink TJ. 1979. Mosquito studies (Diptera, Culicidae) ing evolutionary patterns. Oxford: Blackwell Scientific XXXV. The sabethine genus Johnhelkinia and a preliminary Publications. reclassification of the composite genus Trichoprosopon. Tanaka K. 2002. Studies on the pupal mosquitoes of Japan (6) Contributions of the American Entomological Institute Aedes (Finlaya) (Diptera, Culicidae). Japanese Journal of (Ann Arbor) 17: 1-61. Systematic Entomology 8: 137-177. Zavortink TJ. 1985. Observations on the ecology of treeholes Tanaka K, Mizusawa K, Saugstad ES. 1979. A revision of and treehole mosquitoes in the southwestern United States. the adult and larval mosquitoes of Japan (including the In: Lounibos LP, Roy JR, Frank JH, eds. Ecology of mosqui- Ryukyu Archipelago and the Ogasawara Islands) and Korea toes. Proceedings of a Workshop Florida Medical Entomology (Diptera: Culicidae). Contributions of the American Entomo- Laboratory, Vera Beach, Florida, 473-487. logical Institute (Ann Arbor) 16: 1-987. Zavortink TJ. 1990. Classical taxonomy of mosquitoes - a Taylor FE. 1929. Notes on Australian Culicidae (Dipt). memorial to John N. Belkin. Journal of the American Mos- Bulletin of Entomological Research 20: 271-277. quito Control Association 6: 593-599.

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.

Genus Subgenus Species Distribution

Aedine ingroup taxa Downsiomyia Vargas leonis (Colless) nivea (Ludlow)

APPENDIX 1 Continued

Genus Subgenus Species Distribution

Finlaya Theobald kochi (Doenitz) 5 poicilia (Theobald) 4,5 Haemagogus Conopostegus Dyar leucocelaenus (Dyar & Shannon) 6 Williston leucotaeniatus (Komp) 6 Haeniagogus equinus Theobald 6 splendens Williston 6 Halaedes Belkin australis (Erichson) 5 Ochierotatus Lynch

Arribalizaga Finlaya Theobald albilabris (Edwards) 5 alboannulatus (Macquart) 5 aihocinctus (Barraud) 4 albotaeniatus (Leicester) 4 argenteitarsis (Brug) 5 argyronotum (Belkin) 5 assamensis (Theobald) 4 banksi (Edwards) 4 biocellatus (Taylor) 5 candidoscutellum (Marks) 5 chrysolineatus (Theobald) 4 chungi (Lien) 4 crossi (Lien) 4 echinus Edwards 2 elsiae (Barraud) 4 embuensis (Edwards) 3 geniculatus (Olivier) 2 gilli (Barraud) 4 gubernatoris (Giles) 4 harperi (Knight) 4 hatorii (Yamada) 2,4 ingrami (Edwards) 3 j. japonicus (Theobald) Jugraensis (Leicester) 4 keefei (King & Hoogstraal) 5 loi (Lien) 4 lophoventralis (Theobald) 4 luteostrjatus (Robinson) 3 macfarlanei Edwards 4 notoscriptus (Skuse) 5 oreophilus Edwards 2,4 papuensis (Taylor) 5 prominens (Barraud) 4 pseudotaeniatus (Giles) 4 pulchritorax (Edwards) 3 pulchri venter (Giles) 2,4 rubrithorax (Macquart) 5 seoulensis (Yamada) 2 sherki (Knight) 4 subalbitarsis (King & Hoogstraal) 5 tubbutiensis (Dobrotworsky) 5 Macleaya Theobald (Theobald) tremulus 5 Molpemyia Theobald pecuniosus (Edwards) 5 Ochlerotatus atropalpus (Coquillett) 1,2 deserticola (Zavortink) 1 epactius (Dyar & Knab) 1,6 fluviatilis (Lutz) 6 muelleri (Dyar) 1 sierrensis (Theobald) 1 varipalpus (Coquillett) 1 Protomacleaya Theobald berlini (Zavortink) 6 brelandi (Zavortink) 1 hurgeri (Zavortink) galindoi (Schick)

APPENDIX 1 Continued

Genus Subgenus Species Distribution

hendersoni (Cockerell) 1 homoeopus (Dyar) 6 knabi (Coquillett) 6 kompi (Vargas & Downs) 6 terrens (Walker) 6 triseriatus (Say) 1 zavortinki (Schick) 6 zoosophus (Dyar & Knab) 1

Opifex Hutton Nothoskusea Dumbleton chathamicus (Dumbleton) 5 Opifex fuscus Hutton 5 Tanakaius Reinert, savoryi (Bohart) 2 Harbach & Kitching togoi (Theobald) 1,2,4 Aedine outgroup taxa Abraedes Zavortink papago (Zavortink) 1 Aedes Meigen cinereus Meigen" 1,2 esoensis Yamada 2 Aztecaedes Zavortink ramirezi (Vargas & Downs) 1 Geoskusea Edwards baisasi (Knight & Hull) 4 longiforceps (Edwards) 5 Gymnometopa Coquillett mediovittata (Coquillett) 6 Howardina Theobald sexlineata (Theobald) 6 walkeri (Theobald) 6 Isoaedes Reinert cavaticus (Reinert)" 4 Kenknightia Reinert dissimilis (Leicester) 4 harbachi (Reinert) 4 Kompia Aitkin purpureipes (Aitken) 1 Levua Stone & Bohart geoskusea (Amos) 5 Mucidus Theobald alternans (Westwood) 5 laniger (Wiedemann) 4

Ochlerotatus Rusticoidus Shevchenko & refiki (Medschid) 2 Prudkina rusticus (Rossi) 2 Unassigned to subgenus communis (de Geer) 1,2 infirmatus Dyar & Knab 1 (Rondani) scapularis 1,6 Ochlerotatus Bruceharrisonius Reinert alektorovi (Stackelberg) 2 greenii (Theobald) 4 Chaetocruiomyia Theobald wattensis (Taylor) 5 Pseudoskusea Theobald hancroftianus (Edwards) 5 postspiraculosus (Dobrotworsky) 5 Theobald jamaicensis (Theobald) 6 Psorophora Robineau- Grahhamia Desvoidy Janthinosoma Lynch ferox (von Humboldt) 1,6 Arribaizaga Psorophora ciliata (Fabricius) 1,6 Rhinoskusea Edwards longirostris (Leicester) 4,5 wardi (Reinert) 4 Scutomyia Theobald albolineata (Theobald) 4,5 arboricola Knight & Rozeboom 4 Stegomyia Theobald aegypti (Linnaeus) 1,2,3,4,5,6 africana (Theobald) 3 Zavortinkius Reinert fulgens (Edwards) 3 longipalpis (Grunberg) 3 Non-aedine outgroup taxa Culex Linneaus Culex quinquefasciatus Say Culiseta Felt Culiseta inornata (Williston) 1 Mansonia Blanchard Mansonia titillans (Walker) 1,6 Orthopodomyia Theobald signifera (Coquillett) 1,6

Data matrix for 116 taxa and 232 characters used in the analysis. See text for explanation of coding. 9- - Ii 1111111111222222222233333333334 tITI 1234567890123456789012345678901234567890 I- 0 z LITI 0 Cs. (Gus.) inornata 1011121101012112021101011101110101111011 Cx. (Cux.) quinquefasciatus 1 0 0 1 1 2 1 1 0 1 0 1 2 1 1 2 0 2 0 1 0 1 0 1 0 1 0 0 0 0 0 1 1 0 1 0 1 0 1 1 0 Ma. (Man.) titillans 2211121110010111020110001110101101111010 tj Or signifera 0011001101112112010100001100110000101011 Ab. papago ? ? ? 1000000001001110010111010101000201002 Ae. cinereus 0111121101102112020100101101001101211001 Ae. esoensis ???1121101 1021120201001011010011012 11001 Az. ra,nirezi ???000000010100121000111011101010001010001 0 Ge. baisasi ???0111 101002113010101101100001111111011 0 Ge. longiforceps ???0121001 002113010101101100001111111011 0 Gy. mediovittata 01110000001020010100101110101010002 01002 0CI) 11w. sexlineata 01100100001120010100101111001010002 01002 Hw. walker ???00110021120010100101 10010101000201002 Ia. cavaticus ???0110001 10200101000010100001000001 0110001 0 Ke. dissimi/is ? ? ? 1 1 1 1 0 0 1 1 0 2 1 1 2 0 1 0 1 0 0 1 1 1 0 1 0 010 011 0 1 1 1 1 0 1 1 0 Ke. harbachi ? ? ? 1 1 1 1 0 0 1 1 0 2 0 1 2 0 1 0 1 0 0 1 1 1 0 1 0 0 0 011 0 1 1 1 1 0 1 1 0 0- 0 Ko. purpureipes ? ??0001000102001000010001010101010201002 0 Le. geoskusea ? 11011 1 0 0 0 1 0 2 1 0 2 0 2 0 1 0 ? 0 0 1 100001101111010 0N Mu. alternans 011112100102201002110010110000010 1010110 0 Mu. laniger ? ? 71121001022010021100101100000101010110 Oc. (Brh.) alektorovt ???111 1001102113010101100110010101111011 0 Oc. (Brh.) greenii ? ? ? 1 1 1 1 0 0 1 1 0 2 1 1 3 0 1 0 1 0 1 1 0 0 1 1 0 0 1 011 0 1 1 1 1 0 1 1 Oc. (Cha.) wattensis ? ? ? 0 0 0 0 0 0 0 1 0 1 0 0 1 0 1 1 0 0 0 0 0 1 1 1 0 0 1 0 0 1 0 1 0 1 0 0 1 0 Oc. (Och.) com,nunis 0 1 1 0 1 1 1 1 0 0 0 0 2 0 1 2 0 1 0 010 1 1 0 1 1 1 0 0 1 0 1 0 1 1 011 0 1 1 Oc. (Och.) infirmatus 0111121100102012010000101100010101111010 00 r Oc. (Och.) scapularts 0111111000102002010000101100010101111011 Oc. (Psk.) bancroftianus ? ? ? 0 1 2 1 1 0 1 0 0 3 0 1 3 0 2 0 1 01? 1 1 1 1 0 0 0 1 0 1 0 0 1 0 1 0 1 0 Oc. (Psk.) postspiracu/osus ???0121 101003013020101111100000100101011 0 Oc. (Rus.) re/Iki ???1021 101002112010001101111011101111011 Oc. (Rus.) rusticus 0 1 1 1 012 1 1 0 1 0 0 2 1 1 2 0 1 0 0 0 1 1 0 1 1 1 1 0 0 1 1 0 1 1 1 1 0 1 1 •0 0 0 Ps. (Gra. ) jarnaicensis 0121121101002112020100001101100101111011 0CI) Ps. (Jan.) ferox 0 1 2 1 1 2 1 1 0 1 0 0 2 1 1 2 0 2 0 1 010 0 0 1 1 1 1 1 0 1 1 1 1 1 1 1 0 1 1 C . CJ). Ps. (Pso.) ciliata 0121020000131102121100001100101101111011 Rh. /ongrostris 0 1 1 0 1 1 1 1 1 1 1 0 2 1 1 3 0 2 0 1 010 0 1 1 1 0 0 0 0 0 1 0 1 1 011 0 1 0 0 Rh. wardi ???0121 1111021130201000111000001 01111010 09)•1 Sc. albolineata ???0011 101012102010010111010101 000101002 Sc. arbonicola ???00110010121020100101 10010101 000001002 -Qo St. aegypti 0 1 1 1 0 0 0 0 0 1 0 0 2 0 0 1 1 1 0 0 1 0 1 0 1 0 1 0 1 0 010 0 0 1 0 1 0 0 12 St. africana 0I 01110000010020010100101010010100000101001 00 APPENDIX 2 Continued ,. 0 05 1 1 1 1 1 1 1 1 1 1 2 2 2 2 2 2 2 2 2 2 3 3 3 3 3 3 3.3 3 3 4 .5. 1234567890123456789012345678901234567890 5• Zn. fulgens ? ? ? 1 1 1 1 0 0 1 1 0 2 0 1 2 0 1 0 1 010 0 0 1 1 1 0 1 0 1 1 0 1 1 1 1 0 1 1 cJ] Za. longipalpis 01 1 1111001102012010100001 110101 101111011 0 Do. leonis ???01111011121110200101 11010101100111012 0 Do. nivea ???0111101 112111020010111010101100111012 50 Fl. kochi 0 ? ? 0 0 0 1 0 0 2 112001011010110010111000201012 Fl. poicilia ???00010021 12001011010110010111000201012 Ha. australis 0110001001102001010001101010000111111011 <0 Hg. (Con.) leucocelaenus 0111011000012001010010111100101000001012

CJ) Hg. (Con.) leucotaeniatus ? ? ? 1 0 1 1 0 0 0 0 1 2 0 0 1 0 1 0 0 1 0 1 1 1 1 0 0 1 0 1 0 0 0 010 1 0 1 2 C Hg. (Hag.) equinus 0 1 1 1 0 1 0 0 0 0 0 0 2 0 0 1 011 0 0 1 0 1 1 1 1 0 0 1 0 1 0 0 0 1 0 1 0 1 2 0 Hg. (Hag.) splendens 0 ? ? 1 0 1 0 0 0 0 0 0 2 0 0 1 0 1 0 0 1 0 1 1 1 1 1 0 1 1 1000101012 0 Oc. (Fin.) albilabris ? ? ? 1 0 0 0 0 0 0 1 0 2 0 0 1 0 1 0 0 0 1 1 0 1 1 0 0 1 0 0 1 0 011 011 0 1 1 Oc. (Fin.) aihoannulatus 0111121100101101010000111 1001 11101111011 0 Oc. (Fin.) albocinctus ???1121101122 112020110101010111110111012 C Oc. (Fin.) albotaeniatus ???00001000120101010010110111111000201012 Oc. (Fin.) argenteitarsis ???101101001Q11011000101 11110101101111011 N C 0 Oc. (Fin.) argyronotum ? ? ? 1011100101101210000111100010100101010 CR Oc. (Fin.) assamensis ???10000001020010000001 11100101100111012 Oc. (Fin.) banksi ? ? ? 1 0 1 1 0 0 1 0 0 1 1 0 1 2 1 0 0 0 0 1 1 1 1 0 0 011 0 0 0 0 1 0 1 0 1 1 C Oc. (Fin.) hiocellatus ? 7 ? 1 1 2 1 1 0 1 1 2 2 1 1 2 0 2 0 1 0 0 1 0 1 0 1 0 1 1 0 1 0 1 1 1 1 0 1 01 Oc. (Fin.) candidoscutellum ???11210011020120101101111 10100101 111011 1 Oc. (Fin.) chrysolineatus ???01110101 1011012100101111101110100101011 Oc. (Fin.) chungi ???100100010200100000011110010000011101 1 Oc. (Fin.) crossi ???1121101002 112020101101010101100111011 0 Oc. (Fin.) echinus ? ? ? 1 1 0 1 0 0 1 0 1 2 0 0 2 0 1 0 0 1 ? 1 1 1 1 10111000101012 N Oc. (Fin.) elsiae ???112110110110121001011111011010010101 1 ci) Oc. (Fin.) emhuensis ? ? ? 1 1 1 1 010 1 1 1 2 1 0 2 0 2 0 1 0 1 1 1 1 0 1 0 1 0 1 010 0 1 0 1 0 1 2 Cl) C Oc. (Fin.) geniculatus 0110101001012001010010111110111000201012 Zs Oc. (Fin.) gilli ? ? ? 1111000103013020110100010001111111011 0 Oc. (Fin.) guhernatoris ???1101001102001000000101100101101 111011 Oc. (Fin.) harperi ? ? ? 10000010120010100101 11110111000101012 Oc. (Fin.) hatorii 0 1 1 1 1 2 1 1 0 1 1 0 1 1 0 1 2 1 0 0 010 1 1 1 1 1 0 011 0 1 0 0 1 0 1 0 1 1 C z Oc. (Fin.) ingrami 0 1 1 1 1 2 1 1 0 1 1 1 2 1 0 2 0 2 0 1 011 1 1 1 0 1 0 1 0 1 1 0 0 1 0 1 0 1 2 C Oc. (Fin.) j. japonicus 0 1 1 0 1 2 1 010 1 1 0 1 1 0 1 2 1 0 0 0 0 1 1 1 1 1 0 011 010 0 0 1 0 1 0 1 01 Oc. (Fin.) Jugaraensis ? ? ? 0111000101101210010111110110101111011 Oc. (Fin.) keefei ? ? ? 1 1 2 1 0 0 1 1 0 2 0 0 2 2 2 0 1 0 0 1 0 1 0 1 0 1 0 1 1 0 1 1 1 1 011 1 Oc. (Fin.) loi ? ? ? 0 1 0 0 010 1 1 0 2 0 0 1 0 1 1 0 0 0 1 1 1 1 0 0 1 0 0 1 0 0 1 1 1 0 1 1 CD Oc. (Fin.) lophoventralis ???1101001102001000000111100101101111011 Oc. (Fin.) luteostriatus ? ? ? 1 1 1 1 0 0 1 1 0 2 011 2 0 1 0 1 010 1 0 1 0 1 0 1 0 1 1 0 1 1 1 1 0 1 1 APPENDIX 2 Continued

1111111111222222222233333333334 0 12345678901234 56789012345678901234567890 tTj I- 0 Oc. (Fin.) macfarlanet ? ? ? 1 1 2 1 0 0 1 1 0 1 1 0 1 2 1 0 0 0 0 0 1 1 1 1 0 1 1 0 1 0 0 1 0 1 0 1 01 z tTj 0 Oc. (Fin.) notoscriptus 0 1 1 1 1 0 0 0 0 0 1 0 2 0 0 1 0 1 0 0 011 1 0 1 1 0 0 1 0 1 1 0 0 1 0 1 0 1 1 Oc. (Fin.) oreophilus ? ? ? 1 1 0 1 0 0 0 1 0 2 0 0 120 1 0 0 0 0 1 0 1 1 1 0 1 0 1 1 0 0 1 011 0 1 2 H C Oc. (Fin.) papuensis ? ? ? 1 1 2 1 010 0 1 0 1 1 0 1 2 1 0 0 0 0 1 1 1 1 0 0 011 0 1 0 0 1 0 1 0 1 0 tz1 Oc. (Fin.) prominens ???1101001102001000000101100101101111011 ND Oc. (Fin.) pseudotaeniatus ???101 1001001101210000111100110000101011 O) Oc. (Fin.) pulchrtthorax ? ? ? 1 1 1 1 010 1 1 0 1 1 0 2 0 1 0 1 010 1 0 1 0 1 0 1 0 1 1 0 1 1 1 1 0 1 2 -3 Oc. (Fin.) pulchri venter ? ? ? 1 1 1 1 1 0 1 1 0 2 1 0 1 2 1 0 0 010 1 1 1 1 1 0 1 1 1 0 0 0 1 0 1 0 1 2 C Oc. (Fin.) rubrithorax 011 1 0 2 0 0 0 0 1 0 1 0 0 1 0 1 0 1 ? ? 1 ? ? 1 0 0 0 ? ? ? 0 ? 1 ? 1011 Oc. (Fin.) seoulensis 0 1 ? 1 1 2 1 0 0 0 1 1 2 1 0 2 0 2 0 1 1 0 1 1 1 0 1 0 1 0 1 010 0 010 1 0 1 2 C C Oc. (Fin.) sherki ???0111101101101210010111110101000 101012

U) Oc. (Fin.) subaihitarsis ? ? ? 1 1 2 1 010 0 1 0 1 1 0 1 1 1 0 0 0 0 7 1 1 1 0 0 1 1 1 0 0 0 1 0 1 0 1 2 C 0 Oc. (Fin.) tubbutiensis ? ? ? 1121100101111010000111100111101111011 C Oc. (Mac.) tremulus ???010100010210101000000101011001011101 1 -,C Oc. (Mo!.) pecuniosus ???10000001010 1100000010110010 1101111010 C Oc. (Och.) atropalpus 0 1 1 0 1 1 1 010 1 1 0 2 0 0 1 011 0 0 0 0 1 0 1 1 1 0 011 1 0 0 0 1 011 0 1 1 0 Oc. (Och.) deserticola ???0001000002001010001101110100000100011 C Oc. (Och.) epactius 0 1 1 0 1 1 1 0 0 1 1 0 2 0 0 1 0 1 0 0 0 0 1 0 1 1 1 0 0 0 1 0 0 0 1 011 0 1 1 0N Oc. (Och.) fluviatilts 01111110101101101110000111101010100101011 0 Oc. (Och.) muellert ???0111000012002010001100010101000101 002 0 Oc. (Och.) sierrensis 0 1 1 1 1 0 1 0 0 0 0 0 2 0 0 1 0 1 0 0 0 1 1 0 1 0 1 0 1 0 0 010 0 1 0 1 0 1 1 C Oc. (Och.) varipalpus 0 1 1 1 0 0 1 0 0 0 0 0 2 0 0 2 0 1 0 0 0 1 1 0 1 1 1 0 1 0 010 0 0 1 0 1 0 0 1 C Oc. (Pro.) her/ii ???1111000102 102010000101110111101111011 •1 Oc. (Pro.) brelandi ???1101000012101010000101110101 100111011 zo Oc. (Pro.) burgeri ???101 1000102001010000101010111100111011 Oc. (Pro.) ga/indot ? ? ? 1111000102102010010101110101101111011 Oc. (Pro.) hendersoni 01111010000120010100001Q1010101100111011 0 Oc. (Pro.) ho,noeopus 7 ? ? 1 0 1 1 0 0 0 1 0 2 010 2 0 1 0 0 0 0 1 0 1 1 1 0 1 1 1 1 0 0 1 1 1 0 1 1 Oc. (Pro.) knabi ? ? ? 1 0 1 1 0 0 0 0 0 2 0 0 1 0 1 0 01010 1 0 1 1 0 0 1 0 1 1 0 1 1 1 1 0 1 1 C . C Oc. (Pro.) kompi ???1111000102002020000101 110101100111011 CC) —o Oc. (Pro.) terrens 0111011000102111010000101110111101111011 0. in Oc. (Pro.) triseriatus 0 1 1 1 0 0 1 0 0 0 011 2 0 0 1 0 1 0 0 1 0 1 1 1 0 1 0 1 1 1 0 0 0 2 0 1 0 1 2 0 Oc. (Pro.) zavortinki ???111 1000102112010100101110 1 11101111011 CD •1 © Oc. (Iro.) zoosophus ???10010000020010100001110101 11100101001 Op. (Not.) chathamicus ???0121101 102112010101101010101101110010 Op. (Opi.)fuscus 0 1 1 0 0 0 0 0 0 1 1 0 2 0 0 1 0 0 0 0 0 1 1 0 1 0 1 0 0 0 0 1 0 1 1 1 0 011 0 Ta. savoryi ? ? ? 0 0 2 1 010 1 1 0 1 1 0 1 2 1 0 0 0 0 1 0 1 0 1 0 010 1 1 1 0 1 0 1 0 1 1 OP To. togoi 0110021101101101210000101010001110111011 zc-

APPENDIX 2 Continued 0 0 t 4444444445555555555666666666677777777778 1-. 1234567890123456789012345678901234567890 0 Cs. (Cus.) inornata 0 1 1 1 1 011 1 2 0.0 0 0 2 1 2 0 0 2 1 0 1 0 1 1 0 1 0 2 1 1 0 0 1 2 1 1 1 0 0 CID Cx. (Cux.) quinqueftisciatus 0101101110000002002001011210210000201102 0 < o Ma. (Man.) titillans 1110110111100112101110000101010001211102 Or. signifera 0101110100010202000001000110210000201100 Ab. papago 120100102000000101020000011101000003 10001 Ae. cinereus 0 2 0 1 1 1 0 1 0 0 0 0 1 0 1 2 1 0 121 010 0 1 2 2 1 0 1 0 0 0 0 0 2 1 1 1 0 1 0 Ae. esoensis 0201110110001012101100012210100000211101 Az. ramirezi 1 2 1 1 0 011 0 2 0 1 0 0 0 0 2 0 0 2 0 0 0 0 1 1 1 1 0 1 0 0 0 0 1 3 1 0 0 0 1

J] Ge. baisasi 01111111000010012001001011110200000201101 Ge. longiforceps 0 1 1 1 1 1 0 1 0 0 0 0 1 0 0 2 010 1 0 0 1 0 1 1 1 1 0 2 0 0 0 1 0 2 0 1 1 0 1 Gy. mediovittata 1201101010100201001000001110100101301000 0 Hw. sexlineata 12011010111002000010000111101001013 01001 11w. walkeri 1211101011 1002000010000011201001003 01101 Ia. cavaticus 0 0 1 1 0 1 0 1 1 0 0 0 010 0 120 0 1 0 0 1 0 1 2 1 1 0 1 0 0 0 0 0 1 0 1 0 0 1 Ke. dissimilis 0101111120011102001001011120100000111101 Ke. harbachi 0101111120011112101101011120100000111101 Ko. purpureipes 1211101020100200002000011110100001310101 La. geoskusea 0110111100011112101001011110200010211101 Mu. alternans 0 0 0 0 0 0 0 1 0 0 0 0 010 0 2 0 1 2 0 011 0 1 121 1 0 1 0 0 0 0 0 3 1 1 1 0 2 Mu. laniger 0000000100000002011001012110100000311102 Oc. (Brh.) alektoroui 01001110120001112101111011110100000211001 Oc. (Brh.) greenii 0101110120001212101111011110100000211101 Oc. (Cha) waltensis 00010111200000020020010011101000001 11001 Oc. (Och.) communis 0 1 1 1 1 011 1 1 0 0 0 1 0 0 2 0 0 2 0 0 1 0 1 1 1 1 0 2 0 1 0 0 0 2 1 1 0 0 1 Oc. (Och.) infirmatus 0 1 0 1 1 010 1 2 0 0 0 1 1 0 2 0 0 1 010 0 0 1 1 1 1 0 1 1 1 0 0 0 2 0 1 1 0 1 Oc. (Och.) scapu/aris 0 1 0 1 1 0 0 1 2 0 0 0 010 0 2 0 0 1 0 0 1 0 1 1 1 1 0 1 1 1 0 0 0 2 0 1 1 0 1 Oc. (Psk.) bancroflianus 011100012000020200200011110201000201101 Oc. (Psk.) postspiraculosus 0 011 1 0 0 1 1 2 0 0 0 0 2 0 2 0 0 2 0 0 1 0 1 1 1 1 0 2 0 1 0 0 0 2 0 1 1 0 1 Oc. (Rus.) refiki 0101111120001102001000012210201000211101 Oc. (Rus.) rusticus 0111111120001102001000012210201000211101 Ps. (Gra. ) jamaicensis 0 1 0 1 0 1 1 1 1 0 0 0 1 0 1 2 010 0 1 010 1 1 1 1 2 0 1 1 0 0 0 0 1 1 1 1 0 0 Ps. (Jan.) ferox 0 1 0 1 0 1 1 1 1 0 0 0 1 0 1 2 1 0 0 1 1 0 1 1 1 1 2 0 1 1 0 0 0 0 1 1 1 1 0 01 Ps. (Pso.) ciliata 0 1 0 1 0 011 1 0 0 0 0 1 0 1 2 1 0 0 1 1 1 1 1 2 1 2 0 1 1 0 0 0 0 1 1 1 1 0 0 Rh. longirostris 0 010 1 1 0 1 1 1 0 0 0 011 1 2 0 0 1 0 0 1 0 1 1 1 1 0 2 0 0 0 0 0 1 0 1 1 0 2 Rh. wardi 0 010 1 1 0 1 1 1 0 0 0 1 1 1 2 0 0 1 0 0 1 0 1 1 1 1 0 2 0 0 0 0 0 1 0 1 1 0 2 Sc. aiholineata 1201111011100100001000011110100100301101 Sc. arboricola 1 2 0 1 1 0 1 0 1 1 1 0 0 1 0 0 0 0 1 0 0 0 011 1 1 1 0 1 0 0 1 0 0 3 0 1 1 0 1 St. aegypti 1 2 1 1 0 1 1 012 0 0 0 0 1 0 1 0 0 1 0 0 0 0 0 121 1 0 1 0 0 0 0 0 1 0 1 0 0 1 St. africana 1201111020000100002000001110100000101001 C)1 APPENDIX 2 Continued 0 0 4444444445555555555666666666677777777778 0 1234567890123456789012345678901234567890 I1 0 z 0 Za. fulgens 0111111120001112101101001110100100211101 Za. longipalpis 011 1111120001112101101001110100000211101 0 Do. leonis 1 2 0 1 1 1 1 0 2 0 1 0 1 0 0 2 0 0 120 0 0 0 1 1 1 1 0 1 0 0 1 0 0 2 1 1 1 0 1 t1 0 Do. nivea 1 2 0 1 1 1 1 0 2 0 1 0 1 0 0 2 0 0 120 0 0 0 1 1 1 1 0 1 0 0 1 0 0 2 1 1 1 0 1 © Fl. kochi 1201001121100112002001011 121100100301012 Fl. potcilia 1211001121100112002001011121100100301012 0) Ha. australis 0111111100011012100111001110100010201101 0 Hg. (Con.) leucocelaenus 1 2 0 1 1 0 1 0 120 1 0 0 2 0 1 0 0 2 0 0 0 0 1 1 1 1 0 1 0 0 1 0 0 3 0 1 0 0 1 Hg. (Con.) leucotaentatus 1201101020100201002000011110100100301001 0 0 Hg. (Hag.) equtnus 1201101020100200002000011110100100301001 Hg. (Hag.) splendens 1201101020100200002000011110100100301001 0 U) Oc. (Fin.) albilabris 0 1 1 1 1 1 11 2 0 0 0 010 012 0 0 2 0 0 1 0 1 1 1 1 0 1 0 1 1 0 0 2 0 1 1 0 1 0 Oc. (Fin.) alboannulatus 01111111100011020010010111201000002011?? 0 Oc. (Fin.) albocinctus 0201101020001012001001011120100000111101 1211011121000202 00100 1012110100100301000 -,0 Oc. (Fin.) albotaentatus Oc. (Fin.) argenteitarsis 111 1111120001002001000011110100000201001 0 0 Oc. (Fin.) argyronotum 0101101110001102001001011120100100201101 0 0 0211111110001002001001011110100000311001 0 Oc. (Fin.) assamensis Oc. (Fin.) hanksi 0111011 120000102001001011110100100201001 0 Oc. (Fin.) hiocellatus 0101111120001012002001011120100000210101 Oc. (Fin.) candidoscutellum 0111110120011002102101011110100000211101 1211011 120000202001001011110100100201101 0 Oc. (Fin.) chrysolineatus Oc. (Fin.) chungi 0111111110001002001001011110100100211001 Oc. (Fin.) crossi 0101111120001002002000011110100000111001 Oc. (Fin.) echinus 12110010211102020010000111101000002110?? Zn Oc. (Fin.) elsiae 0111001120000202001001011110100100201 101 0n Oc. (Fin.) embuensis 1201111020000102002000011110100000111001 50 Oc. (Fin.) geniculatus 12 11001011110102001000011110100000211001 ...t-. Oc. (Fin.) gilli 0111111120001 112101111011110100000211101 0 0 Oc. (Fin.) gubernatoris 0111111110001002001001011110100000211101 Oc. (Fin.) harperi 1211011021101112001000012110100100111001 0 011 1 0 011 1 1 0 0 0 0 1 012 0 0 0 0 0 1 0 1 1 1 1 0 1 0 0 1 0 0 2 0 1 0 0 1 cn Oc. (Fin.) hatorti Oc. (Fin.) ingrami 1201111020001102002000011110100000111001 cn Oc. (Fin.)j. japonicus 0 011 1 0 1 1 1 2 00 0 0 2 0 2 0 0 2 0 0 1 0 1 2 1 1 0 1 0 0 1 0 0 2 0 1 0 0 1 Oc. (Fin.) jugaraensts 11110011120000202001001011110100100201001 CD Oc. (Fin.) keefei 11011011200111020020000? 1120100100211101 Oc. (Fin.) loi 011 1 1 1 1 1 1 1 0 0 0 0 0 0 2 0 0 1 0 0 1 0 1 1 1 1 0 1 0 0 1 0 0 1 1 1 0 1 1 Oc. (Fin.) lophoventralis 011111112000011210100101 1110100000211001 Oc. (Fin.) luteostriatus ?1 11111110001012102101011110100000111101 zc-

APPENDIX 2 Continued -0 .9. 4444444445555555555666666666677777777778 12345678901234 5678901234 5678901234567890

Oc. (Fin.) macfarlanei 0111001120000202000001011110100100201101 cn Oc. (Fin.) notoscriptus 011 1 1 1 1 1 1 1 0 010 010 0 2 0 0 2 0 0 1 0 1 1 1 1 0 1 0 1 1 0 0 2 0 1 1 0 1 o Oc. (Fin.) oreophilus 0211111010001002001001011110100000211001 Oc. (Fin.) papuensis 0111101120001002001000011121100100201101 Oc. (Fin.) prorninens 01111111200010020010010111101000002 11001 Oc. (Fin.) pseudotaeniatus 0111011120000102001001011110100100201001 Oc. (Fin.) pulchrithorax 011 1 1 1 1 1 1 2 0 0 0 1 1 1 2 1 0 2 1 0 1 0 1 1 1 1 0 1 0 0 0 0 0 121 1 1 0 1 Oc. (Fin.) pulchriuenter 011 1 1 0 1 1 1 2 0 0 0 0 2 0 2 0 0 2 ? 0 1 0 1 1 1 1 0 1 0 0 1 0 0 2 0 1 0 ? ? WCJ) 0 Oc. (Fin.) rubrithorax 0 1 1 0 1 1 0 1 ? 0 0 0 1 0 1 2 1 0 1 0 0 1 0 1 1 1101000002111?? Oc. (Fin.) seoulensis 0201101020001 10200100101111010000011100? Oc. (Fin.) sherki 22110011110101202001001012110100100201101 0 Oc. (Fin.) subalbitarsis 2111111120001102001001011110100000201101 0 Oc. (Fin.) lubbutiensis 0111111110011112001001011110100000201101 0 0 Oc. (Mac.) tremulus 0011011020000002002000101110100000101001 0 Oc. (Mol.) pecuniosus 0 1 1 0 0 0 0 1 0 0 0 0 010 0 2 0 0 1 0 0 0 0 0 1 1 1 0 1 0 0 0 0 0 3 0 1 0 0 1 0N Oc. (Och.) atropalpus 0 1 1 1 0 011 1 0 0 0 011 0 0 2 0 0 1 0 0 1 0 1 2 1 1 0 1 0 0 0 1 0 2 0 1 0 0 1 0 Oc. (Och.) deserticola 1101011020000102002000011110100010111001 CR Oc. (Och.) epactius 0111001100001002001001012110100010201001 Oc. (Ochj fluviatilis 01111011200001020010010111101000010201101 Oc. (Och.) inuelleri 012 1 1 1 0 1 0 2 0 0 0 0 0 0 2 0 0 2 0 0 0 0 1 1 1 1 0 1 0 0 0 0 0 2 1 0 0 0 1 0 Oc. (Och.) sierrensis 011 1 1 0 1 1 0 2 0 0 0 0 1 0 2 0 0 2 0 0 0 0 1 1 1 1 0 1 0 0 0 1 0 1 1 1 0 0 1 Oc. (Och.) varipalpus 011 1 1 0 011 1 2 0 0 0 0 1 0 2 0 0 2 0 0 0 0 1 1 1 1 0 1 0 0 0 1 0 1 1 1 0 0 1 Oc. (Pro.) herlini 0101111020001102001001011110100000211101 Oc. (Pro.) brelandi 1201111020001102001000011110100000211001 Oc. (Pro.) burgeri 0201111020001002001001011110100000211001 Oc. (Pro.) galindoi 1201111020001002001001011110100000211101 Oc. (Pro.) hendersoni 1201111020001102001000011110100000111001 Oc. (Pro.) homoeopus 11011110200010020010010111101000002 11101 C!) 0 Oc. (Pro.) knabi 1101111120001102001001011110100000211101 Oc. (Pro.) kompi 0201111020001102002001011110100000211101 Oc. (Pro.) terrens 0201111020001012101101011110100000211101 Oc. (Pro.) triseriatus 12011110200011020010000111101000001211101 C) Oc. (Pro.) zavortinki 0 1 0 1 1 1 1 1 2 0 0 0 1 0 1 2 010 1 0 0 1 0 1 1 1 1 0 1 0 0 0 0 0 2 1 1 1 0 1 Oc. (Pro.) zoosophus 1201111020001102001000011110100000211001 Op. (Not.) chathamicus 0 1 1 1 1 1 1 1 0 0 0 1 . 1 0 1 2 1 0 1 1 1 1 0 1 1 1 1 0 1 0 0 0 1 0 1 0 1 1 0 1 Op. (Opi.) fuscus 01110111000110121001110011201000010101100

I- Ta. savoryi 0111011100011102001001011110100110201101 I- Ta. togoi 0111011120010102001001011110100110201101

01 C)1 ND APPENDIX 2 Continued 9- C 111111111111111111111 TI 8888888889999999999000000000011 111111112 TI 1 2 3 4 5 6 7 8 9 0 1 2 3 4 5 6 7 8 9 0 1 2 3 4 5 6 7 8 9 0 1 2 3 4 5 6 7 8 9 0 —4 C z 1 1 1 0 0 0 1 1 1 1 011 0 0 1 1 1 1 0 0 0 120 2 1 0 0 0 1 1 0 0 0 2 Cs. (Gus.) inornata 1 0 1 1 0 0 H 0 Cx. (Cux.) quinquefasciatus 111100111000 10111100 1111000 112 100 01100 02 Ma. (Man.) titillans 00100000101101100111?- 1001? 0021000110102 Or signifera 0011000110000010101111110002021000110002 Ab. papago 0010110010110010000111101101020011211002 Ae. cinereus 0010000110100010110111010001022001110002 0010000110100010110111010001022001110002 C Ac. esoensis t7 4 Az. ramirezi 0010010010010000001111100101020011111002 Ge. haisasi 0 1 2 1 0 0 1 1 1 010 0 0 0 1 0 1 1 0 1 1 1 1 1 0 0 0 1 0 2 0 0 1 0 0 1 0 0 0 2 C C Ge. longiforceps 0120011111000010110110010011020010010001 Gy. mediovittata 1010010010010000001110100011022011201002 0CD 1 1 1 0 0 1 0 1 1 0 1 1 010 1 0 0 1 1 1 1 1 1 1 0 0 0 1 0 1 2 0 2 0 1 1 0 0 0 2 C) Hw. sexlineata C Hw. walkeri 1 1 1 010 1 011 1 0 0 1 1 0 1 010 1 1 1 1 0 1 1 0 1 0 1 0 1 2 0 2 1 1 1 0 0 0 2 2 1 1 010 01 01 1 1 0 1 0 0 0 1 1 1 1 0 1 1 1 1 1 0 0 0 120 2 1 0 0 0 1 1 0 0 0 2 0 Ia. cavaticus Ke. dissimilis 0120001110110011110111110011020111210002 0 Ke. harbachi 0 1 2 0 0 1 0 1 1 0 1 1 0 0 1 1 011 0 1 1 1 1 1 0 0 1 1 0 2 0 1 1 1 2 1 0 0 0 2 0 C Ko. purpureipes 0010010110010000001111100101022011201002 Le. geoskusea 1120001110000110011111110011021000010001 0 Mu. alternans 1 0 1 0 0 1 0 1 1 0 011 0 0 1 0 0 0 0 1 1 1 1 1 0 0 0 1 0 2 2 1 1 0 1 1 0 0 0 2 C Mu. Iciniger 1010000110000111000111110001022120110002 Oc. (Brh.) alekiorovi 2 1 2 010 0 011 1 0 1 0 0 0 1 0 1 1 0 1 0 1 1 1 0 0 011 0 2 1 0 0 0 1 1 0 0 0 2 C Oc. (Brh.) greenii 2121001110100011110101110011021000110002 Or. (Cha.) wattensis 1010010110111110110011010001020000310002 C Oc. (Och.) communis 0 0 2 010 0 1 0 1 0 1 0 0 0 1 1 1 1 0 1 1 1 1 1 0 0 0 1 0 2 1 0 2 1 1 1 0 0 0 2 C Oc. (Och.) infirmatus 00100001100000101 10111110001021001110002 00 Oc. (Och.) scapularis 0011001 110000111010111110001021001110002 . . Oc. (Psk.) bancroftianus 0 0 1 0 0 0 0 1 1 0 0 0 0 0 1 1 ? 1 0 1 1 1 1 10001020001110002 Oc. (Psk.) postspiraculosus 0 0 1 0 0 0 0 1 1 0 0 0 0 0 1 011 1 0 1 1 1 1 1 0 0 0 1 0 2 2 0 0 1 1 1 0 0 0 2 o Oc. (Rus.) refiki 1010001110000011010111110001021101110102 CC C 0011001110001O11010111 110001021101110102 TOO Oc. (Rus.) rusticus •0 Ps. (Gra. ) jamaicensis 1 0 1 0 0 1 0 1 1 0 1 1 1 0 0 0 1 0 011 0 1 1 1 0 0 0 1 1 2 1 1 0 1 1 1 0 0 0 2 C co 0 Ps. (Jan.) ferox 1 1 2 1 0 0 1 1 1 0 1 0 1 0 1 1 1 0 0 1 0 1 1 1 0 0 0 120 2 0 1 1 1 2 1 0 0 0 2 C 0 0 1 0 0 1 1 0 1 0 1 1 0 1 0 1 1 0 1 1 0 1 011 0 0 0 1 1 2 0 1 1 011 1 0 0 0 2 CJD Ps. (Pso.) ciliata Rh. longirostris 1121001011011011100111110002010010010001

CD Rh. wardi 11210001011011011100111110002010010010001 CC) Sc. albolineata 0110010111110010011111110012012001111002 Sc. arhoricola 0 1 1 0 0 0 0 1 1 1 1 1 0 0 1 010 1 1 1 1 0 1 1 0 0 1 2 0 1 2 0 0 1 1 1 1 0 0 2 St. aegypti 0000010110010000001111110001010001211102 0I St. africana 0 0 0 0 0 1 0 0 1 010 1 0 0 1 0 0 0 1 1 1 1 1 1 1 0 0 1 0 1 0 1 1 0 2 1 1 1 0 2

D 0

00 APPENDIX 2 Continued

05 111111111111111111111 8888888889999999999000000000011111111112 1234567890123456789012345678901234567890 cc Za. fulgens 0 0 2 0 0 0 1 1 1 0 1 0 0 0 1 011 1 0 1 0 1 1 1 0 0 1 1 0 2 0 1 1 0 2 1 0 0 0 2 0 Za. longipalpis 00200011101100101101111100010201102 10002 < 0) 0 Do. leonis 1 1 1 0 0 0 0 1 1 0 0 1 0 0 1 0 0 1 1 0 1 1 1 1 0 0 0 120 1 0 0 1 0 2 1 0 0 0 2 50 Do. nivea 1 1 1 0 0 0 0 1 1 0 0 1 0 0 1 0 1 1 1 0 1 1 1 1 0 0 0 120 1 0 0 1 0 2 1 0 0 0 2 or- Fl. kochi 1010000100111010110011110101022000110002 0 Fl. poicilia 1 0 1 0 0 0 0 1 0 0 1 1 1 0 1 011 1 0 0 1 1 0 1 0 0 0 1 0 2 2 0 0 0 1 1 0 0 0 2 Oi) Ha. australis 0110001011001101101111100101021001110002 Hg. (Con.) leucocelaenus 0120010010010000011111110011010011210002 w&) 0 Hg. (Con.) leucotaeniatus 1120010010010010011111110011010011210002 0 1-1g. (Hag.) equinus 1010010010110000001111110001010011210002 0 Hg (Hag.) splendens 1010010010010000001111110001010011210002 Oc. (Fin.) aihilabris 2110010110000110110011110011022000111002 0 0 Oc. (Fin.) alboannulatus ?????????????????????????????21000110002 0 Oc. (Fin.) albocinctus 1020000110100110110111110002010000110002 0 Oc. (Fin.) albolaeniatus 0020010110010010010110110012022000110002 N Oc. (Fin.) argenteitarsis 0110010110010010110011110001022001110002 0 Oc. (Fin.) argyronotum 0 1 2 010 1 011 1 0 011 0 0 1 0 0 1 0 1 1 0 1 1 0 0 0 1 0 2 1 0 0 0 1 1 0 0 0 2 Oc. (Fin.) assamensis 01100011 10000010010011110001020010210002 Oc. (Fin.) hanksi 0 0 1 0 0 1 1 0 1 0 0 1 0 0 0 01011 1 1 1 1 1 1 0 0 1 1 0 2 2 0 0 1 1 1 0 0 0 2 Oc. (Fin.) biocellatus 0020010110000110010111110001021000110002 0 1 Oc. (Fin.) candidoscutellum 0020000110000010110111110001021000110002 Oc. (Fin.) chrysolineatus 1110010110010000110111110011022000110002 Oc. (Fin.) chungi 0121001100100010110111110001020010210002 Oc. (Fin.) crossi 0010010110000110010111110001022000110002 0 Oc. (Fin.) echinus ?????????????????????????????21001110002 0 Oc. (Fin.) elsiae 0010011010010000010110110001021000110002 0 0 0 Oc. (Fin.) e,nhuensis 0 0 1 0 0 0 0 1 1 0 0 0 0 1 1 0 0 1 0 1 1 1 1 1 0 0 0 1.0 2 2 0 1 0 1 1 0 0 0 2 0 Oc. (Fin.) geniculatus 01100001100001100111110110001022001110002 cI C)0 Oc. (Fin.) gilli 0021001110110011110101110011012000110002 C) Oc. (Fin.) gubernatoris 0120001110010010010111110011020000110002 Oc. (Fin.) harperi 001001001001000010110110001020010210002 Oc. (Fin.) hatorii 0020011010010000010111110001021001110002 0) Oc. (Fin.) ingrami 0010000110000110000111110001020010110002 Oc. (Fin.) j. Japonicus 1010010110010010110110110001021000110002 Oc. (Fin.) jugaraensis 2010010010010000110111110011021000110002 Oc. (Fin.) keefei 0120001110100010110111110001021000110002 Oc. (Fin.) br 1010000110010010010001110001021000110002 Oc. (Fin.) bophoventralis 0 1 1 0 0 0 1 1 1 0 1 1 010 1 0 0 1 0 1 1 1 1 1 0 0 1 1 0 2 0 0 0 0 1 1 0 0 0 2 Oc. (Fin.) luteostriatus 0020000110100010110111110001020010212002 01 co cn APPENDIX 2 Continued 9-4 111111111111111111111 8888888889999999999000000000011111111112 tTl 0 1234567890123456789012345678901234567890 z 0 tTI Oc. (Fin.) macfarlanet 1010010101001000001011111000102 1000110002 0 Oc. (Fin.) notoscriptus 2 1 1 0 0 1 0 1 1 0 0 0 0 011 0 1 1 0 1 1 1 1 1 0 0 0 1 0 2 2 0 0 0 1 1 1 0 0 2 © Oc. (Fin.) oreophilus 0120001110110010010111110011022000110002 Oc. (Fin.) papuensis 0120010110010010010110110001021000110002 C Oc. (Fin.) prominens 0120001110110010010111110001020010210002 Oc. (Fin.) pseudotaeniatus 00100100100100000111111110011022000110002 Oc. (Fin.) pulchrithorax 00200011101 10010110111110001022010112002 Oc. (Fin.) pulchri venter ?????????????????????????????21001110002

0 Oc: (Fin.) rubrithorax ?????????????????????????????21001110002 Oc. (Fin.) seoulensis 212000011000011001011111000102000? 110002 (I) Oc. (Fin.) sherki 1110010110010000110110110011022000110002 0 0 Oc. (Fin.) subalbitarsis 0020010110010010110010110011021001110002 0 Oc. (Fin.) tubbutiensis 00200001101000101101 10110001021000110002 C -, Oc. (Mac.) tremulus 0010010011101100011011010001020000110002 C Oc. (Mol.) pecuniosus 00101010100110010001111000001021000110002 Oc. (Och.) atropalpus 0 0 2 0 0 1 0 0 1 0 0 1 1 0 0 0 1 011 1 1 1 1 1 0 1 0 1 0 2 2 0 0 1 1 1 0 0 0 2 0 Oc. (Och.) deserticola 10100101 10010010001111000001022001 110002 Oc. (Och.) epactius 0020010010011000101111110111022001110002 0 Oc. (Och.) fluviatilis 1010010010010000010111110001022001110002 Oc. (Och.) muelleri 1020000110001010011111000001021011110002 0 a Oc. (Och.) sierrensis 101001001000001000111 110000102 12001110002 Oc. (Och.) varipalpus 1 0 1 0 0 1 0 0 1 0 0 0 0 0 1 0 0 0 1 1 1 1 1 0 0 0 0 1 0 2 120 0 1 1 1 0 0 0 2 C Oc. (Pro.) berlini 1 0 2 0 0 1 0 0 1 0 1 1 0 0 1 0 011 0 1 1 1 1 1 0 0 0 1 0 2 2 0 0 0 1 1 0 0 0 2 1 Oc. (Pro.) brelandi 0110010110001010011111110001021001110002 Oc. (Pro.) burgeri 1 0 2 0 0 1 0 1 1 0 0 010 0 1 0 0 1 0 1 1 1 1 1 0 0 0 1 0 2 1 0 0 0 1 1 0 0 0 2 Oc. (Pro.) galindoi 1 0 2 0 0 1 0 0 1 0 1 1 0 0 1 0 011 0 1 1 1 1 1 0 0 0 1 0 2 2 0 0 0 1 1 0 0 0 2 . . Oc. (Pro.) hendersoni 1 0 2 0 0 1 0 1 1 0 0 0 01011 0 0 1 1 1 1 1 1 1 0 0 0 1 0 2 1 0 0 0 1 1 0 0 0 2 o Oc. (Pro.) homoeopus 012001011011001001011111001021000110002 Ca Oc. (Pro.) knabi 0 0 1 0 0 1 0 0 1 0 0 1 0 0 1 0 0 1 0 1 1 1 ? 1 0 0 0 1 0 2 1001110002 a 002001011001001001011111000102 1001110002 a Oc. (Pro.) kompi a Oc. (Pro.) terrens 10200100101100100110111110001021000110002 —o Oc. (Pro.) triseriatus 1 1 2 0 0 1 0 1 1 0 0 1 0 0 1 0 011 1 1 1 1 1 1 0 0 0 1 0 2 1 0 0 0 1 1 0 0 0 2 Oc. (Pro.) zavortinki 1 0 2 0 0 1 0 1 1 0 1 1 0 0 1 0 0 1 0 1 1 1 1 1 0 0 011 0 2 1 0 0 0 1 1 0 0 0 2 C Oc. (Pro.) zoosophus 0110010110000010011? 11110001021000110002 .. Op. (Not.) chat hamicus 2011011010000011111111100002021001110010 -I Op. (Opi.) fuscus 0010011010111001111010100001000012210010 Ta. savoryi 0 1 1 010 1 1 0 1 0 0 1 1 0 0 0 1 1 1 1 1 1 1 1 0 0 0 1 012 1 0 0 1 1 1 0 0 0 2 Ta. togoi 1 0 1 0 0 1 1 0 1 0 0 1 1 0 0 011 1 0 1 1 1 1 1 0 0 0 1 0 2 1 0 0 1 1 1 0 0 0 2 01

U) APPENDIX 2 Continued

050 1111111111111111111111111111111111111111 2222222223333333333444444444455555555556 qg. 12345678901234 5678901234 5678901234567890

wt. Cs. (Cus.) inornata 10201111110000120112100101110101121011111 Cx. (Cux.) quinquefasciatus 1 0 2 1 1 1 1 1 1 1 0 0 0 0 0 1 0 1 1 2 1 0 0 0 0 1 1 0 00 0 0 1 1 0 1 0 0 0 1 CD 0 0) 0 Ma. (Man.) titillans 1021111111000002011210000110100021010001 00 Or signifera 10201111110001111 11110000110010011010101 50 Ab. papago 10201111110001120112211101 101001111110111 Ae. cinereus 01000111110000000101000101 10110011010001 <0 Ae. esoensis 0100011111000000011110010110110011010001 1 zi Az. ramirezi 0020011111011012110012000110111111010111 WCJ) 0 Ge. baisasi 0010011111000000010102100010100020010001 0 0 Ge. longiforceps 01000111110000000101000000101000120010001 Gy. mediovittata 1020110011011112011111010110100101010001 0 Hw. sexlineata 1020010011000112011212010110110111010011 0 Hw. walkeri 1 0 2 0 0 1 0 0 1 1 0 0 0 1 1 1 1 1 0 2 1 120 0 0 1 0 0 1 0 0 0 1 1 0 1 0 0 1 1 0 Ia. cavaticus 0 0 2 0 0 1 1 1 1 1 1 0 0 0 0 1 0 1 1 2 1 0 0 0 0 1 1 0 1 0 0 0 011 0 1 0 0 0 1 Ke. dissimilis 00200100000010000010101 110110100001010001 0N Ke. harbachi 0020010000011000011211110110100001010001 0 Ko. purpureipes 0 0 2 0 0 1 0 0 1 1 0 1 1 0 1 1 0 1 1 2 1 1 0 0 0 1 1 0 0 1 0 1 011 1 1 0 0 1 1

C) Le. geoskusea 0100011111000000010100000010100010010000 0 Mu. alternans 1021111111000002111210010110111111001111 Mu. laniger 1 0 2 1 1 1 1 1 1 1 0 0 0 0 0 2 1 1 1 2 1 0 0 1 0 1 1 0 1 1 1 1 1 1 0 011 1 1 1 a Oc. (Brh.) alektorovi 0 0 2 0 0 1 1 1 1 1 01011 011 2 0 1 1 2 1 0 0 0 0 1 1 0 1 1 0 1 1 1 0 1 0 0 1 1 01 0 Oc. (Brh.) greenii 0010011011101012011210000110100111010011 Oc. (Cha.) wattensis 0 0 2 1 0 1 1 1 1 1 1 0 0 0 0 1 0 1 0 1 0 1 1 0 0 1 1 0 1 0 1 1 1 1 0 1 1 011 1 Oc. (Och.) communis 1 0 2 1 0 1 1 1 011 0 1 1 011 121 1 1 2 1 0 0 1 0 1 1 0 1 1 0 1 121 1 1 011 1 1 0 Oc. (Och.) infirmatus 0 0 2 1 0 1 0 0 0 1 1 0 1 0 011 0 1 1 1 0 0 0 0 0 1 1 0 1 1 0 1 2 1 1 1 1 1 1 1 0 Oc. (Och.) scapularis 0 0 2 1 0 1 0 0 0 1 1 0 1 0 1 1 0 1 1 1 0 0 0 0 0 1 1 0 1 0 0 1 2 1 1 1 011 1 1 0 Oc. (Psk.) bancroftianus 0021011111000012011110000110110021011101 0 0 Oc. (Psk.) postspiraculosus 0021011111000010011210000110110121011101 cI 0 Oc. (Rus.) re/Iki 1 0 2 1 0 1 1 1 1 1 0 1 1 0 1 1 1 1 1 2 1 0 0 1 1 1 1 0 1 1 1 1 2 1 1 1 011 1 1 C) 0 Oc. (Rus.) rusticus 1021111111011011111210011110111121111111 Ps. (Gra.) jamaicensis 1 0 2 1 1 1 1 1 1 1 0 0 0 0 0 2 1 1 1 2 1 0 0 0 0 1 1 1 1 1 0 0 121 0 1 1 1 1 1 tS Ps. (Jan.) ferox 0021011111000002011211100111111111110111 0) Ps. (Pso.) ciliata 0 0 2 1 0 1 1 1 1 1 0 1 1 1 0 1 1 1 1 2 1 2 1 0 0 0 0 1 1 1 1 1 2 1 0 1 011 1 1 Rh. longirostris 01000111110000001101011000001000010010000 Rh. wardi 0100011111000000110101100000100000010000 Sc. aiholineata 001001001 101 1000010101100110100001 100001 Sc. arboricola 00100100110110000101011001 10100001100001 St. aegypti 1 0 2 0 0 1 0 0 1 1 0 1 1 0 1 2 0 1 1 1 2 1 1 1 0 1 1 0 1 0 0 1 011 1 1 0 1 0 1 St. africana 1 0 2 0 0 1 0 0 1 1 0 1 010 0 1 0 1 0 1 2 1 1 1 0 1 1 0 1 0 0 1 0 1 1 1 0 0 0 1 01 C)1 CJI APPENDIX 2 Continued

0 1111111111111111111111111111111111111111 2222222223333333333444444444455555555556 tTl 1234 5678901234 56789012345678901234567890 -I 0 B z 0 Za. fulgens 0020010000011000111211110110100001110111 0 Za. longipalpis 002001000001100001 12111101 10100001110111 Do. leonis 002 1010000000001010201 1001 10100011010111 Do. nivea 0021010000100001010201100110100011010111 Fl. kochi 1021111111000012111211110110100101010111 Fl. poicilia 1021111111000012111212110110100101010111 Ha. australis 1021111111000002011110010110110121011011 Hg. (Con.) leucocelaenus 0020010000011000011211110110110101110111 Hg (Con.) leucotaeniatus 0 0 2 0 0 1 0 0 0 0 0 1 010 0 0 0 1 1 2 1 1 1 1 0 1 1 0 1 1 0 1 0 1 1 1 0 1 1 1 0 Hg. (Hag.) equinus 0010000000000000100201110110010101100111 0 Hg. (Hag.) splendens 0100000000000000100201110110010101100111 cr 0 Oc. (Fin.) albilabris 1 0 2 1 1 1 1 011 1 0 1 1 0 1 2 0 1 1 1 1 1 1 1 0 1 1 0 1 0 0 1 1 1 0 1 0 0 1 1 Oc. (Fin.) alboannulatus 1021111111000012011110010110110111010111 Oc. (Fin.) alhocinctus 0021010001101001011210000110100111010011 0 Oc. (Fin.) alhotaeniatus 0021110011101001011110000110110111010111 0 0 Oc. (Fin.) argenteitarsis 0021010000100000010000010110100101010001 0 Oc. (Fin.) argyronotum 1021010001100001010100010110110111010011 0 Oc. (Fin.) assamensis 0 0 2 0 0 1 0 0 0 1 1 0 0 0 0 1 0 1 1 1 2 1 1 1 0 1 1 0 1 0 0 0 011 0 1 0 0 1 1 Oc. (Fin.) hanksi 1Q21011 111011112011012010110110111010111 C Oc. (Fin.) hiocellatus 1021111101101011011212010110100111010111 Oc. (Fin.) candidoscutellum 0021111111011112011211100000100011010001 C Oc. (Fin.) chrysolineatus 1 0 2 0 1 1 1 0 1 1 0 1 1 1 1 2 0 1 1 2 1 2 0 0 0 1 1 0 1 1 0 0 011 0 1 0 0 1 1 Oc. (Fin.) chungi 0020010011011112011211210110100011010111 Oc. (Fin.) crossi 0021011111011112011210010110110111010011 Zn Oc. (Fin.) echinus 0021011111011111011212010110110111010111 Oc. (Fin.) elsiae 1021111111011112011211010110110111010111 Oc. (Fin.) embuensis 0 02 101011110 10 110 112 10 010 110 10 0 111010 111 Or. (Fin.) geniculatus 0021011110111110121001011011011 1010111 o Oc. (Fin.) gilli 0 0 2 1 0 1 0 0 0 1 1 0 1 0 1 1 0 1 1 2 1 0 0 0 0 1 1 0 1 0 0 0 1 10 1 0 1 1 1 on Oc. (Fin.) gubernatoris 0021010001100001011121210110100011010111 0•0 Oc. (Fin.) harperi 0 0 2 1 1 1 0 0 1 1 0 0 0 0 0 0 0 1 0 1 2 1 0 0 0 1 1 0 1 0 0 0 011 0 0 0 1 1 1 0 Oc. (Fin.) hatorii 1021011111011112011210010110110111010111 C1) Oc. (Fin.) ingrami 0021010011011.011010211110110100111010111 Oc. (Fin.) j. japonicus 1020011111011112011210000110110011010011 Oc. (Fin.) .jugaraensis 1020011011011110110112000110110011010011 1 0 0 Oc. (Fin.) keefei 010 2 1 011 1 1 1 1 0 1 1 1 1 2 0 1 1 0 1 2 0 1 0 1 1 0 1 0 0 0 2 1 0 1 0 0 1 1 Oc. (Fin.) loi 0021011111000012011110000110110111010011 Oc. (Fin.) lophoventralis 0021010001100001010121110110100011010001 Oc. (Fin.) luteostriatus 0020010011011010011210010110100001010011 0I- -. APPENDIX 2 Continued

. 0 1111111111111111111111111111111111111111 o . c 2222222223333333333444444444455555555556 123456789012345678901234 5678901234567890

Ci) Oc. (Fin.) macfarlanei 1021111111011112011210010110110111010111 o Oc. (Fin.) notoscriptus 10211111110111120111111101101001 11010011 < o. 0 •1 Oc. (Fin.) oreophilus 0020011111011 1120112100101101001 11010011 Oc. (Fin.) papuensis 0021010001100001011100010110110111010111 Oc. (Fin.) prominens 0020010001 100001010121110110100011010011 . Oc. (Fin.) pseudotaeniatus 1021011111011112011012010110110011010111 0) Oc. (Fin.) pulchrithorax 0020010011011110011210010110100001010111 cn Oc. (Fin.) pulchriuenter 1021011111101012011112010110111111010111 0 Oc. (Fin.) rubrithorax 1021011111000012011210010110110111011111 Ct Oc. (Fin.) seoulensis 0020010001100001011210010110100111010011 0 Oc. (Fin.) sherki 1020011111011110110012100110110011010011 Oc. (Fin.) subalbitarsis 0021010000100000010000010110110011010011 0 0 Oc. (Fin.) tubbutiensis 1021111111000002011210010110110111010111 0 1020011111000012011212010110110111010011 0 Oc. (Mac.) Ireinulus Oc. (Mol.) pecuniosus 0021011111011011011211110110100111010111

C Oc. (Och.) atropalpus 0010010001000011011210000110110111010111 Oc. (Och.) deserlicola 102 00 111110000 12 0112 112 10110110111010011 0 0010010001000011011210000110110111010111 a Oc. (Och.) epactius Oc. (Och.) fluviatilis 0 0 1 1 0 1 010 0 1 0 0 0 0 1 121 1 011 1 0 0 1 0 1 1 0 1 0 0 1 1 1 0 1 0 1 1 1 Oc. (Och.) muelleri 0021011111011011011210010110111121110111

a Oc. (Och.) sierrensis 1020011111001012011212110110110111010111 a Oc. (Och.) varipalpus 1 0 2 0 1 1 1 1 1 1 0 0 0 0 1 2 0 1 1 2 1 02 12 1 0 1 1 0 1 1 0 1 1 1 0 1 0 1 1 1 Oc. (Pro.) berlini 002101111000001010210010110100011010011 0 Oc. (Pro.) brelandi 0021011111011011011212010110110111010011 Oc. (Pro.) burgeri 0021011111011011011210010110100111110011 a Oc. (Pro.) galindoi 00200111110000010101100101 10100011010011 0a a Oc. (Pro.) hendersoni 00210111100000010112120101 10100111010011 a Ci) Oc. (Pro.) homoeopus 0021011111000001010210010110100111010011 C 0 Oc. (Pro.) knahi 0 0 2 1 0 1 1 0 1 1 0 1 1 0 1 1 0 1 1 2 1 2 0 1 0 1 1 0 1 0 0 011 1 0 1 0 0 1 1 0 Oc. (Pro.) kompi 0021011101011001011210010110110111010011 0021011101000001011210010110110111110011 ND Oc. (Pro.) terrens CD 0 Oc. (Pro.) triseriatus 0 0 2 1 0 1 0 0 1 1 0 0 0 0 0 1 0 1 1 2 1 120 1 0 1 1 0 1 010 1 1 1 0 1 0 0 1 1 0) Oc. (Pro.) zavortinki 0021011111001011011211010110100111010011 Oc. (Pro.) zoosophus 0021010011001011011212010110100111010011 Op. (Not.) chathamicus 0100011111000002010110000110110011011011 Op. (Opi.) fuscus 1110111111000002011211100110110011010101 0 Th. savoryt 0 0 1 0 0 1 1 1 1 1 0 1 1 1 1 2 0 1 1 2 1 0 0 1 0 1 1 0 1 1 0 1 011 0 1 0 0 1 1 Ta. togoi 1 0 1 0 0 1 1 1 1 1 0 1 1 1 1 2 0 1 1 2 1 0 0 1 0 1 1 0 1 1 0 1 1 1 0 1 010 1 1 01 APPENDIX 2 Continued

C 9- 1111111111111111111111111111111111111112 Ti 6666666667777777777888888888899999999990 1234567890123456789012345678901234567890 C Cs. (Gus.) inornata 101101131000001110000022220001101 01 01001 Cx. (Cux.) quinquefasciatus -3 C 100101001000000100000121210001101 01 ? 1001 Ma. (Man.) titillans 100100031000001110010012120002211 00 ?0000 © Or. signifera 000101031000001110111012120001101 01 01001 0 Ab. papago 0001000110100011010100111 1 0 0 1 1 ? 0 1 01 01101 Ae. cinereus 000101001000000000000211110112101 11 01101 Ae. esoensis 000101001000001000000211110112101 11 01101 Az. ramirezi 0001101001000001000000222220011101 01 11102 Ge. baisasi 0 12 10100 1000000000000 0222 10000 111 10 10101 Ge. longiforceps 012101001000000000000022210000111 10 00101 Gy. mediovittata 0 0 0 2 1 0 0 011 0 1 0 0 1 1 1 1 0 0 1 0 2 2 1 2 1 0 0 1 1 1 0 1 01 01101 0) C Hw. sexlineata 000100001000001010010011110010001 01 01101 C, CD Hw. walkeri 000100001000001010010022220010001 01 0 1 1 0 01 Ia. cavaticus 100101001000001000000221210010101 01 01101 C -, Ke. dissimilis 000100001000001000000221210010001 01 1 00111 0 Ke. harhachi 000100001000001000000221210010001 01 1 00111 CL Ko. purpureipes 0 000101001000001000000221210011101 11 01101 0 Le. geoskusea 00010000100000000000001100001 1111 10 10101 N 0 Mu. alternans 10010102 1000101111010222221112111 10 00101 0 Mu. laniger 1 0 0 1 0 1 0 1 1 0 0 0 1 0 1 1 1 0 0 010 2 2 2 2 2 1 1 1 2 1 1 1 10 00101

C, Oc. (Brh.) alektorovi 0 0 0 1 0 1 0 0 1 0 0 0 0 0 1 1 1 0 1 1 1 2 2 2 2 2 0 0 1 0 0 0 1 11 01101 Oc. (Brh.) greenii 0 0 0 1 0 1 0 011 0 0 0 0 0 1 1 1 0 1 1 1 2 2 2 2 2 0 0 1 0 0 0 1 11 01101 Oc. (Cha.) wattensis 000102001001001010010011110011001 01 11101 C Oc. (Och.) communis 101101011000001 100000221211112111 10 01101 Oc. (Och.) in/i rmatus 00110100100000010000022222 1102111 10 00101 Oc. (Och.) scapularis 000101011000000110000222221 102111 10 00101 Oc. (Ask.) hancroftianus 0 0 0 1 0 1 0 011 0 0 0 0 0 0 0 0 0 0 0 0 2 2 2 2 1 0 0 1 2 1 1 1 10 10101 Oc. (Psk.) postspiraculosus 000101011000000000000022210012111 10 10101 Oc. (Rus.) refiki 101101011110001110000222221112111 11 11101 Oc. (Rus.) rusticus 1011010111100011 10000222221112111 11 01101 Ps. (Gra. ) jamaicensis 100000031000101111010022220012210 00 00102 Ps. (Jan.) ferox 1 0 011 0 0 0 0 1 1 1 0 0 0 1 0 0 0 0 0 0 2 2 2 2 2 1 1 1 2 2 1 0 10 00102 Ps. (Pso.) ciliata 100101031010000110010222221102210 00 00102 Rh. longirostris 0 0 0 1 0 010 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 1 1 1 1 0 0 1 0 1 011 10 00101 Rh. wardi CD 0001010000000000000000111100101011 10 00101 0 Sc. albolineata 000101001001001010010011000010001 01 01101 I- Sc. arhoricola 000101001001001010010011100010001 01 01101 St. aegypti 000100011000011010010211000010001 11 01101 St. africana 0I 0001000010000001 10010210101111101 01 01101 - APPENDIX 2 Continued

0 1111111111111111111111111111111111111112 o 6 6 6 6 6 6 6 6 6 7 7 7 7 7 7 7 7 7 7 8 8 8 8 8 8 8 8 8 8 9 9 9 9 9 9 9 9 9 9 0 123456789012345678901234 5678901234567890

Za. fulgens 0001010010000000000102212111 101010101101 o Za. longipalpis 00010000100000011001022 12111101010101101 < C) Do. leonis 0001010010000000000002222200110010101101 •1 0 Do. nivea 0001010010000000000002222200110010101 101 Fl. kochi 0002000210001011111012222200110010101101 Fl. poicilia 0 00020002100010 11111012222200110010101101 Ha. australis 1001010010000011100002212211121011101101 -0 Hg. (Con.) leucocelaenus 0001000010000010000002212100100010101100 ( Cl) C Hg. (Con.) leucotaeniatus 0 0 0 1 0 0 0 0 1 0 0 0 0 0 1 0 0 0 0 0 0 2 2 1 120 0 0 1 0 0 0 1 0 1 0 1 1 0 0 Hg. (Hag.) equinus 0 0 0 1 0 0 0 0 1 0 1 1 0 0 1 0 0 0 0 0 0 2 2 1 2 0 0 0 1 1 0 0 1 0 1 1 1 1 0 01 Hg. (Hag.) splendens 0 0000000010110010000000111100100010111100 - Oc. (Fin.) albilabris 0001000010000111100102222100110010000101 0 0 Oc. (Fin.) aihoannulatus 0001010010000011100102222111120011101100 0 Oc. (Fin.) albocinctus 0001010110000001100102222200111010101101 0 Oc. (Fin.) albotaeniatus 0 0 0 1 0 1 0 1 1 0 0 0 0 0 1 1 1 0 0 1 0 2 2 2 2 2 0 0 1 1 0 0 1 0 1 0 1 1 0 01 CN Oc. (Fin.) argenteitarsis 0001000010000010100102222200110010101100 C Oc. (Fin.) argyronotum 0001010010000010100102222200121011101100 CR Oc. (Fin.) assamensis 0001000110000000101 102222200110010100101 p Oc. (Fin.) hanksi 0001010110000101101 112222200111010111001 Oc. (Fin.) biocellatus 0001010110000011100102222200120010100101 1 Oc. (Fin.) candidoscutellum 0001010010000011100102222200020010100101 C Oc. (Fin.) chrysolineatus 0 0 0 1 0 1 0 1 1 0 0 0 0 0 0 1 1 0 0 1 0 2 2 1 2 1 0 0 1 1 0 0 1 0 1 0 1 1 0 1 Oc. (Fin.) chungi 0 0 0 1 0 1 0 1 1 0 0 0 0 0 1 0 1 0 0 0 0 2 2 2 2 2 ? 0 1 2 0 0 1 0 1 0 0 1 0 1 Oc. (Fin.) crossi 0 0 0 1 0 1 0 0 1 0 0 0 0 0.0 0 0 0 0 0 0 2 2 2 2 2 0 0 1 2 ? 0 1 0 1 1 1 1 0 1 Oc. (Fin.) echinus 0011010010000010000002222100120011101101 0 Oc. (Fin.) elsiae 0001010110000101101112222200121010101101 0 p Oc. (Fin.) embuensis 0001010110000000100102222200121010101001 C Oc. (Fin.) geniculatus 0001010010000010000002222100120010100101 Cl) C Oc. (Fin.) gilli 0001010010000001100102222200120011101101 Oc. (Fin.) gubernatoris 0001000110000010101112222200110010101101 Oc. (Fin.) harperi 0 0 0 1 0 0 0 1 1 0 0 0 0 0 1 1 1 0 0 1 0 2 2 2 2 2 0 0 1 1 0 0 1 0 1 0 1 1 0 01 Oc. (Fin.) hatorii 0001010110000101011122222012011101101 C C) Oc. (Fin.) ingrami 0 0 0 1 0 1 0 011 0 0 0 0 0 0 0 1 0 0 1 0 2 2 2 2 2 0 0 1 2 0 0 1 0 1 0 1 1 0 1 Oc. (Fin.)j. japonicus 0001010010000001100102222200111011101101 Oc. (Fin.) jugaraensis 0001010010000001100102222200111011101101 Oc. (Fin.) keefei 0001010010000011100102222200120010101101 C Oc. (Fin.) loi 0001010010000010100102??? ??01??0101 10101 Oc. (Fin.) lophoventralis 0001000010000010101102222200110010100101 Oc. (Fin.) luteostriatus 0001010010000001100102212111110010100101 C)1 rc APPENDIX 2 Continued 9- C 1111111111111111111111111111111111111112 TI 6666666667777777777888888888899999999990 C 1234567890123456789012345678901234567890 z C Oc. (Fin.) macfarlanei 0001010110000101101112222200121010100101 0 Oc. (Fin.) notoscriptus 0001000010000111100102222100110010000101 t:T1 © Oc. (Fin.) oreophilus 0001010110000010000002222200110010111101 Oc. (Fin.) papuensis 0001010010000011100102222200120011 101 100 C Oc. (Fin.) prom inens 00010101100000111011122222001100101 00101 Oc. (Fin.) pseudotaeniatus 0001010110000101101112222200111010101001 Oc. (Fin.) pulchrithorax 0001010010000001100102222111110010101101 Oc. (Fin.) pulchri venter 0001010110000010000002222200111011101101 0 Oc. (Fin.) rubrithorax 0001010010000011100102222111121011101100

0 Oc. (Fin.) seoulensis 0001010110000001101112222200110011101101 CD Oc. (Fin.) sherki 0001010110000001100102222200110010100101 C 0 Oc. (Fin.) subalbitarsis 00020100100000101001o2222211120011100100 Oc. (Fin.) tubbutiensis 0 0 0 1 0 1 0 011 0 0 0 0 0 1 1 1 0 0 1 0 2 2 2 2 2 1 1 1 2 0 0 1 0 1 0 1 1 0 0 0 Oc. (Mac.) tremulus 0001000010000001100100211100110010110101

0 Oc. (Mol.) pecuntosus 0 0 011 0 1 0 1 1 0 0 0 0 0 1 0 1 0 0 1 0 2 2 2 2 2 0 0 1 1 1 0 1 0 1 ? 1 1 0 1 0 0- Oc. (Och.) atropalpus 0001000110000011101112222200011010101010 0 0 Oc. (Och.) deserticola 0001000110000011101012212100121111000101 Oc. (Och.) epactius 0001000110000011101112222200011010101010 C Oc. (Och.) fluviatilis 0 0 0 1 0 0 0 1 1 0 1 0 0 0 1 1 1 0 0 1 0 2 2 2 2 2 0 0 1 1 1 0 1 0 1 0 1 0 1 0 Oc. (Och.) muelleri 0 0 1 1 0 1 0 0 1 0 1 0 0 0 1 0 0 0 0 0 0 0 1 1 1 1 0 0 1 2 1 1 1 011 0 1 1 0 1 0 Oc. (Och.) sterrensis 0011000110000011101112212 100111111101101 Oc. (Och.) varipalpus 0 0 1 1 0 1 0 1 1 0 1 0 0 0 1 1 1 0 1 1 1 2 2 1 2 1 0 0 1 1 1 1 1 1 0 0 011 0 1

0 Oc. (Pro.) berlini 0001000110000010101102222200121010101101 1 Oc. (Pro.) brelandi 0001010010000010000002222200120010101101 Oc. (Pro.) burgeri 001101001000001000000222220012 1010101101 Oc. (Pro.) galindot 0001000110000010101102212100110010101101 Oc. (Pro.) hendersoni 0001010110000010000002222200120010101101 o Oc. (Pro.) homoeopus 000101011000001010110222220012001 0101101 •1 c Oc. (Pro.) knabi 0 0 0 1 0 1 0 0 1 0 0 0 0 0 1 0 1 0 0 1 0 2 2 2 2 2 1 1 1 ? 0 0 1 011 0 1 1 0 1 Oc. (Pro.) kompi 0 0 0 1 0 0 0 0 1 0 0 0 0 0 1 0 0 0 0 0 0 2 2 1 2 1 0 0 1 2 0 0 1 0 1 0 011 0 1 0 Oc. (Pro.) terrens 000100011000001010110222220012001 0101101 0. Oc. (Pro.) triseriatus 0001010010000010000002212100120010101101 Oc. (Pro.) zavortinki 000101011000001010110222220012 001010 1101 0 Oc. (Pro.) zoosophus 0001010110000010100102222200120010101101 - Op. (Not.) chathamicus 0 0 0 1 0 1 1 0 1 0 0 0 0 0 010 0 0 0 0 0 2 2 1 2 1 1 1 0 1 0 0 1 1 1 0 1 1 1 1 Op. (Opi.) fuscus 1001011010000011100002000001001 01010100111 Ta. savoryi 00010000100000100000022222111 00011111101 -QQ Ta. togoi 1001000110000011101112222211100011111101 0I

C APPENDIX 2 Continued

09C.0 22222222222222222222222222222222 0 •1. 00000000011111111112222222222333 12345678901234 567890123456789012

CI) Cs. (Gus.) inornata 110 12 0 102 1100010 1010 16 01010 12 0 10 CD 0 Cx. (Cux.) quinquefasciatus 1 1 1 1 2 0 1 1 1 0 0 0 0 0 1 0 1 0 1 0 1 2 0 1 - - - 1 0 1 1 03 Ma. (Man.) titillans 110110110101001001012130112001010 Or signifera 1 1 0 1 1 0 1 1 1 1 010 0 1 1 0 1 0 1 0 1 2 0 1 0 0 0 0 1 0 1 3 Ab. papago 01011110111001001010110112000013 Ae. cinereus 0100011010101100100200- 100010000 Ae. esoensis 0100011010101100100200-100010000 -1j A.z. ramirezi 00011010211001001010110112000013

0 Ge. baisasi 01111010211001021010110100002014 Ge. longiforceps 001011010221001001010110100001014 01011110211001001010111013 000013 0 Gy. mediovittata -C Hw. sexlineata 01011110211001001010110111000013 0 Hw. walkeri 0101101021 1001001010010111000013 0 0 Ia. cavaticus 0100001 1110001001010110100010100 Ke. dissimilis 001011011111001001010111010001013 Ke. harhachi 001011011111001001010111010001013 C Ko. purpureipes 00011010211001011010110112000013 ca Le. geoskusea 01011010111011010010160111001014 0 C Mu. alternans 0 1 0 1 1 0 1 1 1 1 1 0 1 1 0 1 1 0 1 0 1 1 0 1 1 3 0 0 0 0 1 01 Mu. laniger 01011011111011011010110112000010 Oc. (Brh.) alektoroui 0101101021101 1011010110123001013 Oc. (Brh.) greenii 01011010211011011010110123001013 Oc. (Cha.) wattensis 01011010111011011010011113000013 Oc. (Och.) communis 00011110211111011010110113001010 Oc. (Och.) infirmatus 00011010211111011010110113002010 > Oc. (Och.) scapularis 00011110211111011010110113001010 C Oc. (Psk.) hancroflianus 0 1 0 1 1 0 1 0 2 1 1 0 0 1 0 0 1 0 1 0 1 1 0 1 0 0 0 0 2 0 1 0 TI C Oc. (Psk.) postspiraculosus 01011010211001001010110100002010 C)) ci 0C Oc. (Rus.) refiki 00011110201111011010110113102010 Oc. (Rus.) rusticus 00011110201111011010110113102010 Ps. (Gra. ) ja,naicensis 00001011111001101013130100002010 0 z Ps. (Jan.) ferox 00001011111001101013130100002010 0 C) Ps. (Pso.) ciliata 00001011111000101013130100002010 Rh. longirostris 00011001021111101011120120000014 Rh. wardi 0 0 0 1 1 0 0 1 012 1 1 1 1 1 0 1 0 1 1 1 2 0 1 2 0 0 0 0 0 1 4 Sc. albolineata 01000011111001001010040100010003 Sc. arboricola 01000011111001001010040100010003 St. aegypti 01000111100001001010110100012103 St. africana 01000111110001001000110100012103 0) APPENDIX 2 Continued

C. 9- 0 22222222222222222222222222222222 Ij 00000000011111111112222222222333 T1 12345678901234 5678901234 56789012 -4 0 z Tj Za. fulgens 010110101211011001010110112002013 Za. longipalpis 0 1 0 1 1 0 1 0 2 1 1 0 1 1 0 0 1 0 1 0 0 1 010 1 2 0 0 1 0 1 3 0 tj Do. leonis 0101100011101 1001110111012001013 Do. nivea 01011000111011001110111012001013 CD Fl. kochi 0101101011100101 1110111113100013 O) Fl. poicilia 01011010111001001110111013100013 Ha. ausiralis 0 1 0 1 1 0 1 1 121 1 0 0 1 0 1 1 0 1 0 1 1 1 1 0 0 0 0 0 0 1 1 Hg. (Con.) leucocelaenus 0000201101 1001001110111113000013 Hg. (Con.) leucotaeniatus 00002011011001001110110113000013 Hg. (Hag.) equinus 00002010111011011110111113000013 Hg. (Hag.) splendens 0000201101101 10111101101 13000013 CD 0 r) Oc. (Fin.) albilabris 01011010111001001010111012001013 Oc. (Fin.) alboannulatus 0 1 0 1 1 0 1 0 2 1 1 0 1 1 0 0 1 0 1 0 1 1 0 1 1 2 0 0 0 0 1 02

0 Oc. (Fin.) albocinctus 01011110101011001010110112002013 Oc. (Fin.) albotaeniatus 0101101 1111011011110111113002013 0 Or. (Fin.) argenteitarsis 01011011011011011010111112100013 0 Oc. (Fin.) argyronoturn 0 1 0 1 1 0 1 1 0 1 1 0 1 1 0 1 1 0 1 0 1 1 0 1 1 2 1 0 1 0 1 23 Oc. (Fin.) assamensis 01011010211001001010111012002013 Oc. (Fin.) banksi 01011111101001001010110112002012 0 Oc. (Fin.) hiocellatus 01011010211011001110110013002013 ci Oc. (Fin.) candidoscutellum 0 1 0 1 1 0 1 0 2 1 1 0 1 1 0 1 1 0 1 0 0 1 0 0 1 3 0 0 1 0 1 23 Oc. (Fin.) chrysolineatus 01011011111001001010110112002013 Oc. (Fin.) chungi 01011010111001001010011013101013 01 Oc. (Fin.) crossi 01011111111011001010110112002013 Z?. Oc. (Fin.) echinus 01011010211001001010110112001013 Oc. (Fin.) elsiae 0 1 0 1 1 0 1 1 1 1 1 0 0 1 0 0 1 0 1 0 1 5 0 1 1 2 0 0 2 0 1 23 Oc. (Fin.) embuensis 01011010211011001010110112001013 Oc. (Fin.) geniculatus 01011010211001001010110112001013 o 0 Oc. (Fin.) gilli 01011010211001001010010112001013 Or. (Fin.) gubernatorts 01011010211001001010111013102013 Or. (Fin.) harpert 01011010111011001110111113002013 Oc. (Fin.) hatorii 01011011111001001011110112000012 Oc. (Fin.) ingrami 01011110211011001010110112001013 Oc. (Fin.) j. japonicus 01011011111001001010110112002013 Or. (Fin.) jugaraensls 01011011111001001010110112002013 Oc. (Fin.) keefei 01011010211011011010110013002013 Or. (Fin.) loi 0 1 0 1 1 0 1 0 2 1 1 0 1 1 0 0 ? 0 1 0 1 1 0 1 1 3 0 0 ? 0 1 3 Oc. (Fin.) lophoventralts 01011010211001001010111012002013 Oc. (Fin.) luteostriatus 00011010211011011010110013002013 11PV 1.

z APPENDIX 2 Continued

22222222222222222222222222222222 o . = 00000000011111111112222222222333 12345678901234567890123456789012

Oc. (Fin.) macfarlanei 0 1 0 1 1 0 1 1 1 1 1 0 0 1 0 0 1 0 1 0 1 5 0 1 1 2 0 0 2 0 1 23 o Oc. (Fin.) notoscriptus 01011010111001001010111012001013 Oc. (Fin.) oreophilus 01011 010211001001010111012102013 - _j Oc. (Fin.) papuensis 0 1 0 1 1 0 1 1 0 1 1 0 1 1 0 1 1 0 1 0 1 1 0 1 1 2 1 0 1 0 1 23 Oc. (Fin.) prominens 0101101021100100101011 1012002013 Oc. (Fin.) pseudotaeniatus 0 1 0 1 1 1 1 1 1 0 1 0 0 1 0 0 1 0 1 0 1 1 0 1 1 2 0 0 2 0 1 23 Oc. (Fin.) pulchrithorax 00011010211011011010110013002013 rJw Oc. (Fin.) pulchri venter 0 1 0 1 1 1 1 1 1 1 1 0 0 1 0 0 1 1 1 0 1 1 0 1 1 2 0 0 1 0 1 23 0 Oc. (Fin.) rubrithorax 0 1 0 1 1 0 1 0 1 1 1 0 1 1 0 0 1 0 1 0 1 1 1 1 1 3 0 0 0 0 1 23 Oc. (Fin.) seoulensis 01011110211011011010110112001013 0 Oc. (Fin.) sherki 01011011111001001010110112002013 — Oc. (Fin.) subalbitarsis 0 0 0 1 1 0 1 1 0 1 1 0 1 1 0 1 1 0 1 0 1 1 1 1 1 2 1 0 0 0 1 23 0 Oc. (Fin.) tubbutiensis 01011010111011011010111112000012 0- 0 Oc. (Mac.) tremulus 01011010111001000010011012002013 0 Oc. (Mol.) pecuniosus 01011010111001001010110013000013 Oc. (Och.) atropalpus 01011011011011011010110112000013 0 Oc. (Och.) deserticola 00011010211011011010110113000013 Oc. (Och.) epactius 01011011011011011010110112000013 a Oc. (Ochj fluviatilis 0 1 0 1 1 0 1 1 0 1 1 0 1 1 0 1 1 0 1 0 1 1 0 1 1 2 0 0 0 0 1 23 Oc. (Och.) muelleri 0 0 0 1 1 011 0 2 1 1 0 1 1 0 1 1 0 1 0 1 1 0 1 1 2 0 0 1 0 1 3 0 Oc. (Och.) sierrensis 00011010221011011010110112001013 a Oc. (Och.) varipalpus 0001101021101011010110113002013 Oc. (Pro.) berlini 01011010211001001010111012002013 0- Oc. (Pro.) brelandi 01011010211001001010111012002013 Oc. (Pro.) burgeri 0 1 0 1 1 0 1 0 121 1 0 0 1 0 0 1 0 1 0 1 1 1 1 1 2 0 0 2 0 1 3 a Oc. (Pro.) galindoi 01011010211001001010111012002013 a Oc. (Pro.) hendersoni 01011010211001001010111012002013 a Oc. (Pro.) homoeopus 01011010211001001010111012002013 0C/) Oc. (Pro.) knabi 010110102? 1 0 0 1 0 0 1 0 1 0 1 ? ? 0 1 2 0 0 2 0 1 3 Oc. (Pro.) kompi 01011010211001001010111012002013 Oc. (Pro.) terrens 01011010211001001010111012002013 Oc. (Pro.) triseriatus 01011010211001001010111012002013 Oc. (Pro.) zavortinki 01011010211001001010111012002013 I- Oc. (Pro.) zoosophus 01011010211001001010111112002013 Op. (Not.) chathamicus 01011011111001001010130100000011 Op. (Opi.) fuscus 21011011110000000002130000002011 Ta. savoryi 00011011111001001110110112002011 Ta. togoi 00011011111001001110110112001011 64 J. F. REINERT ET AL.

APPENDIX 3 mesokatepisternum with broad, pale scales in upper and lower patches, several upper and posterior setae; NEW GENERIC-LEVEL TAXA prealar area with broad, pale scales on upper (absent The new generic-level taxa recognized in the present in Co. banksi) and lower areas, numerous upper setae; study are described below. Additional character data mesepimeron with 1 patch of broad, pale scales on for those species of each genus included in the analysis upper area and extending onto middle, several upper are contained in the data matrix (Appendix 2). The setae, lower setae absent. genera are listed in alphabetical order. Endings of some masculine adjectival species names have been Wing: Dark-scaled with elongate, narrow, pale-scaled changed to agree in gender with new feminine generic patch at or near base of anterior margin of costa; names. 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. COLLESSIUS REINERT, HARBACH & KITCHING, Legs: Midfemur with narrow, longitudinal stripe of GEN. NOV. white-scales from base to or near apex on middle of anterior surface; hindfemur with subapical white Type species: Ochierotatus macfarlanei Edwards, 1914. 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, Females simple. Head: Vertex with decumbent scales narrow, curved, Abdomen: Tergum I with pale-scaled patch on lateral pale and dark; erect forked scales numerous on occiput tergite; terga II—V with narrow, pale-scaled, basal and vertex; ocular line narrow, with narrow, curved, bands. pale scales; normally 4 or 5 interocular setae (occasionally 6 in Co. elsiae elsiae); eyes contiguous or sep- Genitalia: Tergum VIII with apex broadly rounded, arated by less than width of 1 eye facet; antennal numerous broad scales on approximately distal pedicel with small, broad, pale scales on mesal sur- 0.30-0.40; sternum VIII sclerotized, width greater face; clypeus bare; maxillary palpus dark-scaled with than length, apical margin with moderate, median apex pale-scaled; proboscis longer than forefemur, emargination separating broadly rounded lobes (Co. dark-scaled with patch of pale scales on ventral and hatorii and Assemblage B with apex nearly straight), lateral surfaces near midlength (Assemblage A) or numerous short, lanceolate scales and occasionally 1- entirely dark-scaled (Assemblage B). 3 broad scales (Co. hatorii and Assemblage B without lanceolate scales, but with several to numerous broad Thorax: Scutum covered with narrow, curved scales scales); seta 2-S inserted posterior to 1-S; tergum IX except for bare median prescutellar area, scales comprising single, moderately to heavily pigmented, sparse on scutal fossa, scales dark except for narrow, elongate plate with median emargination apically in pale-scaled stripes on acrostichal (anterior and poste- Assemblage A or with plate short and wide in Assem- rior) area, dorsocentral (anterior and posterior) area, blage B, several setae on each side of midline distally; outer and posterior margins of scutal fossa, outer mar- postgenital lobe relatively long, narrow, apex nor- gins of prescutellar area mesad of setae, and longitu- mally flat or with very shallow, median emargination dinal stripe on supraalar area; setae well developed on (Co. hatorii and Assemblage B with postgenital lobe anterior promontory, acrostichal (anterior and poste- somewhat shorter and wider), several setae distally; rior) area, antedorsocentral area, dorsocentral (ante- upper vaginal sclerite small to moderate size; lower rior and posterior) area, scutal fossa, antealar and vaginal sclerite absent; insula liplike, with several supraalar areas, and lateral margins of prescutellar setae in lateral patches; cercus moderately long, area; scutellum with narrow or broad scales on mid- moderately wide throughout length, apex broadly lobe, narrow scales on lateral lobes; antepronota rounded, scales absent (Assemblage B with cercus widely separated, with broad, pale scales, several somewhat shorter and with scales); 3 spermathecal setae; postpronotum with scales, few posterior setae; capsules. prespiracular setae absent; hypostigmal area bare; postspiracular area with scales, several setae; subspi- (Co. banhsi racular area with pale scales without or Males with few scales, Co. pseudotaeniatus without scales); upper proepisternum with pale scales, several setae, Head: Maxillary palpus with 5 palpomeres, slightly lower area bare; paratergite narrow, with pale scales; shorter than proboscis, palpomeres 4 and 5 slightly

Journal compilation © 2006 The Linnean Society of London, Zoological Journal of the Linnean Society, 2006, 148, 1-101 No claim to original US government works. PHYLOGENY AND CLASSIFICATION OFFINLAYA 65 down-turned, palpomeres 3 (distal part) and 4 with Fourth-i nstar larvae moderate number of long setae ventrally; antennal Head: Seta 1-C slender, single, apex attenuate; 4-C flagellar whorls with numerous long setae directed short, slender, with few branches, inserted at approx- dorsally and ventrally. imately same level or slightly anterior to 5,6-C; 5,6-C Legs: Fore- and midtarsi with 2 unequal ungues, relatively short, moderately stout, with several larger unguis with 2 teeth, smaller unguis with 1 branches, 6-C lateral to 5-C, 4-6-C all at approxi- tooth; hindtarsus with 2 equal ungues, simple. mately same level; 7-C moderately long, relatively slender, with several branches, inserted lateral and Genitalia: Tergum LX with poorly sclerotized, moder- posterior to 4-6-C; 12-C short, slender, with several ate lobe bearing several moderately long, slender branches, inserted mesal to 13-C; 13-C single, longer setae on each side of midline of posterior margin; than 12-C; 14-C short, stout, normally single or gonocoxite moderately long, relatively narrow 2-branched, occasionally 3-branched; 19-C absent; throughout length, dorsal surface without apical antenna moderately long, narrow, with several to and basal lobes on mesal margin, with few moder- numerous spicules, seta 1-A moderately long, nor- ately long and long, relatively slender setae scat- mally with 2 or 3 branches (rarely single). tered over dorsal and ventral surfaces, mesal surface membranous, lateral, outer part of dorsal Thorax: Setae 1-3-P on common setal support plate; and most of ventral surfaces with broad scales; 4-P single; 8-P branched, shorter than 4-P; 1-M,T gonostylus attached at apex of gonocoxite, long, nar- stout, single or 2-branched, inserted on large, pig- row, slightly curved, gonostylar claw short, narrow, mented tubercle (Assemblage A) or 1-M,T slender, attached at apex, 1 or more shorter, stout setae sub- with few branches, not on tubercle (Assemblage B); apically (these similar to gonostylar claw but some- 5-M longer than 7-M; 2,6-T single (Lu & Ji, 1997: what shorter and thinner, especially in Co. hatorii) fig. 44, of Co. ningheensis shows 2-T with 3 branches). (Assemblage A) or subapical, stout setae absent Abdomen: Seta 6-1,11 long, with 2 branches, 641 longer (Assemblage B); aedeagus simple, tubelike, distal than 6-I11; 7-1,11 long, single (rarely 2-branched); 12-I part wider than middle (Assemblage A) or approxi- present; 841 short, single or 2-branched; 1-VII long; mately same width as middle (Assemblage B); proc- 12-VII single; 2,4-VIII single; comb comprising numer- tiger with few tiny, cercal setae; claspette with ous scales in patch; segment X with moderate saddle, single, moderately long, columnar stem with small incomplete ventrally, with numerous stout spicules on subapical, thumblike lobe (Assemblage A) or subapi- posterior margin dorsal to seta 1-X, acus absent, 1-X cal lobe absent (Assemblage B), claspette filament inserted on saddle, ventral brush with several long, attached apically, long, relatively narrow, strongly branched setae with short stems, attached to grid with curved, apex pointed; sternum IX with several setae both transverse and lateral bars, some species with 1 on posteromesal area. shorter seta anterior to grid. Siphon: With attached acus; pecten with numerous Pupae closely set spines; seta 1-S single or branched, inserted within distal part of pecten or slightly distal to pecten. Cephalothorax: Setae 1,3-CT similarly developed; 11- CT normally single, rarely 2-branched. Abdomen: Seta 3-1,11 branched (Assemblage A) or Included species normally single (Assemblage B), longer than 64,11; 6- Assemblage A: Collessius elsiae elsiae (Barraud), Co. I shorter than 74; 141 with several branches; 54I elsiae vicarius (Lien), Co. hatorii (Yamada), Co. mac- inserted lateral to 4-1I; 3-I11 single (Co. banksi sin- dougalli (Edwards), Co. macfarlanei (Edwards), Co. gle or 2-branched); 5-V long, stout, single, longer ningheensis (Lei) and Co. shortti (Barraud). Assem- than tergum VI median length; 2-VI inserted mesal blage B: Collessius banksi (Edwards), Co. pseudotae- to 1,3-VT (Assemblage A) or 2-VI inserted lateral to niatus (Giles) and Co. tonkinensis (Galliard & Ngu). 1-VT and mesa! to 3-VI (Assemblage B); 6-Vu inserted posterior and mesa! to 9-WI; 9-VIII inserted slightly anterior to posterolateral corner (Assemblage Distribution A) or on corner (Assemblage B), with multiple branches. Assemblage A: China, India, Indonesia, Japan, Malay- sia, Pakistan, Sri Lanka, South Korea, Taiwan, Thai- Paddle: Apical margin rounded; midrib extends to land and Vietnam. Assemblage B: Bangladesh, India, apex of paddle; without hairlike spicules on margins; Myanmar, Nepal, Philippine Islands, Sri Lanka and seta 1-Pa short, single. Vietnam.

Bionomics fossa covered with pale scales; acrostichal (anterior Assemblage A: Immatures are normally found in rock and posterior) and dorsocentral (anterior and poste- pools but also in artificial containers (e.g. concrete rior) areas with setae; scutellum with narrow, curved water tanks). Females of some species bite humans scales on lateral lobes; paratergite with broad, white scales; antepronota widely separated, with broad and indoors and outdoors. Assemblage B: Immatures have been collected normally from rock pools but also from few moderately broad, pale scales; postpronotum with tree holes, and artificial containers (e.g. cement sinks broad, pale and dark scales, Da. echinus normally also and gutters, drains, iron cisterns, tires, etc.). Females with few narrow, curved scales dorsally; broad, white of Co. pseudotaeniatus bite humans indoors and scales on postspiracular area, subspiracular area, outdoors. upper proepisternum, mesokatepisternum (upper and lower posterior patches), upper and lower prealar areas, mesepimeron (1 large patch); mesepimeron Discussion without lower setae. Species of Collessius can be grouped into Assemblages Wing: Dark-scaled; upper calypter with several setae based on morphological characters (see above). Bar- on margin; alula with row of narrow scales on margin; raud (1934: figs 37, 39) illustrated the development of remigium with 2 or 3 setae on dorsal surface distally. the gonostylus and fourth-instar larva of Co. elsiae and Co. shortti. See Tanaka et al. (1979: figs 97, 98, Legs: Ante- and postprocoxal membranes bare; hind- 219) for illustrations of the male genitalia, fourth- femur with well-developed apical, white-scaled band; instar larva and adults of Co. hatorii, Tanaka (2002: hindtibia and tarsus dark-scaled, ungues equal, sim- fig. 1) for the pupa and Moriya et al. (1973) for the egg ple; fore- and midungues equal, each with 1 tooth. of this species. Abdomen: Tergum I with pale-scaled patch on laterotergite; segment VII laterally compressed. Etymology Genitalia: Tergum VIII heavily pigmented, basolat- Collessius is named in honour of Dr Donald H. Colless eral corners expanded into short, proximally curved, in recognition of his contributions to the taxonomy of thumblike projections, apex broadly rounded, numer- Oriental mosquitoes. The generic name is masculine, ous broad scales on distal 0.56-0.58; sternum VIII formed from his surname and the masculine Latin suf- heavily pigmented, base straight or very slightly con- fix -ius. Recommended abbreviation = Co. cave mesally, apex noticeably wider than base, nearly straight but with shallow, median emargination, DA.HLIANA REINERT, HARBACH & KITCHING, broad scales absent or very few (Da. geniculata) or GEN. NOV. several scales forming lateral patches (Da. echinus), 2 moderately long setae slightly subapical on lateral Type species: Culex geniculatus Olivier, 1791. part of apical margin, width greater than length; tergum IX heavily pigmented with median area slightly lighter, apex with moderately deep, V-shaped, median Females emargination, about 6 moderately long, stout setae Head: Median area of vertex with narrow, curved, pale apically on each side of midline; postgenital lobe mod- scales; occiput and vertex with numerous erect forked erately long, relatively narrow, apex rounded, setae on scales; ocular line narrow, with narrow, white scales; distal part; upper vaginal sclerite moderately large to eyes above antennal pedicels separated by width of 2- large, heavily pigmented; lower vaginal sclerite 4 ocular facets; antennal pedicel with few short, fine absent; insula liplike, with few setae in lateral setae, few small, broad scales sometimes present on patches; cercus relatively long, moderately wide, apex mesa] surface; clypeus bare; maxillary palpus and pro- broadly rounded, without scales; 3 spermathecal boscis dark-scaled, proboscis longer than forefemur. capsules. Thorax: Scutum with narrow, curved scales; pale scales forming narrow, longitudinal stripe on anterior and posterior acrostichal area and extending posteri- Males orly to or near prescutellar area, posterior dorsocen- Head: Maxillary palpus with 5 palpomeres, slightly tral area extending posteriorly laterad of prescutellar shorter than proboscis, dark-scaled, numerous long setae to posterior margin of scutum, and on anterior and moderately long setae ventrolaterally on distal and lateral margins of prescutellar bare area; white part of palpomere 3 and ventrolaterally on palpomeres scales on antealar area and extending posteriorly as 4 and 5; antennal flagellomeres with numerous long narrow, longitudinal stripe on supraalar area; scutal setae directed dorsally and ventrally.

Legs: Foretarsus with 2 unequal ungues, larger slightly anterior to 7-C; 7-C moderately long, moder- unguis with 2 teeth, smaller with 1 tooth; midtarsus ately stout, with 2 or 3 branches; 12-C branched, with 2 unequal ungues, each with 1 tooth; hindtarsus inserted mesa] to 13-C; 13-C single, longer than 12-C; with 2 equal ungues, simple. 14-C short, branched; 19-C absent; antenna moderately long, narrow, without spicules, seta 1-A moder- Genitalia: Tergum IX comprising 2 relatively broad, ately long, single or 2-branched. lateral plates each with short, narrow, strongly sclerotized lobe on posteromesal area bearing 2-6 short, Thorax: Setae 1-3-P inserted on common setal sup- stout, flattened, dark setae; gonocoxite moderately port plate; 5-P with 2 branches, longer than 6-F; 8-P long, relatively narrow, dorsal surface without apical short, multiple-branched; 4-M single; 5-M longer than and basal lobes on mesal margin, ventral surface with- 7-M; 2,6-T single. out specialized setae or scales, mesal surface membranous, numerous scales on dorsal, lateral and ventral Abdomen: Seta 6-1,11 with 2 long, stout, aciculate surfaces; gonostylus attached at apex of gonocoxite, branches; 7-1,11 long, stout, single; 12-I present; 84I relatively long, narrow, distal part gently curved single; 1-VII moderately long, stout, stellate, multiple- mesally, single gonostylar claw, long, narrow, attached branched; 12-VII single; 2,4-VIII single; segment VIII at apex; aedeagus simple, tubelike, lateral margins with comb comprising several scales in single curved gently curved to apex, distal 0.50 narrowest; proctiger row; segment X with large saddle, incomplete ven- with few to several cercal setae; claspette with single trally, acus absent, seta 1-X long, stout, branched, moderately long to long, narrow, columnar stem, inserted on saddle, ventral brush with several long, claspette filament attached apically, long, somewhat normally 2-4-branched setae with long stems, flattened, slightly curved to narrow, pointed apex; inserted on grid with both transverse and lateral bars, sternum IX with several setae on posteromedian area. 2 precratal setae. Siphon: With attached acus; pecten with several, closely set spines; seta 1-S moderately long, multiple- Pupae (based on Da. geniculat& branched, inserted distal to pecten. Cephalothorax: Setae 1,3-CT similarly developed, normally with 2 branches; 11-CT single. Included species Abdomen: Seta 3-I single, shorter than 6-I; 6-I longer Dahliana echinus (Edwards), Da. geniculata (Olivier) than 74; 241 inserted lateral to 1,3-I1; 341 moderately and Da. gilcolladoi (Sanchez-Covisa Villa, Rodriguez long, stout, normally single (occasionally 2-forked), Rodriguez & Guillen Llera). shorter than 6-I1; 541 long, stout, single, noticeably longer than 3-I1, slightly shorter than 6-I1, inserted lateral to 4-1I; 3,6411 long, stout, single; 5-V long, Distribution stout, single, longer than tergum VI median length; Albania, Algeria, Austria, Belarus, Belgium, Bulgaria, 2-VI inserted anterior and slightly lateral to 1-VI and Croatia, Czech Republic, Denmark, France, European mesal to 3-VI; 6-WI short, slender, inserted posterior Russia, Finland, Germany, Greece, Hungary, Iran, and slightly mesal to 9-VII; 9-VII moderately long, Italy, Lithuania, Macedonia, Morocco, Netherlands, stout, aciculate, usually with 2 branches; 9-VIII mod- Norway, Poland, Portugal, Romania, Slovenia, Slova- erately long, stout, aciculate, with 2-5 (usually 2 or 3) kia, Spain, Sweden, Switzerland, Turkey, Ukraine, branches. United Kingdom and Yugoslavia. Paddle: Length noticeably greater than width; midrib well developed, reaching apex of paddle; without hairlike spicules on margins; seta 1-Pa short, slender, Bionomics single. Immature stages normally inhabit treeholes but have been found in open tree stumps, root holes of olive trees and pools in rocks. Fourth-instar larvae Head: Seta 1-C single, moderately stout; 4-C moder- Discussion ately long, slender, multiple-branched, inserted mesal See discussion under Classification of Evaluated Taxa. and slightly anterior to 6-C; 5-C long, stout, single, Marshall (1938: figs 12, 57(g), 69, 94 and p1. IV) pro- inserted posterior to 4,6,7-C; 6-C moderately long to vided illustrations of the egg, first- and fourth-instar long, stout, single to 3-branched, inserted anterior and larvae, male genitalia and female of Da. geniculata, approximately same lateral level to 5-C and mesa] and and Reinert (2002e: fig. 3) illustrated the female gen-

Journal compilation © 2006 The Linnean Society of London, Zoological Journal of the Linnean Society, 2006, 148, 1-101 No claim to original US government works. 68 J. F. REINERT ETAL. italia of this species. Villa, Rodriguez & Liera (1985) patches of broad, pale scales, 1 on upper and 1 on mid- provide photographs of the fourth-instar larvae of dle, scale patches nearly meet in some specimens, sev- Da. echinus, Da. geniculata and Da. gilcolladoi. eral upper setae, lower setae absent; metameron bare. Wing: Dark-scaled, some specimens with 1-3 pale Etymology scales at base of costa; upper calypter with several Dahliana is named in honour of Dr Christine Dahl in setae on margin; alula with row of narrow, dark scales recognition of her many important contributions to on margin; remigium with 1-3 setae distally on dorsal the taxonomy and bionomics of Palaearctic mosqui- surface. toes. The generic name is feminine, formed from her Legs: Hindfemur with pale-scaled apex; hindtarsus surname and the feminine Latin suffix -iana. Recom- with broad, pale-scaled, basal bands on tarsomeres mended abbreviation = Da. 1-4; fore-, mid- and hindtarsi with 2 equal ungues, each with 1 tooth. DoBRorwoRsIYIus REINERT, HARBACH & Abdomen: Tergum I with patch of pale scales on lat- KITCHING, GEN. NOV. eral tergite; terga dark-scaled with dorsomesal pale- scaled areas basally and basolaterally, not forming Type species: Aedes tubbutiensis Dobrotworsky (1959) complete bands. Genitalia: Tergum VIII with apex broadly rounded, Females numerous broad scales on distal half; sternum VIII Head: Vertex with decumbent scales narrow, curved, sclerotized, length equal to or slightly longer than pale and dark; erect forked scales numerous on occiput width, apical margin with median emargination sep- and vertex; ocular line narrow, covered with narrow, arating 2 moderate lobes, several to moderate number curved, pale scales; 2-4 long, interocular setae; eyes of broad scales on lateral areas, seta 2-S inserted pos- above antennal pedicels contiguous (but separated terior to 1-S; tergum IX comprising 2 pigmented, lat- by approximately width of 3.5 eye facets in eral plates normally connected mesally by lightly Dh. rubrithorax); antennal pedicels with several pigmented, narrow, basal strip, inner margin of plates small, broad, pale scales on mesa] surface; clypeus darkly pigmented distally, setae absent (Db. ,nilsoni bare; maxillary palpus dark-scaled with pale scales at occasionally with 1 seta distally on 1 or both plates); apex of palpomeres 3 and 4; proboscis longer than postgenital lobe moderately long, moderately wide but forefemur, dark-scaled (few pale scales intermixed in distal part narrower, apex normally flat or with Db. alboannulatus and Db. tubbutiensis). minute emargination, several setae distally; upper vaginal sclerite small; lower vaginal sclerite absent; Thorax: Covered with narrow, curved scales insula liplike, transversely U-shaped (depressed), sev- (Db. alboannulatus and Db. milsoni also with few eral setae in lateral patches; cercus moderately long, broad, pale scales on lateral margins of bare prescutel- moderately wide distally but proximal part wider, lar area), both dark and pale scales present, pale apex broadly rounded, without scales; 3 spermathecal scales on lateral margin of bare prescutellar area capsules. mesal to setae; setae on following: few on anterior promontory, antedorsocentral area, scutal fossa and antealar area, numerous on acrostichal (anterior and Males posterior), dorsocentral (anterior and posterior), supraalar and prescutellar areas; scutellum with nar- Head: Maxillary palpus with 5 palpomeres, slightly row, curved scales on all lobes; antepronota widely shorter than proboscis, palpomeres 4 and 5 slightly separated, with pale scales, normally narrow, curved; down-turned, numerous long setae on distal part of postpronotum with both narrow, curved and broad palpomere 2 ventrally and palpomere 4 ventrolater- scales, few posterior setae; prespiracular setae absent; ally; antenna flagellar whorls with numerous long hypostigmal area bare; postspiracular area with setae directed dorsally and ventrally. scales, several setae; subspiracular area with broad, Legs: Fore- and midtarsi with 2 unequal ungues, pale scales; upper proepisternum with broad, pale larger unguis with 2 teeth, smaller unguis with 1 scales, several setae on upper area, lower area bare; tooth, some species with larger midunguis with 1 paratergite with pale scales; mesokatepisternum with tooth; hindtarsus with 2 equal unguis, each with 1 broad, pale scales in upper and lower patches, 3 or 4 tooth. upper and several posterior setae; prealar area with broad, pale scales in patch on lower area and few on Genitalia: Tergum IX with narrow, median area sepa- knob, several upper setae; mesepimeron with 2 rating broad, lateral areas, posterior margin with scle-

rotized, small, short, narrow lobe bearing few short, Abdomen: Seta 6-1,11 long, with 2 branches, 6-I1 longer slender setae on each side of midline; gonocoxite mod- than 6-111; 7-I long, single; 12-I present; 7-I1 short, slen- erately long, moderately wide, mesal margin of dorsal der, with several branches; 8-11 short, branched; 1-VII surface with small basal lobe bearing few moderately moderately long, moderately stout; 12-VII normally long and short, relatively slender setae, without apical single; 2,4-VIII single (2-branched in single available lobe, mesal surface membranous, lateral, outer part of specimen of Dh. rubrithorax); segment VIII with comb dorsal and most of ventral surfaces with broad scales; comprising numerous scales in patch; segment X with gonostylus attached at apex of gonocoxite, long, nar- moderate saddle, incomplete ventrally, acus absent, row, slightly curved, single gonostylar claw, long, nar- seta 1-X moderately long, inserted on saddle, ventral row, attached at apex; aedeagus simple, tubelike; brush with several long, multiple-branched setae with proctiger with few tiny, cercal setae; claspette with short stems, inserted on grid with well-developed single, moderately long, narrow, columnar stem, clas- transverse and lateral bars, anterior 2 setae shorter pette filament attached apically, long, somewhat flat- and occasionally 1 of these inserted anterior to grid. tened beyond middle, distal part tapering to narrow, Siphon: With attached acus; pecten with numerous, curved point; sternum IX with several setae on pos- closely set spines, seta 1-S moderately long, moder- teromedian area. ately stout, aciculate, with several branches, inserted slightly distal to pecten or rarely level with distal 1 or 2 pecten spines. Pupae Cephalothorax: Seta 1,3-CT similarly developed; 11-CT single or 2-branched. Included species Dohrotworskyius alboannulatus (Macquart), Db. Abdomen: Seta 3-1,11 branched, 3-I longer than 6-I, milsoni (Taylor), Db. occidentalis (Skuse), Db. 341 longer than 64I; 6-I longer than 7-I; 141 with mul- rubrithorax (Macquart), Dh. rupestris (Dobrotworsky), tiple branches; 541 inserted lateral to 44I; 3-I11 single, Db. subbasalis (Dobrotworsky) and Db. tubbutiensis longer than 5-I11; 5-V long, stout, single, longer than (Dobrotworsky). tergum VI median length; 2-VI inserted mesal to 1,3-VI; 6-VII inserted posterior and slightly mesa] to 9-Vu, both with multiple branches; 9-VIII inserted on pos- Distribution terolateral corner, with multiple branches. Australia. Paddle: Apical margin rounded; midrib extending to or near apex of paddle; without hairlike spicules on Bionomics margins; seta 1-Pa short, slender, single. 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 habi- Fourth-instar larvae tats some species also were collected from old wells, water collections around houses, artificial containers, Head: Seta 1-C dark, spiniform; 4-C short, slender, rot-hole in a tree and cavities in logs. Females of most with few branches, inserted mesal and approximately species have been collected biting humans during the in line or slightly posterior or anterior to 5-C and day, but also in the evening and at night. 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 Discussion stout, with several branches, inserted at approxi- See Dobrotworsky (1959, 1962, 1965) for illustrations mately same level or slightly anterior to 7-C; 7-C mod- and descriptions of most life stages of species in- erately long, relatively slender, aciculate, with several cluded in Dobrotworskyius. Russell (1996) provided branches, inserted lateral and anterior to 5-C; 12-C photographs and illustrations of females of short, slender, with several branches, inserted mesal Db. alboannulatus, Db. rubrithorax and Db. rupestris. to 13-C; 13-C single, longer than 12-C; 14-C short, Linley et al. (1991a) provided descriptions and single; 19-C absent; antenna moderately long, with photographs of the eggs of Db. alboannulatus and numerous spicules, seta 1-A moderately long, acicu- Dh. rubrithorax. late, with several branches. Thorax: Setae 1-3-P on common setal support plate Etymology (in few available species examined); 4,8-P branched; Dobrotworskyius is named in honour of Dr Nikolai V. 6-T normally single. Dobrotworsky in recognition of his important contri-

butions to the taxonomy of Australian mosquitoes, tively short, moderately wide, apex broadly rounded, especially of Victoria. The generic name is masculine, without scales; 3 spermathecal capsules. formed from his surname and the masculine Latin suffix -ius. Recommended abbreviation = Db. Males Head: GEORGECRAIGIUS REINERT, HARBACH & Maxillary palpus with 5 palpomeres, shorter KITCHING, GEN. NOV. than proboscis; antennal flagellar whorls with numerous long setae directed dorsally and ventrally. Type species: Culex atropalpus Coquillett, 1902. Legs: Fore- and midtarsi with 2 unequal ungues, larger unguis with 2 teeth, smaller with 1 tooth; hind- Females tarsus with 2 equal ungues, simple. Genitalia: Head: Vertex with decumbent scales narrow, curved, Tergum IX with pair of lateral lobes, rela- pale mesally, broad laterally, few erect forked scales; tively narrowly connected mesally, posterior margin occiput with numerous erect forked scales; ocular line relatively flat but each side gently sloping to midline, narrow with narrow, curved, pale scales; eyes moder- without setae; gonocoxite moderately long, relatively ately to broadly separated above antenna] pedicels; narrow, mesal margin of dorsal surface without apical antennal pedicel with few, small, broad scales and few lobe, with small, darkened, basal lobe bearing 2-4 short, slender setae mesally; clypeus bare; maxillary long, relatively stout setae and several slender setae, palpus and proboscis dark-scaled. ventral surface with row of long, relatively stout, more or less evenly spaced setae on mesal margin, mesal Thorax: Acrostichal area without anterior and poste- surface membranous; gonostylus attached at apex of rior setae; anterior dorsocentral area without setae, gonocoxite, long, narrow, curved mesally, with single, posterior dorsocentral setae present; supraalar area relatively short, narrow, gonostylar claw at apex; with longitudinal patch of pale scales; prescutellar aedeagus simple, tubelike, distal half slightly wider, area with pale scales on lateral margins of median, apex convex; proctiger with cercal setae; claspette bare space; scutellum with narrow, curved scales on with single moderately long to long, narrow, columnar all lobes; scales present on antepronotum, postprono- stem, claspette filament attached apically, long, some- tum, subspiracular area (broad), upper proepisternum what flattened, slightly curved; sternum IX with 2-12 (broad), mesokatepisternum (broad in upper and setae. lower posterior patches), lower prealar area (broad) and mesepimeron (broad); mesepimeron without lower setae; metameron bare. Pupae Wing: Dark-scaled with white-scaled patch at base of Cphalothorax: Seta 11-CT single, longer than costa; upper calypter with several setae on margin; 10,12-CT. alula with row of narrow scales on margin; remigium Abdomen: Seta 3-I long, stout, single (rarely 2- without setae. branched on 1 side of Ge. atropalpus), noticeably Legs: Hindfemur with band of pale scales at apex; longer than 64; 7-I longer than 64 141 very slender, hindtarsomeres 1 and 2 with narrow, pale-scaled, multiple-branched; 241 short, lateral to 1,3-I1; 3-I11 basal band, hindungues simple; fore- and midungues single, long, stout, inserted at same level as 1-I11; 5-I11 each with 1 tooth. inserted mesal to 4-I11; 6-WI short, slender, single or 2-branched, inserted mesa] and posterior to 9-Vu; Genitalia: Tergum VIII with apex broadly rounded, 9-VII stout, branched, longer than 6-Vu; 9-VIII mod- numerous scales on much of surface; sternum VIII erately long, stout, multiple-branched. scierotized, width slightly greater than length, apical Paddle: Wide; without hairlike spicules on lateral margin broadly rounded, numerous scales on much of margin; midrib extends to apex of paddle; seta 1-Pa surface, seta 2-S inserted posterior to 1-S; tergum IX single. 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 Fourth-instar larvae setae distally; upper vaginal scierite small, moder- Head: Seta 1-C slender to moderately stout, apex ately pigmented; lower vaginal sclerite absent; irisula attenuate, single; 4-C short, slender, branched, liplike, with few setae in lateral patches; cercus rela- inserted mesal and slightly posterior to 6-C; 5-C

Journal compilation © 2006 The Linnean Society of London, Zoological Journal of the Linnean Society, 2006, 148, 1-101 No claim to original US government works. PHYLOGENY AND CLASSIFICATION OF FINLAYA 71 moderately long to long, stout, inserted posterior to Bionomics 6-C; 6-C moderately long, stout; 7-C moderately Immature stages usually are collected from rockholes long, moderately stout, branched, inserted approxi- and rock pools, but also are found in artificial contain- mately at same level or slightly posterior to 5-C; ers, and occasionally treeholes, ground pools, stream 12-C short, branched, shorter than and inserted pools and cavities in agave leaves. Wood et al. (1979) mesal to 13-C; 13-C single; 14-C short, single; 19-C provide a summary of egg development for Gc. absent; antenna moderately long, with few to sev- atropalpus. eral spicules. Thorax: Seta 4-P single; 5-M longer than 7-M; 2,6-T single. Discussion Georgecraigius includes two subgenera, George- Abdomen: Seta 12-I present; 6-1-111 long, aciculate, craigius and Horsfallius (see descriptions below). branched; 7-I long, stout, aciculate; 7,10,12-VII single; Zavortink (1972) provided descriptions and illustra- segment VIII with comb comprising numerous scales tions of the members of this genus (as Atropalpus in patch, seta 2-VIII longer than 1-VIII, 2,4-VIII sin- Section). Yamaguti & LaCasse (1951), Carpenter & gle; segment X with relatively small saddle, incom- LaCasse (1955) and Wood et al. (1979) also provide plete ventrally, acus absent, seta 1-X short, slender, descriptions and illustrations of Gc. atropalpus. See single, inserted ventral to saddle (occasionally on ven- Linley & Craig (1994) for descriptions and photo- tral margin of saddle in Gc. fluviatilis), ventral brush graphs of Gc. atropalpus and Gc. epactius eggs. attached to grid with both transverse and lateral bars, without precratal setae. Siphon: With acus attached; pecten with numerous Etymology spines, evenly spaced or with distal 1 or 2 spines more Georgecraigius is named in honour of Dr George widely spaced; seta 1-S moderately long, aciculate, Brownlee Craig, Jr in recognition of his pioneering multiple-branched. 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 Included species his first name, surname and the masculine Latin suf- Gc. Georgecraigius atropalpus (Coquillett), Gc. epaclius fix -ius. Recommended abbreviation = (Dyar & Knab) and Gc. fluviatilis (Lutz). GEORGECRAJGJUS SUBGENUS GEORGECRAIGIUS REINERT, HARBACH & KITCHING Distribution Females Argentina (Misiones), Bolivia (Cochabamba), Brazil (Bahia, Goias, Minas Gerais, Para, Rio de Janeiro, Sao Thorax: Antealar scales pale; paratergite without Paulo, Sergipe), Canada (Labrador, Newfoundland, scales; postpronotum with broad scales; postspiracu- Ontario, Quebec), Colombia (Boyaca, Cundinamarca, lar area with patch of broad, white scales. Meta), Costa Rica (Alajuela, Cartago, Guanacaste, San Jose), Ecuador (Zamora), El Salvador, French Legs: Postprocoxal membrane without scales; hindtar- Guiana (Guyane, Inini), Guatemala (Guatemala), somere 2 with narrow, apical, pale-scaled band. Guyana (Essequibo), Honduras (Colon), Mexico (Baja California Sur, Chiapas, Guerrero, Morelos, Nuevo Abdomen: Tergum I without scales on laterotergite. Leon, Oaxaca, Puebla, Veracruz), Nicaragua (Zelaya), Panama (Canal Zone, Chiriqui, Code, Colon, Darien, Panama), Surinam (Suriname), United States (Ala- Males bama, Arizona, Arkansas, Colorado, Connecticut, Head: Maxillary palpus with few short setae ventrally District of Columbia, Georgia, Indiana, Kansas, on distal part of palpomere 3 and ventrolaterally on Kentucky, Louisiana, Maine, Maryland, Massachuset- palpomere 4, palpomeres 3 and 4 not down-turned. tus, Michigan, Minnesota, Missouri, Nebraska, New Hampshire, New Jersey, New Mexico, New York, North Carolina, Ohio, Oklahoma, Pennsylvania, Pupae Rhode Island, South Carolina, Tennessee, Texas, Utah, Vermont, Virginia, West Virginia, Wisconsin) Abdomen: Seta 5-11 inserted mesal to 4-11; 64I shorter and Venezuela (Aragua, Distrito Federal, Territorio than 7-I1; 6-Ill-V short; 5-IV—VI each less than or Amazonas). approximately equal to following tergum; 2-VI,VII

inserted lateral to or at same level as 1-VI,VTI; Legs: Postprocoxal membrane with pale scales; hind- 8-VI,VII inserted ventrally. tarsomere 2 without apical, pale-scaled band. Abdomen: Tergum I with broad, pale scales on Fourth-instar larvae laterotergite. Head: Seta 5-C long, single. Thorax: Setae 1-3-P and 5,6-P not inserted on com- Males mon setal support plate; 4-M single. Head: Maxillary palpus with several to numerous Abdomen: Seta 741 long, developed similar to 74; moderate to long setae ventrally on distal part of 13-Ill—V moderately long, branched; 1-WI short, slen- palpomere 3 and ventrolaterally on palpomere 4, der, single. palpomeres 3 and 4 down-turned. Siphon: Pecten with distal 1 or 2 spines more widely spaced; seta 1-S inserted within pecten. Pupae Abdomen: Seta 541 inserted lateral to 4-I1; 641 longer Included species than 7-I1; 6-111-V relatively long; 5-IV-VI very long, Georgecraigus atropalpus and Gc. epactius. stout, each noticeably longer than following tergum; 2-VI,V1I inserted mesal to 1-VI,VII; 8-VI,Vfl inserted dorsally. Distribution Canada (Labrador, Newfoundland, Ontario, Quebec), Costa Rica (AJajuela, Guanacaste), El Salvador, Gua- Fourth-i nstar larvae temala (Guatemala), Mexico (Baja California Sur, Chi- apas, Guerrero, Morelos, Nuevo Leon, Oaxaca, Puebla, Head: Seta 5-C moderately long, multiple-branched. Veracruz), Panama (Panama) and United States Thorax: Setae 1-3-P and 5,6-P inserted on common (Alabama, Arizona, Arkansas, Colorado, Connecticut, setal support plates; 4-M branched. District of Columbia, Georgia, Indiana, Kansas, Ken- tucky, Louisiana, Maine, Maryland, Massachusetts, Abdomen: Seta 741 short, developed differently than Michigan, Minnesota, Missouri, Nebraska, New 74; 13-Ill—V long, single; 1-VII long, stout, aciculate, Hampshire, New Jersey, New Mexico, New York, with 2 branches. North Carolina, Ohio, Oklahoma, Pennsylvania, Siphon: Pecten with spines evenly spaced; seta 1-S Rhode Island, South Carolina, Tennessee, Texas, inserted slightly distal to pecten. Utah, Vermont, Virginia, West Virginia, Wisconsin).

Included species Bionomics Georgecraigius flu viatilis. Immature stages usually occur in rockholes and rock pools, but also have been found in artificial containers, and occasionally in treeholes, ground pools, stream Distribution pools and cavities in agave leaves. Argentina (Misiones), Bolivia (Cochabamba), Brazil (Bahia, Goias, Minas Gerais, Para, Rio de Janeiro, Sao Recommended abbreviation = Gec. Paulo, Sergipe), Colombia (Boyaca, Cundinamarca, Meta), Costa Rica (Alajuela, Cartago, San Jose), Ecua- GEORGECRAIGIUS SUBGENUS HORSFALLIUS dor (Zamora), French Guiana (Guyane, mini), Guyana REINERT, HARBACH & KITCHING, SURGEN. NOV. (Essequibo), Honduras (Colon), Mexico (Veracruz), Nicaragua (Zelaya), Panama (Canal Zone, Chiriqui, Type species: Culex flu viatilis Lutz, 1904. Code, Colon, Darien, Panama), Surinam (Suriname) and Venezuela (Aragua, Distrito Federal, Territorio Amazonas). Females Thorax: Antealar scales dark; paratergite with broad, pale scales (rarely absent?); postpronotum with nar- Bionomics row, curved scales dorsally; postspiracular area with- Immature stages usually are collected from rockholes, out scales. rock pools and stream pools. They are found occasion-

Journal compilation © 2006 The Linnean Society of London, Zoological Journal of the Linnean Society, 2006, 148, 1-101 No claim to original US government works. PHYLOGENY AND CLASSIFICATION OF FINLAYA 73 ally in artificial containers, ground pools and Wing: Dark-scaled, with or without small, pale-scaled treeholes. 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 Etymology distally on dorsal surface. Subgenus Horsfallius is named in honour of Dr Will- iam Robert Horsfall in recognition of his outstanding Legs: Femora without subapical pale-scaled bands; contributions to the biology of New World mosquitoes midfemur without median, longitudinal white-scaled and descriptions of the eggs of aedine species of the stripe on anterior surface; hindfemur with apex with Nearctic Region. The name is masculine, formed from white scales; tibiae and tarsi dark-scaled; fore- and his surname and the masculine Latin suffix -ius. midtarsi with 2 equal ungues, each with 1 tooth; hind- Recommended abbreviation = Hor. tarsi with 2 equal ungues, simple. Abdomen: Tergum I with pale-scaled patch on latero- GILEsIus REINERT, HARBACH & KITCHING, tergite; terga dark-scaled with basal pale-scaled GEN. NOV. areas; segment VII dorsoventrally flattened. Type species: Culex pulchri venter Giles, 1901 Genitalia: Tergum VIII with apex broadly rounded, nearly covered with broad scales; sternum VIII sclerotized, width slightly greater than length, apical mar- Females gin with small, median emargination, several broad Head: Vertex with decumbent scales narrow and scales on lateral areas, seta 2-S inserted posterior curved, several broad scales present or absent on ante- to 1-S; tergum IX comprising 2 moderately pigmented, rolateral area; erect forked scales numerous on occiput lateral plates connected mesally by lightly pigmented and vertex; ocular line narrow, with narrow, curved, area, several short setae distally on each plate; post- pale scales; 4 or 5 interocular setae; eyes separated by genital lobe relatively long, moderately wide, apex approximately width of 3 ocular facets; antennal flat, several setae distally; upper vaginal sclerite small pedicel with small, broad, pale scales on mesal sur- to moderate size; lower vaginal sclerite absent; insula face; clypeus bare; maxillary palpus dark-scaled, with liplike, few setae in lateral patches; cercus moderately or without pale scales on subbasal area; proboscis long, moderately wide throughout length, apex longer than forefemur, dark-scaled. broadly rounded, numerous broad scales on dorsal surface; 3 spermathecal capsules. Thorax: Scutum covered with narrow, curved scales except bare median prescutellar area; scales pale on following: combined anterior acrostichal and anterior Males dorsocentral areas, moderate stripe on posterior acrostichal area, narrow stripe on posterior dorsocentral Head: Maxillary palpus with 5 palpomeres, shorter area, narrow stripe on outer margins mesad of setae than proboscis, palpomeres 4 and 5 slightly down- on prescutellar area, and moderate, longitudinal turned; antennal flagellar whorls with numerous long stripe on supraalar area; acrostichal (anterior and setae directed dorsally and ventrally. posterior) and dorsocentral (anterior and posterior) Legs: Fore- and midtarsi with 2 unequal urigues, setae short; prescutellar area with several setae lat- larger unguis with 2 teeth, smaller unguis with 1 erally; scutellum with narrow, curved scales on all tooth; hindtarsus with 2 equal ungues, simple. lobes, median lobe also with or without few broad scales; antepronota widely separated, with broad, pale Genitalia: Tergum IX with median part relatively scales, several setae; postpronotum with scales, few narrow with bibbed, pincushion-like structure bear- posterior setae; prespiracular setae absent; hypostig- ing numerous, short to moderately long, very slen- mal area with broad, pale scales; postspiracular area der, curved setae, lateral area developed as large, with pale scales, several setae; subspiracular area oblong lobe projecting cephalad; gonocoxite moder- with broad, pale scales; upper proepisternum with ately long, dorsal surface without apical and basal broad, pale scales, several setae, lower area bare; lobes on mesal surface, middle part of dorsal sur- paratergite with pale scales; mesokatepi sternum with face wide with mesal area broad, curved, projecting upper and lower patches of broad, pale scales, upper ventrally with row of short, stout, closely set, comb- and posterior setae; prealar area with broad, pale like setae on mesal margin, much of dorsal surface scales on upper and lower areas, numerous upper covered with short to moderately long, very slender setae; mesepimeron with 2 patches of broad, pale setae, ventral surface with elongate patch of numer- scales, 1 on upper and 1 on middle areas, several ous, long, moderately wide, fusiform scales on most upper setae, lower setae absent. of distal 0.50 of mesa] margin, mesa] surface mem-

Journal compilation © 2006 The Linnean Society of London, Zoological Journal of the Linnean Society, 2006, 148, 1-101 No claim to original US government works. 74 J. F. REINERT ET AL. branous, lateral, outer part of dorsal and most of Abdomen: Seta 6-1,11 long, stout, aciculate, with 2 ventral surfaces with broad scales; gonostylus branches, 6-I1 longer than 6-I11; 7-1,11 long, stout, attached at apex of gonocoxite, long, narrow but aciculate, single; 12-I present; 8-11 short, with 2 proximal part somewhat thicker, somewhat sigmoid- branches; 1-VU long, stout, aciculate, single; 12-Vu shaped, single gonostylar claw short, curved, spini- single; 2,4-VIII single; segment VIII with comb com- form; aedeagus simple, tubelike; proctiger with few prising numerous scales in patch; segment X with tiny, cercal setae; claspette with single, short to mod- moderate saddle, incomplete ventrally, with numer- erately long, narrow, columnar stem with small sub- ous stout spicules on posterior margin dorsal to apical lobe, claspette filament attached apically, long, seta 1-X, acus absent, 1-X moderately long to long, curved, relatively slender throughout length, apex stout, aciculate, single, inserted on saddle, ventral pointed; sternum IX with few setae (Gi. pulchriven- brush with several long, 2-5-branched setae with ter also with 2 or 3 broad scales) on posteromesal long stems, attached to grid with both transverse area. and lateral bars, anterior few setae somewhat shorter.

Pupae Siphon: With attached acus; pecten with numerous closely set spines; seta 1-S relatively long with several Cephalothorax: Setae 1,3-CT similarly developed; aciculate branches, inserted very slightly distal to 11-CT single, longer than 10-CT. pecten. Abdomen: Seta 3-1,11 single, longer than 6-1,11; 6-I shorter than 7-I 1-11 long, with 3 branches; 5-I1 inserted lateral to 4-I1; 3-111 relatively short, single, Included species approximately equal to 5-I11 length; 5-V moderately Gilesius alius (Lien) and Gi. pulchri venter (Giles). long, stout, single, shorter than tergum VI median length; 2-VI inserted mesa! to 1,3-VI; 6-VII single, inserted slightly posterior and mesal to 9-WI; 9-Vu Distribution with 3 branches; 9-VIII inserted on posterolateral cor- China, India, Nepal, Pakistan and Taiwan. ner, with multiple branches. Paddle: Apical margin rounded; midrib extends to Bionomics apex of paddle; without hairlike spicules on margin; Immature stages inhabit rock pools and treeholes but seta 1-Pa short, single. 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. pulchri venter Fourth-instar larvae have been collected from humans at dusk. 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, Discussion lightly aciculate, with several branches, inserted See Lien (1968b: figs 1, 6) for illustrations of the male mesal to 6-C; 6-C moderately long, stout, with 3 or 4 genitalia, pupa and fourth-instar larva of Gi. alius. 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 Etymology stout, lightly aciculate, with 4 or 5 branches; 12-C Gilesius is named in honour of Dr (Lt-Col) George short, with 2 or 3 slender branches, inserted mesal to Michael James Giles in recognition of his contribu- 13-C; 13-C single, longer than 12-C; 14-C short, mod- tions to the taxonomy of mosquitoes, primarily of erately stout, single or 2-branched; 19-C absent; India and surrounding areas. The generic name is antenna moderately long, narrow, with several small masculine, formed from his surname and the mascu- spicules, seta 1-A moderately long, with 3 or 4 (rarely line Latin suffix -ius. Recommended abbreviation = 2) slender branches. Gi. Thorax: Setae 1-3-P on common setal support plate, 3-P multiple-branched; 4-P relatively long; 5-P long, HIMMius REINERT, HARJ3ACH & KITCHING, stout, with 2 or 3 aciculate branches; 8-P short, stellate, with 5 or 6 branches, shorter than 4-P; 4-M sin- GEN. NOV. gle; 5-M longer than 7-M; 2,6-T single. Type species: Finlaya gilli Barraud, 1924.

Journal compilation © 2006 The Linnean Society of London, Zoological Journal of the Linnean Society, 2006, 148, 1-101 No claim to original US government works. PI{YLOGENY AND CLASSIFICATION OF FINLAYA 75

Females apex rounded, several setae distally; upper vaginal scierite small, lightly pigmented; lower vaginal scler- Head: Vertex with decumbent, narrow, curved, golden- ite absent; insula liplike, with few setae in lateral white to white scales mesally, broad, golden-white to patches; cercus relatively moderately long, moderately white scales laterally; occiput with numerous erect wide, apex rounded, mesal margin gently concave, forked scales; clypeus bare; maxillary palpus and pro- without scales; 3 spermathecal capsules. boscis dark brown-scaled, proboscis approximately equal to forefemur length. Thorax: Scutum with narrow, curved scales, golden- Males white scales covering approximately anterior 0.60 of Head: Maxillary palpus with 5 palpomeres, 0.86-0.95 scutum, similar scales forming narrow stripe on pos- length of proboscis, dark-scaled, distal part of pal- terior acrostichal area extending from anterior patch to anterior part of prescutellar area where it forks and pomere 3 bearing several long setae ventrolaterally, palpomeres 4 and 5 short, slightly down-turned, with extends along lateral margins of prescutellar bare area, similar narrow stripe on posterior dorsocentral several short and moderately long setae primarily on area extending posteriorly along lateral margin of ventral and distal areas. prescutellar setae, longitudinal stripe on supraalar Legs: Fore- and midtarsi with 2 unequal ungues, area, scutal fossa with small, posteromedian patch of larger unguis with 2 teeth, smaller unguis with 1 white scales, prealar area with small patch of some- tooth; hindungues equal, simple. what broader, narrow, white scales; setae on posterior dorsocentral area; scutellum with narrow, curved, Genitalia: Tergum IX comprising 2 moderately pig- golden-white scales on all lobes; paratergite without mented, lateral plates, each with small, short, narrow, scales; antepronota widely separated, with numerous heavily scierotized lobe on posterior margin bearing 5 moderately broad to broad, golden-white scales; or 6 short, stout setae; gonocoxite moderately long, postpronotum with golden-white scales, broad relatively narrow, dorsal surface without apical and scales on most of area except narrow, curved scales basal lobes on mesal margin, several scales present, dorsally; broad, white scales on upper proepisternum, ventral surface without specialized setae or scales, mesokatepisternum (upper and lower posterior mesal surface membranous; gonostylus long, attached patches), lower prealar area and mesepimeron (single at apex of gonocoxite, narrow but distal part narrower, large patch), other areas without scales; mesepimeron nearly straight, with single relatively short, narrow, without lower setae. gonostylar claw at apex; aedeagus simple, tubelike, moderately long, relatively narrow; proctiger with dis- Wing: Dark-scaled; upper calypter with several setae tal part heavily pigmented, apex flattened with 2 or 3 on margin; alula with row of narrow, dark scales on short, stout teeth, cereal setae present; claspette with margin; remigium with 2 or 3 setae on dorsal surface single long, narrow columnar stem, claspette filament distally. attached at apex of stem, long, relatively narrow, Legs: Ante- and postprocoxal membranes without slightly flattened, distal part terminating in long, scales; hindfemur with apex dark-scaled; hindtibia slender, pointed apex; sternum IX relatively short, dark-scaled with basal band of white scales, band nar- with several setae on posteromedian area. row dorsally, broader on anterior, posterior and ventral surfaces; hindtarsomeres 1-3 with broad, basal bands of white scales, hindurigues equal, simple; fore- Pupae and midungues equal, each with 1 tooth. Cephalothorax: Seta 1-CT very long, stout, with 2 Abdomen: Tergum I with white-scaled patch on lat- branches; 3-CT very long, with 3-5 branches; 4-CT erotergite; segment VII laterally compressed. shorter than 5-CT; 7-CT long, with 3-5 branches; 11-CT relatively long, normally with 2 branches Genitalia: Tergum VIII with broad scales covering all (single on 1 side of 1 specimen). but narrow basal strip; sternum VIII sclerotized, slightly wider than long, several broad scales on prox- Abdomen: Seta 3-I long, moderately stout, with 3 or 4 imal area, apical margin nearly straight with several branches; 6-I long, slender, single or 2-branched, moderately stout setae decreasing in length mesally, longer than 7-I; 2-I1 inserted lateral to 1-IT and mesal seta 2-S inserted posterior to 1-S; tergum IX compris- to 3-I1; 3-I1 long, stout, aciculate, with 2 or 3 ing lightly to moderately pigmented scierite with rel- branches; 4-I1 inserted mesal to 5-I1; 6-I1 long, slen- atively deep, median, apical emargination separating der, single to 3-branched, longer than 7-I1; 3-I11 long, pair of narrow lobes each bearing 3 or 4 short, slender stout, with 2 branches; 5-IV,V exceptionally long; 24V setae; postgenital lobe relatively long and narrow, inserted mesal to 1,3-TV; 6-VII short, slender, inserted

mesal and slightly anterior to 9-Vu; 9-VII moderately Included species long, stout, aciculate, with 3-5 branches; 9-VIII mod- Himalaius gilli (Barraud) and Hi. simlensis erately long, stout, aciculate, with 10 or 11 branches, (Edwards). The latter species is provisionally placed in inserted noticeably far anteriorly from posterior Himalaius. margin. Paddle: Ovoid; midrib well developed, reaching apex Distribution of paddle; without hairlike spicules on lateral margins; seta 1-Pa very long, 0.65-1.02 length of paddle, Himalaius gilli: India (Western Himalayas, Himachal single. Pradesh State, Krol Mountains near Solan, Naini Tal District, Uttar Pradesh State, Himachal Pradesh State, Uttaranchal State) and Nepal (Okaldunga Fourth-i nstar larvae District, Ramechhap District). Himalaius simlensis: India (Western Himalayas, Bhowali, Kumaon, Lansd- Head: With scooplike pouch anteroventral to antennal owne, Muktesar, Simla) (Barraud, 1934), and Him- base; seta 1-C relatively short, moderately thickened achal Pradesh and Uttaranchal (Kaur, 2003). throughout length but distal part slightly wider, apex bluntly pointed, single; 4-C very short, slender, with 6-9 branches, inserted near, mesa] and slightly poste- Bionomics rior to 5-C; 5,6-C very long, stout, dark, single, 5-C Barraud (1924, 1934) reported that the immature inserted near, mesal and slightly posterior to 6-C; 7-C stages of Hi. gilli (including those of the type series) long, stout, lightly aciculate, with 2 branches (3- were collected from water in treeholes. However, Kaur branched on 1 side of 1 specimen), inserted anterior (1992) states that Barrauds paratypes deposited in and lateral to 4-6-C and posterior to antennal base; the National Institute of Communicable Diseases, 12-C short, slender, with 6-9 branches, inserted mesa] New Delhi, India, were reared from larvae collected in to 13-C; 13-C moderately long, slender, with 3-5 bamboo. Darsie, Pradhan & Vaidya (1992) collected branches; 19-C absent; antenna long, narrow, with few pupae from turbid, chocolate-coloured water in tree- small spicu!es, seta 1-A single, extending beyond apex holes. The immatures of Hi. simlensis are not known. of antenna] shaft. Thorax: Seta 2-P longer than 1,3-F; 4-P moderately Discussion long, slender, with 3 or 4 branches; 6-P longer than Few specimens of Hi. gilli were available for examina- 7-P; 6,7-P longer than 5-P; 8-P relatively short, slender, tion but these bear distinct characters in all life stages with 2 or 3 branches; 1-M moderately long, slender, that readily distinguish the genus (see above descrip- with 3-6 branches, approximately 2.9 times length of tion based on Hi. gilli). Several abdominal segments of 2-M; 4-M with 3-6 branches; 5-M longer than 7-M; the larval specimens are in poor condition and could 2-T with 8-12 branches; 6-T with 3-5 branches. not be described. The species simlensis is provisionally included in Himalaius Abdomen: based on the characters men- Seta 12-I present; 6,74 moderately long, tioned in the brief and incomplete descriptions of stout, 6-I with 4-6 branches, 7-I with 4 or 5 branches, Edwards (1922) and Barraud (1924). only slightly shorter than 64; 641 moderately long, stout, with 3 or 4 branches; 741 relatively moderately long, slender, with 4-6 branches; 841 single; 1-Vu Etymology very long, moderately stout, with 2 or 3 branches; The generic name Himalaius is formed from Hima- 3-VII short, slender with 3-8 branches, inserted laya, the mountains, and the masculine Latin suffix !ateral and slightly anterior to 1-Vu; 7,10,12-Vu -ius, meaning pertaining to the Himalayas. Recom- branched; 1-VIII with 5 or 6 branches; 2,4-VIII mended abbreviation = Hi. branched; segment VIII with comb comprising 80-94 scales in patch; segment X with moderately large saddle, incomplete ventrally, acus absent, 1-X moder- JARNELLIUS REINERT, HARBACH & KITCHING, ately long, slender, with 3 branches, inserted on GEN. NOV. saddle, 3-X very long, stout, single, ventral brush Type species: Culex varipalpus Coquillett, 1902. inserted on grid with both transverse and lateral bars, 2 precratal setae. Siphon: Acus attached; pecten comprising 24-36 Females evenly spaced spines; seta 1-S short, slender, with 6-8 Head: Vertex with decumbent scales narrow (few branches, inserted distal to pecten. broad scales occasionally present laterally in some

Journal compilation © 2006 The Linnean Society of London, Zoological Journal of the Linnean Society, 2006, 148, 1-101 No claim to original US government works. PHYLOGENY AND CLASSIFICATION OF FINLAYA 77 species); occiput and vertex with erect forked scales; tally; upper and lower vaginal sclerites absent; insula eyes moderately separated above antennal pedicels; liplike, with setae in lateral patches; cercus moder- antennal pedicel with few to several, often partially ately long, relatively narrow with apex sharply overlapping, small, broad scales on mesal surface; rounded, without scales (moderately wide with apex clypeus bare; maxillary palpus dark with pale-scaled broadly rounded and 1-3 scales/cercus in Ja. muel- areas (all dark-scaled in Ja. muelleri); proboscis with- len); 3 spermathecal capsules. out median, pale-scaled band.

Thorax: Scutum with moderately wide to wide, Males median, pale-scaled stripe or patch extending over anterior acrostichal area laterally to inner dorsocen- Head: Maxillary palpus with 5 palpomeres, longer tral area or wider and extending laterally to mesal than proboscis, moderate number of moderately long margin of scutal fossa, pale-scaled, median area nor- to long setae ventrolaterally on distal part of pal- mally continuing posteriorly to prescutellar area pomere 3 and ventrolaterally on palpomere 4; anten- (except in Ja. vanipalpus); narrow, curved, white scales na] flagellar whorls with numerous long setae directed on scutal fossa (lateral and posterior margins), dorsally and ventrally. supraalar area (elongate longitudinal line) and pres- Legs: Fore- and midtarsi with 2 unequal ungues, each cutellar space (lateral and anterior margins of median, with 1 tooth (smaller ungues simple in Ja. muelleri); bare area); acrostichal and dorsocentral areas with hindungues equal, both simple. both anterior and posterior setae; paratergite with broad, pale scales; postpronotum with narrow, curved Genitalia: Tergum IX with posterior margin with pair scales dorsally and broad scales ventrally (except all of short, relatively narrow, heavily sclerotized lobes narrow scales in Ja. muelleni); postspiracular, upper bearing several short, stout setae; gonocoxite moder- proepisternal, lower prealar, mesokatepisternal (large ately long, relatively narrow, mesal margin of dorsal upper patch and moderate lower posterior patch), surface without apical lobe but with small basal lobe mesepimeral (upper and lower patches, except 1 large bearing several long setae, 1 or more of these setae patch in Ja. muelleri) and metameral (except Ja. deser- stout and somewhat flattened, ventral surface without ticola) areas with broad, pale scales. specialized scales or setae, mesal surface membranous; gonostylus attached at apex of gonocoxite, mod- Wing: Dark-scaled with small white-scaled patch at base of costa (pale-scaled patch reduced to 2 or 3 scales erately long, narrow, distal part slender and slightly curved mesally, with single moderately long, narrow or occasionally absent in Ja. muelleri); upper calypter with several setae on margin; alula with row of narrow gonostylar claw at apex; aedeagus simple, tubelike, moderately long, apex broadly rounded; proctiger with scales on margin; remigium without setae ventrally and normally absent dorsally (1 or 2 setae distally in cercal setae; claspette with single moderately long, slender, columnar stem, claspette filament attached Ja. ,nuelleri and Ja. vanipalpus). apically, relatively long, relatively narrow, somewhat Legs: Anteprocoxal membrane bare, postprocoxal flattened with distal part narrowing to pointed, membrane normally with broad, pale scales (absent in slightly recurved apex; sternum IX with setae on Ja. desenticola and Ja.sierrensis); hindtarsomere 1 posteromedian area. with broad, basal and apical, white-scaled bands, hindtarsomeres 2 and 3 with broad, apical, white- scaled bands (hindtarsus entirely dark-scaled in Ja. Pupae muelleni), ungues simple. Cephalothorax: Seta 1-CT longer than 3-CT, single or Abdomen: Tergum I with white-scaled patch on lat- 2-branched; 11-CT single. erotergite; segment VIII dorsoventrally flattened. Abdomen: Setae 3,64 single, moderately stout; 241 Genitalia: Tergum VIII sclerotized, width greater shorter and inserted mesal to 1-I1; 341 single, moder- than length, setae on distal 0.70 or more of tergum; ately long, stout, inserted mesa! to 2-1I; 541 less than sternum VIII sclerotized, width greater than length, 341 length, inserted mesal to 4-1I (lateral to 4-11 in seta 2-S inserted posterolateral to 1-S, scales normally Ja. muelleri); 3-I11 single, stout, longer than 5-I11; 5-V present (not seen in only available specimen of Ja. single, less than tergum VI median length (slightly monticola); tergum IX comprising single sclerite with greater than tergum length in Ja. muelleni); 2,3-VI moderately deep, median, apical emargination, 3-12 inserted lateral to 1-VT (? in Ja. laguna); 6-VII setae distally on each side of midline; postgenital lobe inserted posterior and mesa] to 9-VII (at same level in moderately long, relatively broad, apex broad with Ja. muelleri); 9-VII longer and stouter than 6-WI; very small, median emargination, 16-25 setae dis- 9-VIII stout, with 2 or 3 branches.

Paddle: Length greater than width; apex broadly Distribution rounded or with minute, broad, median emargination; Canada (British Columbia), Mexico (Baja California midrib extending 0.60-0.70 from base; without fringe Sur, Distrito Federal, Durango, Hidalgo, Mexico, of hairlike spicules; seta 1-Pa short, single. Sonora, Zacatecas) and United States (Arizona, Cali- fornia, Idaho, Montana, Nevada, New Mexico, Oregon, Texas, Utah, Washington). Fourth-instar larvae Head: Seta 1-C single, spiniform; 4-C short (barely Bionomics moderately long in Ja. muelleri), multiple-branched, inserted mesal and slightly anterior to 6-C; 5-C mod- Immature stages are usually found in treeholes. Jarnellius muelleri erately long (approaching long), single, inserted mesal are found occasionally in artificial and slightly anterior to 7-C and posterior to 4,6-C; 6-C containers and were once collected from leaf axils of moderately long, with 2 branches (single in Ja. muel- maguey (Zavortink, 1972). Insight into the ecology of treeholes in which most species of Jarnellius len); 7-C moderately long, multiple-branched; 12-C are found is given in Zavortink (1985). branched, inserted mesa! 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. Discussion The above generic description of Jarnellius is addi- Thorax: Setae 1-3-P inserted on common setal sup- tional to the description (Varipalpus Group) given by port plate; 4-P branched, longer than 8-P; 5-P Arnell & Nielsen (1972), who should be referred to for branched; 4-M single (rarely 2-branched in Ja. sierren- the systematic background, descriptions and illustra- sis); 5-M longer than 7-M; 2,6-T single. tions of the included species. Zavortink (1972) provided a description and illustrations of Ja. muelleni Abdomen: Setae 3-1—VI and 2,4-VIII single; 6-I—I11 (as Ae. (Och). muelleri), the type species of the new long, aciculate, branched; 7-I moderately long to subgenus Lewnielsenius. The genus includes two sub- long, stout, aciculate, single; 12-I absent (present in genera (see below). Myers (1967) described and illus- Ja. deserticola and Ja. sierrensis); 74I moderately trated eggs of Ja. sierrensis and Ja. varipalpus. long, single, except relatively short and branched in Ja. muelleni; 1-VT! longer than dorsal length of segment X (see method of measurement for character Etymology 49 in Character Descriptions); 2-VII inserted near 1-VT!; 3-VT! moderately long, inserted anterolat- Jarnellius is named in honour of Dr J. Hal Arnell in erad of 1-VT!; segment VIII with comb comprising recognition of his important work on species of this several scales in 1 or 2 irregular rows; segment X genus and other New World aedines. The generic with saddle incomplete ventrally, without acus, seta name is masculine, formed from his first initial J, sur- 1-X single or 2-branched, inserted on saddle in name and the masculine Latin suffix -ius. Recom- mended abbreviation = Ja. small unsclerotized area (reduced in Ja. muelleni), 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, JARNELLJLJS SUBGENUS JARNELLIUS REINERT, except relatively short in Ja. muelleri, 4 relatively HARBACH & KJTCHING long, broad anal papillae with apices broadly Females rounded. Head: Maxillary palpus dark-scaled with pale-scaled Siphon: Acus present; pecten comprising several areas. evenly spaced spines, on proximal 0.50 of siphon; seta 1-S branched, aciculate, inserted distal to pecten; 6-S Thorax: Scutal fossa with white-scaled lines on lateral short, single. 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 Included species white scales. Jarnellius deserticola (Zavortink), Ja. laguna (Arnell & Nielsen), Ia. monticola (Belkin & McDonald), Ja. Legs: Hindtibia dark-scaled with white scales basally; muelleri (Dyar), Ja. sierrensis (Ludlow) and Ja. van- hindtarsomere 1 with broad basal and apical, white- palpus (Coquillett). scaled bands, hindtarsomeres 2 and 3 with broad,

apical, white-scaled band; fore- and midungues each Genitalia: Cercus moderately long, moderately wide, with 1 tooth. apex broadly rounded, with 1-3 scales. Genitalia: Cercus moderately long, moderately wide,

apex sharply rounded, without scales. Males Head: Maxillary palpus dark-scaled. Males Legs: Fore- and midtarsi with larger ungues with 1 Legs: Fore- and midungues each with 1 tooth tooth, smaller ungues simple.

Pupae Pupae Abdomen: Seta 541 inserted lateral to 4-I1; 5-V Abdomen: Seta 541 inserted mesa] to 4-I1; 5-V longer shorter than median length of tergum VI; 6-WI than median length of tergum VI; 6-WI inserted mesal inserted posterior and mesal to 9-VII. and at approximately same level as 9-VII.

Fourth-instar larvae Fourth-instar larvae Head: Seta 6-C with 2 branches; antenna without spicules. Head: Seta 6-C single; antenna with few spicules. Abdomen: Seta 7-I1 developed similarly to 7-I, single Abdomen: Seta 741 developed differently from 74, rel- (occasionally 2-branched in Ja. sierrensis), moderately atively short, branched; ventral brush attached to grid long to long; ventral brush attached to grid with only with both transverse and lateral grid bars, seta 4a-X transverse grid bars, seta 4a-X long. relatively short.

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

Included species Included species Jarnellius muelleri (Dyar). Jarnellius deserticola, Ja. laguna, Ja. monticola, Ja. sierrensis and Ja. varipalpus. Recommended abbreviation = Jar. Etymology Subgenus Lewnielsenius is named in honour of Dr Lewis T. Nielsen in recognition of his many contribu- JARNELLIUS SUBGENUS LEWNIELSENIUS REINERT, tions to the taxonomy, distribution and bionomics of HARBACH & KITCHING, SUBGEN. NOV. North American mosquitoes, especially aedine species. The subgenus is masculine, formed from the combina- Type species: Aedes (Heteronycha) muelleri Dyar, tion of his first name, surname and the Latin mascu- 1920. line suffix -ius. Recommended abbreviation = Lew.

Females JIHLIENIus REINERT, HARBACH & KITCHING, Head: Maxillary palpus dark-scaled. GEN. NOV. Thorax: Scutal fossa with white-scaled areas on lat- Type species: Aedes (Finlaya) chungi Lien, 1968. eral and posterior margins that are expanded mesally; postpronotum with narrow scales; hypostigmal area with pale scales; mesepimeron with single, large patch Females of white scales extending over upper and lower areas. Head: Vertex with decumbent scales broad, pale on Legs: Hindtibia and hindtarsus entirely dark-scaled; median area and dark on lateral areas; erect forked fore- and midungues simple. scales numerous on occiput, few semierect scales on

vertex caudal to ocular setae; ocular line narrow, midtarsi with 2 equal ungues, each with 1 tooth; hind- covered with broad, pale scales; 2 long interocular tarsus with 2 equal ungues, simple. setae; eyes above antenna] pedicels contiguous or separated by less than width of 1 eye facet; anten- Abdomen: Tergum I with pale-scaled patch on lateral nal pedicels with few small, broad, dark scales on tergite; terga dark-scaled with small, basolateral, mesal surface; clypeus bare; maxillary palpus and pale-scaled patch; segment VII laterally compressed. proboscis dark-scaled; proboscis longer than Genitalia: Tergum VIII with apex broadly rounded, forefemur. numerous broad scales on most of surface; sternum VIII sclerotized, width greater than length, apical Thorax: Covered with narrow, relatively straight margin with median emargination, scales absent, seta scales except bare, median, prescutellar area and elon- 2-S inserted posterior to 1-S; tergum IX comprising 2 gate strip between supraalar and prescutellar setae, moderately pigmented, moderately long, lateral plates scales dark except for following pale-scaled areas: nar- connected mesally by basal pigmented strip, base row stripe on anterior promontory and extending cau- wider than apex, few short setae distally on each dally on acrostichal area to approximately level of plate; postgenital lobe long, relatively narrow, apex scutal angle where it becomes less distinct for short rounded, several setae distally; upper vaginal sclerite distance and then distinct on caudal part of posterior small; lower vaginal sclerite absent; insula liplike, acrostichal area, narrow line on caudal part of ante- with few setae in lateral patches; cercus moderately rior dorsocentral area and extending over posterior long, relatively wide, outer distal margin gently dorsocentral area to caudal margin of scutum, stripe oblique, without scales; 3 spermathecal capsules. broadened at juncture of prescutal suture into small triangular patch which connects with narrow stripe extending outward along scutal suture and cephalad Males along margin of scutal fossa onto antedorsocentral area, more or less elongate stripe on supraalar area, Head: Maxillary palpus with 5 slender palpomeres in and indistinct stripe on lateral margins of prescutellar nearly straight line, shorter than proboscis, few area mesad of setae; setae on following: few on ante- moderately long setae on distal part of palpomere 3 rior promontory, antedorsocentral area, dorsocentral ventrally and palpomere 4 ventrolaterally; antennal (anterior and posterior) area, scutal fossa and flagellar whorls with numerous long setae directed antealar area, numerous on supraalar and prescutel- dorsally and ventrally. lar areas; scutellurn with broad or broad and narrow Thorax: Scaled areas of scutum with narrow, rela- scales on lobes; antepronota widely separated, with tively long, pale scales except small area of scutal broad, pale scales, several setae; postpronotum with fossa with narrow, dark scales. pale scales, few posterior setae; prespiracular setae absent; hypostigmal and subspiracular areas bare; Legs: Fore- and midtarsi with 2 unequal ungues, postspiracular area with few setae, scales absent; larger unguis with 2 teeth, smaller unguis with 1 upper proepisternum with broad, pale scales, several tooth; hindtarsus with 2 equal ungues, simple. setae on upper area, lower area bare; paratergite nar- Genitalia: Tergum IX with median area narrow, lat- row, with broad, pale scales; mesokatepisternum with eral area developed as broad, rounded lobe projecting broad, pale scales in upper and lower patches, 3 or 4 cephalad, posterior margin with very small, sclero- upper and several posterior setae; prealar area with tized lobe on each side of midline bearing few short, broad, pale scales in patch on lower area and extend- slender, curved setae; gonocoxite moderately long, rel- ing onto lower part of knob, several upper setae; atively narrow, dorsal surface without apical and mesepimeron with 1 patch of broad, pale scales on basal lobes on mesa] margin, however, with small, upper area and extending onto middle, several upper dark, oval area basomesally bearing straight setae, setae, lower setae absent. few of these setae on most proximal part long and Wing: Dark-scaled with small pale-scaled patch at somewhat flattened, middle part with several moder- base of costa; upper calypter with several setae on ately long setae and distal part with several short margin; alula with row of narrow, dark scales on mar- setae, ventral surface with narrow, elongate patch of gin (Ji. chungi); remigium with 2 or 3 setae distally on moderately long, slender, lanceolate setae on distal dorsal surface. area of mesa] margin (Lu & Ji, 1997: fig. 34, shows narrow fusiform scales in this area of Ji. gonguoensis), Legs: Hindfemur with narrow basal and moderate mesal surface membranous, lateral and outer parts of preapical bands dark-scaled, remainder pale-scaled; dorsal and ventral surfaces with broad scales; gono- hindtibia and hindtarsus dark-scaled except tarsom- stylus attached at apex of gonocoxite, relatively short, ere 1 with broad, pale-scaled, basal band; fore- and narrow, single gonostylar claw, long, narrow, attached

Journal compilation © 2006 The Linnean Society of London, Zoological Journal of the Linnean Society, 2006, 148, 1-101 No claim to original US government works. PHYLOGENY AND CLASSIFICATION OF FINLAYA 81 at apex; aedeagus simple, tubelike, apex concave; nor margin dorsal to insertion of seta 1-X, acus proctiger with few tiny, cercal setae; claspette with absent, seta 1-X moderately long, slender, single or single, moderately long, narrow, columnar stem, clas- 2-branched, inserted on saddle, ventral brush with pette filament attached apically, long, somewhat flat- long, normally 2-4-branched setae with long stems, tened beyond midlength, with transverse striations on inserted on grid with both transverse and lateral bars, much of median area, apex curved into narrow point; 2 shorter, precratal setae. sternum IX with few setae on posteromedian area. Siphon: With attached acus; pecten with numerous, closely set spines; seta 1-S moderately long, slender, multiple-branched, inserted slightly distal to pecten. Pupae Cephalothorax: Setae 1,3-CT similarly developed; 10- CT single; 11-CT with 2 branches, shorter than 10-CT. Included species Jihlienius chungi (Lien) and Ji. unicinctus (Edwards). Abdomen: Seta 34,11 single, shorter than 6-1,11; 6-I Jihlienius gonguoensis (Gong & Lu) is provisionally longer than 74; 141 short, slender, with 2 branches; included based on illustrations in Lu & Ji (1997). 541 inserted lateral to 441; 3-I11 single, longer than 5-I11; 5-V long, stout, single, longer than tergum VI median length; 2-VI inserted mesal to 1,3-VI; 6-Vu Distribution 6-Vu inserted slightly posterior and mesal to 9-VII I China, India, Nepal and Taiwan. 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 Bionomics apex of paddle; without hairlike spicules on margins; Immatures have been collected from bamboo stumps seta 1-Pa short, single or 2-forked. and treeholes.

Fourth-instar larvae Discussion Much of the generic description is based on Ji. chungi Head: Seta 1-C stout, spiniform; 4-C short, slender, because published descriptions of other species were multiple-branched, inserted slightly mesal and inadequate, several life stages are unknown and/or approximately in line with 6-C; 5-C moderately long, specimens were not available for study. See Lien stout, single, inserted posterior to 4,6,7-C; 6-C moder- (1968a) for illustrations of the male genitalia, pupa ately long but shorter than 5-C, moderately stout, sin- and fourth-instar larva of Ji. chungi. gle, 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 Etymology slightly anterior to 5-C; 12-C short, with few branches, Jihlienius is named in honour of Dr Jih Ching Lien inserted mesal to 13-C; 13-C single, longer than 12-C; in recognition of his important contributions to the 14-C short, slender, single; 19-C absent; antenna mod- taxonomy of the mosquitoes of Taiwan. The generic erately long, narrow, without spicules, seta 1-A mod- name is masculine, formed from his first name, erately long, slender, single. surname and the Latin suffix -ius. Recommended Thorax: Setae 1-3-P inserted on common setal sup- abbreviation = Ji. port 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 PATMARKSIA REINERT, HARBACH & KITCHING, than 7-M; 2,6-T single. GEN. NOV. Abdomen: Seta 64,11 with 2 or 3 moderately long, Type species: Leucomyia australiensis variety papuen- moderately stout branches; 7-I moderately long, mod- sis Taylor, 1914. erately stout, with 2 branches; 6-I1 longer than 6-I11; 12-I present; 741 short, slender, with 3 or 4 branches; Females 841 with 2 or 3 branches; 1-VII moderately long, stout, single; 12-VII single; 2,4-VIII single; comb on segment Head: Vertex with median area covered with narrow, VIII comprising patch of more than 20 scales; segment curved, pale scales; occiput and vertex with erect X with moderately large saddle, incomplete ventrally, forked scales; interocular space with patch of narrow, with several moderately long, stout spicules on poste- curved, pale scales projecting ventrad over mesal mar-

Journal compilation © 2006 The Linnean Society of London, Zoological Journal of the Linnean Society, 2006, 148, 1-101 No claim to original US government works. 82 J. F. REINERT ETAL. gins of eyes; eyes contiguous or nearly so; antenna] Males pedicel with few short, fine setae and small, broad, Head: nonoverlapping scales on mesa] surface; clypeus bare; Maxillary palpus with 5 palpomeres, slightly maxillary palpus and proboscis dark-scaled. shorter to slightly longer than proboscis, numerous long setae ventrolaterally on distal part of palpomere Thorax: Scutum with narrow, curved, dark scales 3 and ventrolaterally on palpomere 4; antennal fiagel- except for moderate to broad median, longitudinal lar whorls with numerous long setae directed dorsally patch of white scales on approximately anterior and ventrally. 0.55-0.70 (white-scaled patch on acrostichal area and extending laterally over mesal part of dorsocentral Legs: Fore- and midtarsi with 2 ungues, both toothed. area or to near lateral part of scutal fossa, but not on Genitalia: Tergum IX comprising 2 narrow, lateral lateral part of scutal fossa), other scutal areas nor- plates narrowly joined mesally, without setae; gono- mally dark-scaled including supraalar and prescutel- coxite moderately long, moderately wide, mesal mar- lar areas (but not median, prescutellar bare area) (few gin of dorsal surface without apical lobe but with species with white scales along lateral margin of pres- small basal lobe bearing several short and moderately cutellar bare area); scutellum with narrow, curved long, slender setae, ventral surface without special- scales on all lobes; acrostichal (both anterior and pos- ized setae or scales on mesal margin, mesa] surface terior) areas without setae, anterior dorsocentral area membranous; gonostylus attached at apex of gonocox- normally without setae (occasionally 1 seta on caudal ite, moderately long, narrow, with single, short, part), posterior dorsocentral area with several setae; narrow, gonostylar claw at apex; aedeagus simple, paratergite with broad, pale scales; postpronotum nor- tubelike, moderately long, distal 0.50 widest; proctiger mally with patch of broad, scales ventrally and nar- with cercal setae; claspette with single moderately row, curved scales dorsally (no narrow scales seen in long, slender, columnar stem, claspette filament Pm. dobodura but this area rubbed in specimens attached apically, moderately long to long, somewhat examined; only narrow, curved, pale scales in Pm. ang- flattened, relatively narrow with distal part tapering giensis); mesokatepisternum with upper and lower to slender apex, middle part with transparent sheaf posterior patches of broad, pale scales; metameron with transverse striations encircling spiniform; ster- bare. num IX with setae on posteromedian area. Wing: Costa dark-scaled; upper calypter with several setae on margin; alula with row of narrow scales on margin; remigium with dorsal setae distally. Pupae Legs: Ante- and postprocoxal membranes bare; hind- Cephalothorax: Seta 11-CT single (rarely 2-branched femur with dorsoapical patch of pale scales; tibiae in Pm. argyronotum). dark-scaled (except narrow, pale-scaled, basal band in Abdomen: Seta 3-I long, stout, single, longer than 64; Pm. anggiensis); hindtarsus with broad, basal, pale- 6-I single, longer than 74; 241 short, inserted lateral scaled bands on tarsomeres 1-4. to 141; 341 long, stout, single, longer than 6-I1; 3-I11 Genitalia: Tergum VIII sclerotized, with numerous long, stout, single, longer than 5411; 5-IV,V long, stout, broad scales on distal part; sternum VIII sclerotized, longer than following tergum; 6,9-VII both relatively length greater than width, normally moderate to large stout, aciculate, branched; 9-VIII stout, aciculate, number of broad scales in lateral patches, apex with multiple-branched. small to moderately deep, median emargination sepa- Paddle: Broad; apex broadly rounded or flattened; rating pair of small to broad lobes, apical margin with midrib nearly reaching apex; without hairlike spicules numerous short, curved setae, seta 2-S inserted pos- on margin; seta 1-Pa single. terior to seta 1-S; tergum IX comprising 2 moderately pigmented, narrow, lateral plates connected by basal or subbasal pigmented or membranous band, distal Fourth-instar larvae part of mesal surfaces of plates darkly pigmented and strongly fused to basolateral margins of cerci, without Head: Seta 1-C relatively stout, single; 4-C short, setae; postgenital lobe with several setae distally; branched, inserted slightly posterior and mesal to upper vaginal sclerite small, moderately pigmented; 5-C; 5,6-C moderately long, multiple-branched, lower vaginal sclerite absent; insula liplike, V-shaped displaced far anteriorly, 6-C inserted lateral and medially (depressed), with setae in lateral patches; slightly anterior to 5-C; 7-C moderately long, multiple- cercus moderately long, moderately wide, apex branched, inserted lateral and posterior to 4-6-C; broadly rounded, without scales (rarely 1 or 2 scales 12-C short, branched, inserted mesal to 13-C; 13-C present); 3 spermathecal capsules. single; 14-C short, normally single; 19-C absent;

Journal compilation © 2006 The Linnean Society of London, Zoological Journal of the Linneon Society, 2006, 148, 1-101 No claim to original US government works. PHYLOGENY AND CLASSIFICATION OF FINLAYA 83 antenna with few to numerous spicules, 1-A nor- Etymology mally branched. Patmarksia is named in honour of Dr Elizabeth Nesta Marks in recognition of her pioneering and outstand- Thorax: Setae 1-3-P inserted on common setal sup- ing contributions to the taxonomy of Australasian port plate; 8-P short, normally branched; 1-M,T not on mosquitoes. The generic name is feminine, formed tubercle, 1-M moderately long, 1-T short; 6-T single. from her nickname Pat, her surname and the femi- Abdomen: Seta 7-I long, stout, normally single; 12-I nine Latin suffix -ia. Recommended abbreviation = present; 7-I1 short, branched; 7,10,12-Vu single; 2,4- Pm. VIII single; segment VIII with comb normally with numerous scales in patch (some species with several KITCHING, scales in 1 or 2 irregular rows); segment X with saddle RAMPAMYJA REINERT, HARBACH & incomplete ventrally, moderately developed spicules GEN. NOV. on posterior margin dorsal to seta 1-X (occasionally Type species: Culex notoscriptus Skuse, 1889. 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 Females Pm. dobodura), and inserted on saddle. Head: Vertex with decumbent scales both broad and Siphon: With attached acus; pecten with spines narrow; occiput and vertex with erect forked scales; evenly spaced; seta 1-S branched, inserted distal to eyes contiguous or separated by less than or equal to 1 pecten. eye facet; antenna] pedicel with patch of overlapping, broad, silvery scales on mesal surface; clypeus bare; maxillary palpus with apex pale-scaled; proboscis Included species with well-developed pale-scaled band at about Patmarksia anggiensis (Bonne-Wepster), Pm. argen- midlength. teitarsis (Brug), Pm. argyronotum (Belkin), Pm. bux- Thorax: Acrostichal area with narrow stripe of pale toni (Belkin), Pm. clintoni (Taylor), Pm. derooki scales on anterior and posterior areas; posterior dor- (Brug), Pin. dohodura (King & Hoogstraal), Pm. hol- socentral area with narrow stripe of pale scales; scutal landia (King & Hoogstraal), Pm. mackerrasi (Taylor), fossa with narrow stripe of pale scales on lateral and Pm. novalbitarsis (King & Hoogstraal), Pm. pal- usually posterior margins; supraalar area with nar- marum (Edwards), Pm. papuensis (Taylor) and Pm. row, longitudinal patch of pale scales; prescutellar subalbitarsis (King & Hoogstraal). median, bare area with pale scales laterally mesal to setae; acrostichal and dorsocentral areas with both anterior and posterior setae; scutellum with broad Distribution scales on all lobes; scutal fossa with 1 or 2 posterior Australia (Queensland), Indonesia (Irian Jaya, setae; paratergite with broad, pale scales; broad scales Moluccas), Papua New Guinea and Solomon Islands on antepronotum, postpronotum, subspiracular area, (Guadalcanal, New Georgia). upper proepisternum, lower prealar area, upper and lower posterior mesokatepisternum, and upper and lower mesepimeron, other pleural areas without Bionomics scales; mesepimeron without lower setae. Immature stages of Patmarksia species are found in plant containers (e.g. fallen palm fronds, coconut Wing: Dark-scaled; upper calypter with several setae husks and leaves, treeholes, holes in fallen logs, cut on margin; alula with row of narrow scales on margin; bamboo, funnel-shaped fungi), artificial containers remigium without dorsal and ventral setae. (e.g. metal cans, drum heads, flower bowls), and small Legs: Ante- and postprocoxal membranes bare; mid- puddles, ruts, borrow pits and rock pools in stream and hindfemora with median stripe of pale scales on beds. anterior surface; hindtarsus with wide, basal, pale- scaled bands on tarsomeres 1-4; fore- and midungues equal, each with 1 tooth; hindungues equal, simple. Discussion See King & Hoogstraal (1946: pls 20, 21, 23) for illus- Abdomen: Tergum I with silvery-scaled patch on lat- trations of the male genitalia, fourth-instar larva and eral tergite; segment VII laterally compressed. adults of Pm. papuensis, Penn (1949: fig. 35) for the Genitalia: Tergum VIII sclerotized, width greater than pupa and Reinert (2002e: fig. 9) for the female length, without (rarely with 1 or 2 adventitious) genitalia. scales, setae on distal 0.30 or less of tergum; sternum

VIII scierotized, width greater than length, without Paddle: Apex broadly rounded; without fringe of hair- (rarely with 1 or 2 adventitious) scales, apex broad, like spicules; seta 1-Pa short, single. 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, Fourth-instar larvae median, apical emargination, 2-7 setae distally on Head: each side of midline; postgenital lobe relatively long, Seta 1-C stout, single, sharply curved, spini- narrow, with spicules well developed on lateral mar- form; 4-C short, with very slender branches, inserted gins, few setae distally; upper vaginal scierite well slightly mesal and slightly posterior to 6-C; 4,6-C dis- developed, moderately to heavily pigmented; lower placed far anterior; 5-C inserted posterior and slightly vaginal sclerite absent; insula liplike, with 5-7 setae lateral to 4,6-C and mesal and slightly anterior to 7-C; in lateral patches; cercus moderately long, moderately 7-C moderately long, with few branches; 5,6,13-C sin- wide, apex broadly rounded, without scales; 3 sper- gle; 19-C present; antenna without spicules, 1-A short, mathecal capsules. single (rarely 2-branched), inserted on distal third of antennal shaft. Thorax: Setae 1-3-P not on common setal support Males plate; 8-P short, multiple-branched; 1-M,T slender, branched, not on tubercle; 5-M very long, single, Head: Maxillary palpus with 5 palpomeres, longer noticeably longer than 7-M; 6-T single. than proboscis, numerous long setae ventrolaterally on distal part of palpomere 3 and ventrolaterally on Abdomen: Seta 7-I long, single; 12-I present; 741 palpomere 4; antennal flagellar whorls with numerous short, branched; 1,3,7,10,12-Vu single; 2,4-VIII sin- long setae directed dorsally and ventrally. gle; segment VIII with comb comprising numerous scales in patch; segment X with saddle incomplete Legs: Fore- and midtarsi with 2 unequal ungues, both ventrally, with well-developed spicules on posterior toothed; hindtarsus with 2 equal ungues, simple. margin dorsal to seta 1-X, ventral brush with several Genitalia: Posterior margin of tergum IX with pair of long, branched setae with relatively long stems, short, narrow, acute lobes bearing few slender to mod- attached to grid with both transverse and lateral bars, erately stout setae; gonocoxite relatively long, rela- precratal setae absent, 1-X multiple-branched, tively narrow, mesal margin of dorsal surface without inserted on saddle. apical and basal lobes, ventral surface with patch of Siphon: Relatively short; pecten with spines evenly long, narrow, lanceolate setae on distal part of mesal spaced; seta 1-S branched, inserted distal to pecten. margin, mesal surface membranous; gonostylus attached at apex of gonocoxite, relatively short, less than or equal to 0.25 length of gonocoxite, narrow, dis- Included species tal part slender and slightly curved mesally, with sin- Rampamyia albiabris (Edwards), Ra. notoscripta gle long, narrow gonostylar claw at apex; aedeagus notoscripta (Skuse), Ra. notoscripta montana (Brug) simple, tubelike, noticeably long and narrow; and Ra. quinquelineata (Edwards). paramere relatively long and narrow; proctiger with cercal setae; claspette with single, long, slender, columnar stem, claspette filament attached apically, Distribution flattened, curved with distal part tapering to bluntly Australia, Bismarck Archipelago (New Britain, New pointed apex; sternum IX with setae on posterome- Ireland), Indonesia (Java, Moluccas), New Caledonia dian area. (Ile des Pins, Loyalty Islands), New Zealand (North Island, South Island), Papua New Guinea and Solomon Islands (Bellona, Bougainville, Florida, Pupae Guadalcanal, Kolombangara, Malaita, New Georgia, Cephalothorax: Seta 11-CT single. Rendova, Rennell, Roviana, Sterling, Ugi). Abdomen: Setae 3,6,7-I single; 6-I longer than 74; 3-I1 long, single, inserted mesal to 2-I1; 541 long, single; Bionomics 5-IV,V stout, aciculate, each longer than the following Immature stages occur in plant containers, including tergum; 2-VI inserted mesal to 1-VI; 6-VII inserted treeholes, bamboo, plant leaf axils, palm fronds, coco- posterior and mesal to 9-Vu, noticeably shorter and nut shells and husks, and various small and large arti- thinner than 9-Vu; 9-VIII with multiple, aciculate ficial containers made of metal, cement and wood, and branches. rockholes.

Discussion Legs: Ante- and postprocoxal membranes bare; hind- See Belkin (1962: figs 226-229) for illustrations of the femur without pale scales dorsoapically; hindtibia male genitalia, pupa and fourth-instar larva of Ra. with broad, basal, white-scaled patch; hindtarsus with notoscripta, Russell (1996: 73) for photographs of the white-scaled, basal bands on tarsomeres 1 and 2, nar- female and Linley et al. (1991a) for a description and row on tarsomere 1, broad on tarsomere 2; fore-, mid- photographs of the egg. and hindungues equal, each with 1 tooth. Abdomen: Tergum I with silvery-scaled patch on lateral tergite; segment VII laterally compressed. Etymology Rampamyia is named in honour of Dr Rampa Rat- Genitalia: Tergum VIII with numerous broad scales tanarithikul in recognition of her outstanding contri- on much of surface; sternum VIII sclerotized, width butions to the taxonomy of Oriental mosquitoes. The greater than length, apical margin nearly flat or with generic name is feminine, formed from her first name very shallow, broad, median emargination, with sev- and the feminine Greek noun -myia, meaning fly. eral stout setae decreasing in length mesally, few to Recommended abbreviation = Ra. 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 VANSOMERENIS REINERT, HARBACH & KITCHING, 0.70 by moderately pigmented band, 2 or 3 short setae GEN. NOV. apically on each side of midline; postgenital lobe long, Type species: Aedes (Finlaya) pulchrithorax Edwards, relatively narrow, with several setae distally; upper 1939. and lower vaginal sclerites absent; insula liplike, with few setae in lateral patches; cercus moderately long, relatively narrow, apex sharply rounded, without Females scales; 3 spermathecal capsules. Head: Vertex with median, pale-scaled patch; vertex and occiput with erect forked scales; ocular line nar- Males row, with broad, silvery scales; eyes contiguous; clypeus bare; maxillary palpus and proboscis dark- Head: Maxillary palpus with 5 palpomeres, all rela- scaled. tively slender and in approximately straight line, dark-scaled, slightly longer to slightly shorter than Thorax: Scutum with narrow, curved scales; pale proboscis, palpomeres 4 and 5 with few setae; anten- scales forming relatively narrow stripe on anterior nal flagellar whorls with numerous long setae directed and posterior acrostichal areas and extending to dorsally and ventrally. prescutellar area, similar stripe on posterior dorsocentral area extending caudally laterad of pres- Legs: Fore-, mid- and hindtarsi each with 2 ungues, cutellar setae to posterior margin of scutum, each with 1 tooth. elongate, longitudinal patch on supraalar area, prescutellar area with narrow patches on lateral Genitalia: Tergum VIII with small, median lobe on and anterior parts of median, bare area; scutal posterior margin; tergum IX comprising 2 very small fossa covered with narrow, curved, dark scales; dor- sclerites connected mesally by tonguelike, median, socentral area with anterior and posterior setae; cephalad-directed lobe of tergum X, each sclerite with acrostichal area without setae; scutellum with nar- small, strongly sclerotized lobe on posterior margin row, curved scales on all lobes; mesopostnotum bearing few short, stout setae, lateral part of lobe con- bare; paratergite with broad, silvery scales; broad, nected by narrow band with sternum IX; gonocoxite silvery scales on antepronotum, postpronotum moderately long, moderately wide, dorsal surface with (large patch), upper proepisternum, mesokatepister- scales all approximately same length, mesal margin num (small upper and lower posterior patches), without apical lobe, with moderate-size, basal lobe, lower prealar area and mesepimeron (single large distal part of lobe narrow and bearing 2 or 3 moder- patch extending over upper and lower areas); sub- ately long to long, moderately stout setae whereas spiracular and postspiracular areas without scales; proximal part of lobe is expanded with series of wrin- upper proepisternum with 3-5 setae; mesepimeron kles at base and bearing several short to moderately without lower setae. long setae (some of which may be narrowly lanceolate), ventral surface without specialized scales Wing: Dark-scaled; upper calypter with several setae or setae, mesa] surface membranous; gonostylus on margin; alula with row of narrow scales on margin; attached at apex of gonocoxite, relatively short, slen- remigium with few dorsal setae distally. der, single gonostylar claw moderately long, narrow,

Journal compilation © 2006 The Linnean Society of London, Zoological Journal of the Linnean Society, 2006, 148, 1-101 No claim to original US government works. 86 J. F. REINERT ET AL. attached at apex; aedeagus simple, tubelike, moder- branches, differently developed than 7-I; segment VIII ately long; claspette with single moderately long, slen- with comb comprising numerous scales in patch; 2- der, columnar stem, claspette filament long, proximal VIII branched; segment X with saddle incomplete ven- part narrow, distal part expanded, flattened but ter- trally, acus absent, setae 1-X short, branched, inserted minating in pointed apex; proctiger with few cercal on saddle, ventral brush with several long, branched setae; sternum IX with few setae on posteromedian setae with short stems, attached to grid with well area. developed transverse grid bars, lateral bars absent but represented by faint depression, 2 precratal setae.

Pupae Siphon: With acus attached; pecten with number of evenly spaced spines; seta 1-S branched, inserted dis- Cephalothorax: Setae 1,3-CT branched; 7-CT very tal to pecten. 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, Included species with 2-6 branches; 10-CT relatively short, very slen- Vansomerenis hancocki (van Someren), Va. luteostri- der, with 6-19 branches; 11-CT long, stout, single. ata (Robinson) and Va. pulchrithorax (Edwards). Abdomen: Seta 3-I long, with 2 or 3 branches; 6-I very long, stout, aciculate, single, noticeably longer than Distribution 7-I; 7-I long, stout, aciculate, normally with 2 branches (rarely single or 3-branched); 241 shorter and inserted Angola, Kenya, Tanzania, Uganda, Zaire and Zambia. lateral to 141 and mesal to 3-1I; 3-11,111 long, stout, single; 541 shorter than 3-11; 641 long, stout, acicu- Bionomics late, single, longer than 7-I1; 741 long, branched; 5-IV—VI long, stout, aciculate, longer than following Immature stages have been found in treeholes, a rot tergum; 2-VT inserted mesal to 1,3-Vi; 6-VII short, hole in the fork of a tree, a tree buttress and bored slender, with 4-10 branches, inserted posterior and bamboo. mesal to 9-Vu; 9-VII stout, aciculate, branched, longer than 6-Vu; 9-VIII stout, aciculate, multiple-branched, Discussion inserted near posterolateral corner of segment VIII. See Edwards (1941: figs 6m, 176 and p1. 111-14) for Paddle: Length greater than width, index 1.20-1.42; illustrations of the female genitalia, fourth-instar without hairlike spicules on margins; midrib reaching larva and female of Va. pulchrithorax. Hopkins (1952: apex of paddle; seta 1-Pa single. figs 62, 63) also illustrated the fourth-instar larva of this species.

Fourth-instar larvae Etymology Head: Seta 1-C single, spiniform; 4-C short, very slen- Vansomerenis der, branched, inserted mesal and slightly posterior to is named in honour of Mrs E. C. C. van 6-C; 5-C long, stout, aciculate, often single (range sin- Someren in recognition of her many important contributions to the taxonomy of Afrotropical mosquitoes, gle to 3-branched), inserted posterior and mesal to 6-C and posterior and lateral to 4-C; 6-C long, stout, especially those in tribe Aedini. The generic name is feminine, formed from her surname and the feminine aciculate, longer than 5-C, inserted mesal and very Greek suffix -is denoting daughter of, i.e. daughter of slightly anterior to 7-C; 7-C moderately long, rela- Va. tively stout, aciculate, with 3-6 branches; 12-C van Someren. Recommended abbreviation = branched, inserted mesal to and shorter than 13-C; 13-C branched; 19-C absent; antenna relatively long, APPENDIX 4 slender, slightly curved mesally, with few scattered spicules, seta 1-A moderately long, single or 2- CHECKLIST AND CONSPECTUS OF RECLASSIFICATION branched (occasionally 3-branched). OF AEDINE TAXA Finlaya and certain allied taxa were the focus of the Thorax: Seta 5-P branched, shorter than 6-P; 8-P present study, which resulted in the recognition of 17 short; 4-M branched; 6-T single. genera, of which 11 are new to science, three elevated Abdomen: Setae 6-1,11 long, branched (6-I1 rarely sin- from subgeneric status and three restored from syn- gle in Va. luteostriata); 6-I11 branched, shorter than 6- onymy. As in our previous study (Reinert et al., 2004), II; 7-I long, with 2 or 3 branches; 741 short, with 4-15 some species of Finlaya and their allies cannot be

assigned to the new genera with confidence because of culicinus Edwards, 1922 a lack of specimens for examination and/or inadequate cumminsii (Theobald, 1903) published descriptions. Consequently, these species issp. mediopunctata (Theobald, 1909) (followed by an asterisk in the list below) are only pro- daizieli (Theobald, 1910) visionally placed in the newly proposed genera. As davidi Basio, 1971 previously, where it is not possible to assign species to dentatus (Theobald, 1904) a genus formally recognized herein or by Reinert et at. dialloi Hamon & Brengues, 1965 (2004) they are retained for the time being in the old domesticus (Theobald, 1901) broad concept of the genus-level taxon in which they durbanensis (Theobald, 1903) in were previously placed. For example, unassigned spe- ssp. angolae Ribeiro & Ramos, 1974 cies previously placed in Ochierotatus (Ochierotatus) ebogoensis Rickenbach & Ferrara, 1965 are listed under this genus and subgenus of previous eritreae Lewis, 1942 authors (sensu auctorum). The following checklist and falabreguesi Hamon, 1957 conspectus includes all aedine species and subspecies fihicis Ingram & de Meillon, 1927 recognized as valid at the end of March 2005. Where fowleri (Charmoy, 1908) necessary, the terminations of species names have gandarai da Cunha Ramos, Capela & Ribeiro, 1995 been changed to agree in gender with generic names. gibbinsi Edwards, 1935 gouldi Reinert, 1972 Aedini sensu Belkin, 1962 grenieri Hamon, Service, Adam & Taufflieb, 1961 Abraedes Zavortink, 1970 grjebinei Hamon, Taufflieb & Maillot, 1957 papago (Zavortink, 1970) hirsutus (Theobald, 1901) ssp. adenensis Edwards, 1941 Aedes Meigen, 1818 holocinctus Edwards, 1941 cinereus Meigen, 1818 hopkinsi Edwards, 1936 dahuricus Danilov, 1987 insolens Edwards, 1936 esoensis Yamada, 1921 irritans (Theobald, 1901) geminus Peus, 1970 jamesi (Edwards, 1914) mubiensis Luh & Shih, 1958 karooensis Muspratt, 1961 rossicus Dolbeskin, Gorickaja & Mitrofanova, 1930 lamborni Edwards, 1923 sasai Tanaka, Mizusawa & Saugstad, 1975 leesoni Edwards, 1932 yarnadai Sasa, Kano & Takahasi, 1950 leptolabis Edwards, 1936 leucarthrius (Speiser, 1909) Aedes sensu auctorum lokojoensis Service, 1959 Subgenus Aedimorphus Theobald, 1903 sensu longiseta Edwards, 1936 auctorum lottei Hamon & Brengues, 1965 abnorrnalis (Theobald, 1909) lowisii (Theobald, 1910) ssp. kabwachensis Edwards, 1941 mansouri Qutubuddin, 1959 adami Geoffroy, 1971 masoalensis Fontenille & Brunhes, 1985 aerarius McIntosh, 1975 mathioti Fontenille & Brunhes, 1985 aihocephalus (Theobald, 1903) mattinglyi Hamon & Rickenbach, 1954 albodorsalis Fontenille & Brunhes, 1985 mediolineatus (Theobald, 1901) alboscutellatus (Theobald, 1905) microstictus Edwards, 1936 aihoventralis (Theobald, 1910) minutus (Theobald, 1901) apicoannulatus (Edwards, 1912) mixtus Edwards, 1936 argenteopunctatus (Theobald, 1901) mutilus Edwards, 1936 argenteoscutellatus Carter & Wijesundara, 1948 natronius Edwards, 1932 hambiotai Geoffroy, 1987 nigricephalus (Theobald, 1901) bancoi Geoffroy, 1987 nigrostriatus (Barraud, 1927) bedfordi Edwards, 1936 nyounae Hamon & Adam, 1959 bevisi (Edwards, 1915) oakleyi Stone, 1939 boneti Gil Collado, 1936 ochraceus (Theobald, 1901) ssp. kumhae Chwatt, 1948 orbitae Edwards, 1922 caecus (Theobald, 1901) ovazzai Hamon & Adam, 1959 caliginosus (Graham, 1910) pachyurus Edwards, 1936 centropunctatus (Theobald, 1913) pallidostriatus (Theobald, 1907) chamboni Cornet, 1968 pampangensis (Ludlow, 1905)

phyllolabis Edwards, 1929 ngong (van Someren, 1950) pipersalatus (Giles, 1902) stokesi (Evans, 1929) pseudotarsalis van Someren, 1946 teesdalei (van Someren, 1954) pubescens Edwards, 1925 punctifemoris (Ludlow, 1921) Armigeres Theobald, 1901 punctothoracis (Theobald, 1909) Subgenus Armigeres quasiunivittatus (Theobald, 1901) alkatirii Toma, Miyagi & Syafruddin, 1995 reali Hamon & Adam, 1959 apoensis Bohart & Farner, 1944 rickenbachi Hamon & Adam, 1959 aureolineatus (Leicester, 1908) semlikiensis van Someren, 1950 azurini Basio, 1971 senyavinensis Knight & Hurlbut, 1949 baisasi Stone & Thurman, 1958 seychellensis (Theobald, 1912) bhayungi Thurman & Thurman, 1958 simulans (Newstead & Carter, 1911) breinli (Taylor, 1914) smithhurni van Someren, 1950 candelabrifer Brug, 1939 stenoetrus (Theobald, 1907) confusus Edwards, 1915 stenoscutus Edwards, 1912 conjungens Edwards, 1914 subdentatus Edwards, 1936 denbesteni Brug, 1925 syntheticus Barraud, 1928 durhami (Edwards, 1917) taeniorhynchoides (Christophers, 1911) ejercitoi Baisas, 1935 tarsalis (Newstead, 1907) fimbriatus Edwards, 1930 tauffiebi Rickenbach & Ferrara, 1965 foliatus Brug, 1931 tricholabis Edwards, 1941 giveni Edwards, 1926 ssp. bwamba van Someren, 1950 hybridus Edwards, 1914 trimaculatus (Theobald, 1905) joloensis (Ludlow, 1904) trukensis Bohart, 1957 jugraensis (Leicester, 1908) veeniae McIntosh, 1975 kesseli Ramalingam, 1987 vexans (Meigen, 1830) kinabaluensis Ramalingam, 1972 ssp. arabiensis (Patton, 1905) kuchingensis Edwards, 1915 ssp. nipponii (Theobald, 1907) lacuum Edwards, 1922 wendyae Service, 1959 laoensis Toma & Miyagi, 2003 wigglesworthi Edwards, 1941 maiae (Edwards, 1917) yangambiensis de Meillon & Lavoipierre, 1944 malayi (Theobald, 1901) yvonneae Edwards, 1941 manalangi Baisas, 1935 Subgenus Cancraedes Edwards, 1929 sensu auctorum maximus Edwards, 1922 cancricomes Edwards, 1922 milnensis Lee, 1944 curtipes Edwards, 1915 moultoni Edwards, 1914 indonesiae Mattingly, 1958 obturbans (Walker, 1859) kohkutensis Mattingly, 1958 pallithorax Dong, Zhou & Dong, 2004 mamoedjoensis Mattingly, 1958 papuensis Peters, 1963 masculinus Mattingly, 1958 setifer Delfinado, 1966 palawanicus Mattingly, 1958 seticoxitus Luh & Li, 1981 penghuensis Lien, 1968 suhalbatus (Coquillett, 1898) simplex (Theobald, 1903) var. chrysocorporis Hsieh & Liao, 1956 thurmanae Mattingly, 1958 theobaldi Barraud, 1934 yunnanensis Dong, Zhou & Dong, 1995 Alanstonea Mattingly, 1960 Subgenus Leicesteria Theobald, 1904 brevitibia (Edwards, 1914) annulipalpis (Theobald, 1910) treubi (de Meijere, 1910) annulitarsis (Leicester, 1908) balteatus Macdonald, 1960 Albuginosus Reinert, 1987 cingulatus (Leicester, 1908) capensis (Edwards, 1924) dentatus Barraud, 1927 gilliesi (van Someren, 1962) digitatus (Edwards, 1914) haworthi (Edwards, 1923) dolichocephalus (Leicester, 1908) kapretwae (Edwards, 1941) flavus (Leicester, 1908) kennethi (Muspratt, 1956) inchoatus Barraud, 1927 marshallii (Theobald, 1901) lepidocoxitus Dong, Zhou & Dong, 1995

longipalpis (Leicester, 1904) ningheensis (Lei, 1989), comb. nov. magnus (Theobald, 1908) pseudotaeniatus (Giles, 1901), comb. nov. menglaensis Dong, Zhou & Dong, 2002 shortti (Barraud, 1923), comb. nov. omissus (Edwards, 1914) tonkinensis (Galliard & Ngu, 1947), comb. nov. pectinatus (Edwards, 1914) pendulus Edwards, 1914 Dahliana Reinert, Harbach & Kitching, gen. traubi Macdonald, 1960 nov. vimoli Thurman & Thurman, 1958 echinus (Edwards, 1920), comb. nov. Nomina duhia geniculata (Olivier, 1791), comb. nov. striocrurus (Giles, 1904) gilcolladoi (Sanchez-Covisa Villa, Rodriguez ventralis (Walker, 1860) Rodriguez & Guillen Llera, 1985), comb. nov.

Ayurakitia Thurman, 1954 Diceromyia Theobald, 1911 griffithi Thurman, 1954 adersi (Edwards, 1917) peytoni (Reinert, 1972) agastyai (Tewari & Hiriyan, 1992) bananea (Wolfs, 1958) Aztecaedes Zavortink, 1972 cordellieri (Huang, 1986) ramirezi (Vargas & Downs, 1950) coulangesi (Rodham & Boutonnier, 1983) fascipalpis (Edwards, 1912) Belkinius Reinert, 1982 flavicollis (Edwards, 1928) aurotaeniatus (Edwards, 1922) franciscoi (Mattingly, 1959) furcifer (Edwards, 1913) Bothaella Reinert, 1973 grassei (Doucet, 1951) alongi (Galliard & Ngu, 1947), comb. nov. iyengari (Edwards, 1923) brownscutuma (Dong, Zhu & Dong, 1999) kanarensis (Edwards, 1934) eldridgei (Reinert, 1973) madagascarensis (van Someren, 1949) helenae (Reinert, 1973) mefouensis (Ferrara, 1974) kleini (Reinert, 1973) meronephada (Dyar & Shannon, 1925) micropterus (Giles, 1901) Bruceharrisonius Reinert, 2003, stat. nov., raised nummata (Edwards, 1923) to genus rank periskelata (Giles, 1902) alektorovi (Stackelberg, 1943), comb. nov. pseudonummata (Reinert, 1973) aureostriatus (Doleschall, 1857), comb. nov. punctipes (Edwards, 1921) christophersi (Edwards, 1922), comb. nov. ramachandrai (Reuben, 1967) doonii (Wattal, Bhatia & Kalra, 1958), comb. reginae (Edwards, 1922) nov. reubenae (Tewari & Hiriyan, 1992) greenii (Theobald, 1903), comb. nov. scanloni (Reinert, 1970) huributi (Lien, 1967), comb. nov. sylvatica (Brunhes, 1983) okinawanus (Bohart, 1946), comb. nov. taylori (Edwards, 1936) taiwanus (Lien, 1968), comb. nov. tiptoni (Grjebine, 1953) whartoni (Mattingly, 1965) Christophersiomyia Barraud, 1923 zethus (de Meillon & Lavoipierre, 1944) annulirostris (Theobald, 1905) chionodes (Belkin, 1962) Dobrotworskyius Reinert, Harbach & Kitching, gombakensis (Mattingly, 1959) gen. nov. ibis (Barraud, 1931) alboannulatus (Macquart, 1850), comb. nov. thomsoni (Theobald, 1905) milsoni (Taylor, 1915), comb. nov. occidentalis (Skuse, 1889), comb. nov. Collessius Reinert, Harbach & Kitching, gen. rubrithorax (Macquart, 1850), comb. nov. nov. rupestris (Dobrotworsky, 1959), comb. nov. banksi (Edwards, 1922), comb. nov. subbasalis (Dobrotworsky, 1962), comb. nov. elsiae (Barraud, 1923), comb. nov. tubbutiensis (Dobrotworsky, 1959), comb. nov. ssp. vicarius (Lien, 1968), comb. nov. hatorii (Yamada, 1921), comb. nov. Downsiomyia Vargas, 1950 macdougalli (Edwards, 1922), comb. nov. albolateralis (Theobald, 1908) macfarlanei (Edwards, 1914), comb. nov. albonivea (Barraud, 1934)

axitiosa (Kulasekera, Knight & Harbach, 1990) inornatus Newstead, 1907 dorseyi (Knight, 1946) intermedius Edwards, 1936 ganapathi (Colless, 1958) jani Rickenbach & Lombrici, 1976 harinasutai (Knight, 1978) lacani Rickenbach & Eouzan, 1970 idjenensis (Brug, 1934) leucopous Graham, 1909 inermis (Colless, 1958) mahaffyi van Someren, 1949 lactea (Knight, 1946) marcellei Adam & Hamon, 1959 laoagensis (Knight, 1946) mattinglyi Hamon & van Someren, 1961 leonis (Colless, 1958) melanopous Graham, 1909 PH litorea (Colless, 1958) mortiauxi Cunha Ramos & Ribeiro, 1990 mikrokopion (Knight & Harrison, 1988) oedipodeios Graham, 1909 mjobergi (Edwards, 1926), comb. nov. parvipluma Edwards, 1941 mohani (Knight, 1969) pauliani Grjebine, 1950 nipponica (LaCasse & Yamaguti, 1948) penicillatus Edwards, 1941 nipoononivea (Sasa & Nakahashi, 1952) plioleucus Edwards, 1941 nishikawai (Tanaka, Mizusawa & Saugstad, 1979) ssp. brevis Edwards, 1941 nivea (Ludlow, 1903) productus Edwards, 1941 niveoides (Barraud, 1934) quinquevittatus Theobald, 1901 novonivea (Barraud, 1934) ravissei Rickenbach & Eouzan, 1970 omorii (Lien, 1968) rickenbachi Ferrara & Eouzan, 1974 pexa (Colless, 1958) salauni Rickenbach, Ferrara & Eouzan, 1968 pseudonivea (Theobald, 1905) semisimplicipes Edwards, 1914 saperoi (Knight, 1946) silvestris Ingram & de Meillon, 1927 shehzadae (Qutubuddin, 1972), comb. nov. ssp. conchobius Edwards, 1941 sinensis (Chow, 1950) subsimplicipes Edwards, 1914 subnivea (Edwards, 1922) tendeiroi da Cunha Ramos, Ribeiro & Machado, vana (Colless, 1958) 1992 watteni (Lien, 1968) tonsus Edwards, 1941 vansomereni Hamon, 1962 Edwardsaedes Belkin, 1962 wansoni Edwards, 1941 bekkui (Mogi, 1977) ssp. douceti Adam & Hamon, 1959 imprimens (Walker, 1860) pingpaensis (Chang, 1965) Finlaya Theobald, 1903 alocasicola (Marks, 1947) Eretmapodites Theobald, 1901 ananae (Knight & Laffoon, 1946) adami Ferrara & Eouzan, 1974 avistylus (Brug, 1939) angolensis da Cunha Ramos & Ribeiro, 1992 bougainvillensis (Marks, 1947) argyrurus Edwards, 1936 burnetti (Belkin, 1962) brenguesi Rickenbach & Lombrici, 1975 crocea (Knight & Laffoon, 1946) brottesi Rickenbach, 1967 dobrotworskyi (Marks, 1958) caillardi Rickenbach, Ferrara & Eouzan, 1968 fijiensis (Marks, 1947) chrysogaster Graham, 1909 flavipennis Giles, 1904 corbeti Hamon, 1962 franclemonti (Belkin, 1962) dracaenae Edwards, 1916 freycinetiae (Laird, 1957) dundo da Cunha Ramos & Ribeiro, 1992 fuscipalpis (Belkin, 1962) eouzani Rickenbach & Lombrici, 1974 fuscitarsis (Belkin, 1962) ferrarai Rickenbach & Eouzan, 1970 gahnicola (Marks, 1947) forcipulatus Edwards, 1936 gani (Bonne-Wepster, 1940) germaini Rickenbach & Eouzan, 1970 gressitti (Bohart, 1947) gilletti van Someren, 1949 hollingsheadi (Belkin, 1962) grahami Edwards, 1911 horotoi (Taylor, 1972) grenieri Hamon & van Someren, 1961 hui (Bohart, 1957) haddowi van Someren, 1949 josephinae (Marks, 1958) hamoni Grjebine, 1972 knighti (Stone & Bohart, 1944) harperi van Someren, 1949 kochi (Donitz, 1901) hightoni van Someren, 1947 lewelleni (Starkey & Webb, 1946)

lutea (Ludlow, 1905) anastasionis Dyar, 1921 maffli (Taylor & Tenorio, 1974) andinus Osorno-Mesa, 1944 medleri (Knight & Laffoon, 1946) argyromeris Dyar & Ludlow, 1921 neogeorgiana (Belkin, 1962) baresi Cerqueira, 1960 oceanica (Belkin, 1962) hoshelli Osorno-Mesa, 1944 poicilia Theobald, 1903 capricornii Lutz, 1904 samoana Grunberg, 1913 celeste Dyar & Nunez Tovar, 1927 schiosseri (Belkin, 1962) chalcospilans Dyar, 1921 solomonis (Stone & Bohart, 1944) chrysochiorus Arnell, 1973 sorsogonensis (Banez & Jueco, 1966) equinus Theobald, 1903 stonei (Knight & Laffoon, 1946) iridicolor Dyar, 1921 tutuilae (Ramalingam & Belkin, 1965) janthinomys Dyar, 1921 wallacei (Edwards, 1926) lucifer (Howard, Dyar & Knab, 1913) mesodentatus Komp & Kumm, 1938 Fredwardsius Reinert, 2000 nebulosus Arnell, 1973 vittatus (Bigot, 1861) panarchys Dyar, 1921 regalis Dyar & Knab, 1906 Georgecraigius Reinert, Harbach & Kitching, soperi Levi-Castillo, 1955 gen. nov. spegazzinii Brethes, 1912 Subgenus Georgecraigius splendens Williston, 1896 atropalpus (Coquillett, 1902), comb. nov. tropicalis Cerqueira & Antunes, 1938 epactius (Dyar & Knab, 1908), comb. nov. Subgenus Hors fallius Reinert, Harbach & Halaedes Belkin, 1962 Kitching, subgen. nov. ashworthi (Edwards, 1921) fluviatilis (Lutz, 1904), comb. nov. australis (Erichson, 1842) wardangensis (Brust, Ballard, Driver, Hartley, Geoskusea Edwards, 1929 Galway & Curran, 1998) baisasi (Knight & Hull, 1951) becki (Belkin, 1962) daggyi (Stone & Bohart, 1944) Heizmannia Ludlow, 1905 finibripes (Edwards, 1924) Subgenus Heizmannia kabaenensis (Brug, 1939) aurea Brug, 1932 longiforceps (Edwards, 1929) aureochaeta (Leicester, 1908) lunulata (King & Hoogstraal, 1946) carteri Amerasinghe, 1993 perryi (Belkin, 1962) chandi Edwards, 1922 timorensis (Miyagi, Toma & Lien, 2004), comb. chengi Lien, 1968 nov. communis (Leicester, 1908) tonsa (Edwards, 1924) complex (Theobald, 1910) covelli Barraud, 1929 Gilesius Reinert, Harbach & Kitching, gen. nov. demeilloni Mattingly, 1970 alius (Lien, 1968), comb. nov. funerea (Leicester, 1908) pulchri venter (Giles, 1901), comb. nov. greenii (Theobald, 1905) heterospina Gong & Lu, 1986 Gymnometopa Coquillett, 1905 himalayensis Edwards, 1922 mediovittata (Coquillett, 1906) indica (Theobald, 1905) kana Tanaka, Mizusawa & Saugstad, 1979 Haemagogus Williston, 1896 kanhsienensis Tung, 1955 Subgenus Conopostegus Dyar, 1925 Iii Wu, 1936 clarki (Galindo, Carpenter & Trapido, 1953) macdonaldi Mattingly, 1957 leucocelaenus (Dyar & Shannon, 1924) inattinglyi Thurman, 1959 leucophoebus (Galindo, Carpenter & Trapido, 1953) menglianensis Lu & Gong, 1986 leucotaeniatus (Komp, 1938) menglianeroides Dong, Dong & Zhou, 2003 Subgenus Haemagogus persimilis Mattingly, 1970 acutisentis Arnell, 1973 propin qua Mattingly, 1970 aeritinctus Galindo & Trapido, 1967 proxima Mattingly, 1970 albomaculatus Theobald, 1903 reidi Mattingly, 1957

ruiliensis Dong, Zhou & Wang, 1997 Huaedes Huang, 1968 scanloni Mattingly, 1970 medialis (Brug, 1932) scintillans Ludlow, 1905 variepictus (King & Hoogstraal, 1946) taiwanensis Lien, 1968 wauensis (Huang, 1968) tengchongensis Dong, Wang & Zhou, 2002 thelmae Mattingly, 1970 Hulecoeteomyia Theobald, 1904, stat. nov., viridis Barraud, 1929 restored to genus rank Subgenus Mattinglyia Lien, 1968 chrysolineata (Theobald, 1907), comb. nov. achaetae (Leicester, 1908) formosensis (Yamada, 1921), comb. nov. No catesi (Lien, 1968) harveyi (Barraud, 1923), comb. nov. discrepans (Edwards, 1922) japonica (Theobald, 1901), comb. nov. occidentayunnana (Gong & Lu, 1991) ssp. shintienensis (Tsai & Lien, 1950), comb. tripunctata (Theobald, 1908) nov. ssp. amamiensis (Tanaka, Mizusawa & Saug. stad, 1979), comb. nov. Himalaius Reinert, Harbach & Kitching, gen. ssp. yaeyamensis (Tanaka, Mizusawa & Saug. nov. stad, 1979), comb. nov. gilli (Barraud, 1924), comb. nov. jugraensis Leicester, 1908, comb. nov. simlensis (Edwards, 1922) , comb. nov. koreica (Edwards, 1917), comb. nov. nigrorhynchus (Brug, 1931), comb. nov. pallirostris (Edwards, 1922), comb. nov. Howardina Theobald, 1903 reinerti (Rattanarithikul & Harrison, 1988), albonotata (Coquillett, 1906) comb. nov. allotecnon (Kumm, Komp & Ruiz, 1940) rizali (Banks, 1906), comb. nov. arborealis (Bonne-Wepster & Bonne, 1920) saxicola (Edwards, 1922), comb. nov. argyriti (Dyar & Nunez Tovar, 1927) sherki (Knight, 1948), comb. nov. aureolineata (Berlin, 1969) yunnanensis (Gaschen, 1934), comb. nov. auriti Theobald, 1907 aurivittata (Cerqueira, 1943) Indusius Edwards, 1934 bahamensis (Berlin, 1969) pulverulentus (Edwards, 1922) brevis (Berlin, 1969) brevivittata (Berlin, 1969) Isoaedes Reinert, 1979 busckii (Coquillett, 1906) cavaticus (Reinert, 1979) cozumelensis (Diaz Najera, 1966) ecuadoriensis (Berlin, 1969) Jarnellius Reinert, Harbach & Kitching, gen. eleanorae (Berlin, 1969) nov. fulvithorax (Lutz, 1904) Subgenus Jarnellius grabhami (Berlin, 1969) deserticola (Zavortink, 1969), comb. nov. guatemala (Berlin, 1969) laguna (Arnell & Nielsen, 1972), comb. nov. guerrero (Berlin, 1969) monticola (Belkin & McDonald, 1957), comb. inaequalis Grabham, 1907 nov. ioliota (Dyar & Knab, 1913) sierrensis (Ludlow, 1905), comb. nov. leei (Berlin, 1969) varipalpus (Coquillett, 1902), comb. nov. lorraineae (Berlin, 1969) Subgenus Lewnielsenius Reinert, Harbach & marinkellei (Berlin, 1969) Kitching, subgen. nov. martinezi (Berlin, 1969) muelleri (Dyar, 1920), comb. nov. osornoi (Berlin, 1969) pseudodminicii (Komp, 1936) Jihlienius Reinert, Harbach & Kitching, gen. quadrivittata (Coquillett, 1902) nov. septemstriata (Dyar & Knab, 1907) chungi (Lien, 1968), comb. nov. sexlineata (Theobald, 1901) gonguoensis (Gong & Lu, 1986), comb. nov. spinosa (Berlin, 1969) unicinctus (Edwards, 1922), comb. nov. stenei (Thompson, 1956) vanerndeni (Martini, 1931) Kenknightia Reinert, 1990 walkeri (Theobald, 1901) dissimilierodes (Dong, Zhou & Dong, 2002) whitmorei (Dunn, 1918) dissimilis (Leicester, 1908)

gaffigani (Reinert, 1990) ferinus (Knight, 1947) harbachi (Reinert, 1990) grahamii Theobald, 1909 karwari (Barraud, 1924) laniger (Wiedemann, 1820) lerozeboomi (Reinert, 1990) lucianus (Muspratt, 1959) leucomeres (Giles, 1904) mucidus (Karsch, 1887) litwakae (Reinert, 1990) nigerrimus Theobald, 1913 luzonensis (Rozeboom, 1946) painei (Knight, 1948) paradissimilis (Rozeboom, 1946) quadripunctis (Ludlow, 1910) pecori (Reinert, 1990) quasiferinus (Mattingly, 1961) wilkersoni (Reinert, 1990) scatophagoides Theobald, 1901 sudanensis Theobald, 1908 Kompia Aitken, 1941 tonkingi (Gebert, 1948) purpureipes (Aitken, 1941) Neoinelaniconion Newstead, 1907 Leptosomatomyia Theobald, 1905 albicosta (Edwards, 1913) aurimargo (Edwards, 1922) albothorax (Theobald, 1907) aurovenatum (Worth, 1960) Levua Stone & Bohart, 1944 hequaerti (Wolfs, 1947) dufouri (Hamon, 1953) bergerardi (Pajot & Geoffroy, 1971) fryeri (Theobald, 1912) bolense (Edwards, 1936) geoskusea (Amos, 1944) carteri (Edwards, 1936) circumluteoluin (Theobald, 1908) Lorrainea Belkin, 1962 crassiforceps (Edwards, 1927) amesii (Ludlow, 1903) ellinorae (Edwards, 1941) celebica (Mattingly, 1959) flavimargo (Edwards, 1941) dasyorrhus (King & Hoogstraal, 1946) fusci nerve (Edwards, 1914) fumida (Edwards, 1928) jamoti (Hamon & Rickenbach, 1954) lamellifera (Bohart & Ingram, 1946) lineatopenne (Ludlow, 1905) ssp. aureum (Gutsevich, 1955) Macleaya Theobald, 1903, stat. nov., restored to luridum (McIntosh, 1971) genus rank luteolaterale (Theobald, 1901) Subgenus Chaetocruiomyia Theobald, 1910, trans- mcintoshi (Huang, 1985) ferred from Ochierotatus monotrichus (Edwards, 1936) calabyi (Marks, 1963), comb. nov. palpale Newstead, 1907 elchoensis (Taylor, 1929), comb. nov. pogonurum (Edwards, 1936) humeralis (Edwards, 1922), comb. nov. punctocostale (Theobald, 1909) macmillani (Marks, 1964), comb. nov. taeniaroslre (Theobald, 1909) moloiensis (Taylor, 1929), comb. nov. unidentatum (McIntosh, 1971) spinosipes Edwards, 1922, comb. nov. tulliae (Taylor, 1929), comb. nov. Ochierotatus Lynch Arribaizaga, 1891 wattensis (Taylor, 1929), comb. nov. Subgenus Rusticoidus Shevchenko & Prudkina, 1973 Subgenus Macleaya, transferred from Ochlerot at us albescens (Edwards, 1921) littlechildi (Taylor, 1933), comb. nov. bicri status (Thurman & Winkler, 1950) stoneorum (Marks, 1977), comb. nov. krymmontanus (Alekseev, 1989) tremula Theobald, 1903, comb. nov. lepidonotus (Edwards, 1920) provocans (Walker, 1848) Molpemyia Theobald, 1910, stat. nov., restored to quasirusticus (Torres Canamares, 1951) genus rank refiki (Medschid, 1928) auridorsum (Edwards, 1922), comb. nov. rusticus (Rossi, 1790) pecuniosa (Edwards, 1922), comb. nov. var. suhtrichurus (Martini, 1927) purpurea Theobald, 1910, comb. nov. subdiversus (Martini, 1926) Unassigned to subgenus Mucidus Theobald, 1901 atlanticus (Dyar & Knab, 1906) alternans (Westwood, 1835) communis (de Geer, 1776) aurantius (Theobald, 1907) dorsalis (Meigen, 1830) ssp. chrysogaster Taylor, 1927 fulvus (Wiedemann, 1828)

ssp. pallens (Ross, 1943) albineus (Seguy, 1923) infirmatus (Dyar & Knab, 1906) aloponotum (Dyar, 1917) scapularis (Rondani, 1848) ambreensis (Rodham & Boutonnier, 1983) sollicitans (Walker, 1856) andersoni (Edwards, 1926) angustivittatus (Dyar & Knab, 1907) Ochierotatus sensu auctorum annulipes (Meigen, 1830) Subgenus Finlaya Theobald, 1903 sensu auctorum antipodeus Edwards, 1920 albocinctus (Barraud, 1924) atactavittatus (Arnell, 1976) alhotaeniata (Leicester, 1904) auratus (Grabham, 1906) ra alticola (Bonne-Wepster, 1948) aurifer (Coquiflett, 1903) auronitens (Edwards, 1922) behningi (Martini, 1926) australiensis (Theobald, 1910) bejaranoi (Martinez, Carcavallo & Prosen, 1960) barnardi (Edwards, 1924) berlandi (Seguy, 1921) biocellatus (Taylor, 1915) himaculatus (Coquillett, 1902) britteni (Marks & Hodgkin, 1958) biskraensis (Brunes, 1999) candidoscutellum (Marks, 1947) bogotanus (Arnell, 1976) crossi (Lien, 1967) breedensis (Muspratt, 1953) eatoni Edwards, 1916 burjaticus (Kuchartshuk, 1973) embuensis (Edwards, 1930) hurpengaryensis (Theobald, 1905) fengi (Edwards, 1935) caballus (Theobald, 1912) gracilelineatus (Bonne-Wepster, 1937) cacozelus (Marks, 1963) harperi (Knight, 1948) calcariae (Marks, 1957) ingrami (Edwards, 1930) calumnior (Belkin, Heinemann & Page, 1970) keefei (King & Hoogstraal, 1946) campestris (Dyar & Knab, 1907) koreicoides (Sasa, Kano & Hayashi, 1950) camptorhynchus (Thomson, 1869) lauriei Carter, 1920 canadensis (Theobald, 1901) lepchana (Barraud, 1923) ssp. mathesoni (Middlekauff, 1944) loi (Lien, 1968) cantans (Meigen, 1818) mallochi (Taylor, 1944) var.subvexans (Martini, 1922) monocellatus (Marks, 1948) cantator (Coquillett, 1903) nyasae (Edwards, 1930) caspius (Pallas, 1771) oreophilus Edwards, 1916 ssp. meirai Ribeiro, da Cunha Ramos, Capela & peipingensis (Feng, 1938) Pires, 1980 plagosus (Marks, 1959) var. hargreavesi Edwards, 1920 quasirubithorax (Theobald, 1910) cataphylla (Dyar, 1916) roai (Belkin, 1962) var. rostochiensis (Martini, 1920) rubiginosus (Belkin, 1962) chelli Edwards, 1915 seoulensis (Yamada, 1921) churchillensis (Ellis & Brust, 1973) sintoni (Barraud, 1924) clelandi (Taylor, 1914) stanleyi (Peters, 1963) clivis (Lanzaro & Eldridge, 1992) subauridorsum (Marks, 1948) coluzzii (Rioux, Guilvard & Pasteur, 1998) suffusus (Edwards, 1922) comitatus (Arnell, 1976) toxopeusi (Bonne-Wepster, 1948) condolescens (Dyar & Kriab, 1907) tsiliensis (King & Hoogstraal, 1946) continentalis (Dobrotworsky, 1960) versicolor (Barraud, 1924) crinifer (Theobald, 1903) wasselli (Marks, 1947) cunabulanus (Edwards, 1924) wellmanii (Theobald, 1905) cyprioides (Danilov & Stupin, 1982) cyprius (Ludlow, 1920) Subgenus Ochierotatus Lynch Arribalzaga, 1891 decticus (Howard, Dyar & Knab, 1917) sensu auctorum deficiens (Arnell, 1976) aboriginis (Dyar, 1917) detritus (Haliday, 1833) abserratus (Felt & Young, 1904) diantaeus (Howard, Dyar & Knab, 1913) aculeatus (Theobald, 1903) duplex (Martini, 1926) aenigmaticus (Cerqueira & Costa, 1946) dupreei (Coquillett, 1904) akkeshiensis (Tanaka, 1998) dzeta (Seguy, 1924) albifasciatus (Macquart, 1838) edgari (Stone & Rosen, 1952)

eidsvoldensis (Mackerras, 1927) nigrocanus (Martini, 1927) eucephalaeus (Dyar, 1918) nigromaculis (Ludlow, 1906) euedes (Howard, Dyar & Knab, 1913) niphadopsis (Dyar & Kiiab, 1918) euiris (Dyar, 1922) nivalis (Edwards, 1926) euplocamus (Dyar & Knab, 1906) normanensis (Taylor, 1915) excrucians (Walker, 1856) nubilis (Theobald, 1903) explorator (Marks, 1964) obturbator (Dyar & Knab, 1907) fitchii (Felt & Young, 1904) oligopistus (Dyar, 1918) flavescens (Muller, 1764) patersoni (Shannon & del Ponte, 1928) flavifrons (Skuse, 1889) pectinatus (Arnell, 1976) grossbecki (Dyar & Krab, 1906) pennai (Antunes & Lane, 1938) gutzevichi (Dubitsky & Deshevykh, 1978) perkinsi (Marks, 1949) hakusanensis (Yamaguti & Tamaboko, 1954) pertinax (Grabham, 1906) harrisoni (Muspratt, 1953) perventor (Cerqueira & Costa, 1946) hastatus (Dyar, 1922) phaecasiatus (Marks, 1964) hesperonotius (Marks, 1959) phaeonotus (Arnell, 1976) hexodontus (Dyar, 1916) phoeniciae (Coluzzi & Sabatini, 1968) hodgkini (Marks, 1959) pionips (Dyar, 1919) hokkaidensis (Tanaka, Mizusawa & Saugstad, 1979) procax Skuse, 1889 hortator (Dyar & Knab, 1907) pseudonormanensis (Marks, 1949) hungaricus (Mihalyi, 1955) puicritarsis (Rondani, 1872) imperfectus (Dobrotworsky, 1962) ssp. asiaticus (Edwards, 1926) impiger (Walker, 1848) pullatus (Coquillett, 1904) ssp. daisetsuzanus (Tanaka, Mizusawa & Saugstad, punctodes (Dyar, 1922) 1979) punctor (Kirby, 1837) implicatus (Vockeroth, 1954) purpuraceus (Brug, 1932) incomptus (Arnell, 1976) purpureifemur (Marks, 1959) increpitus (Dyar, 1916) purpuriventris (Edwards, 1926) inexpectatus (Bonne-Wepster, 1948) ratcliffei (Marks, 1959) intermedius (Danilov & Gornostaeva, 1987) raymondi (del Ponte, Castro & Garcia, 1951) intrudens (Dyar, 1919) rempeli (Vockeroth, 1954) jacobinae (Serafim & Davis, 1933) rhyacophilus (da Costa Lima, 1933) jorgi (Carpintero & Leguizamón, 2000) riparioides (Su & Zhang, 1987) juppi (McIntosh, 1973) riparius (Dyar & Knab, 1907) kasachstanicus (Gutsevich, 1962) sagax (Skuse, 1889) lasaensis (Meng, 1962) sapiens (Marks, 1964) ssp. gyirongensis (Ma, 1982) schizopinax (Dyar, 1929) lepidus (Cerqueira & Paraense, 1945) schtakelbergi (Shingarev, 1928) leucomelas (Meigen, 1804) scutellalbum (Boshell-Manrique, 1939) linesi (Marks, 1964) sedaensis (Lei, 1989) longifilamentus (Su & Zhang, 1988) sergievi (Danilov, Markovich & Proskuryakova, luteifemur (Edwards, 1926) 1978) macintoshi (Marks, 1959) serratus (Theobald, 1901) mariae (Sergent & Sergent, 1903) shannoni (Vargas & Downs, 1950) martineti (Senevet, 1937) silvestris (Dobrotworsky, 1961) mcdonaldi (Belkin, 1962) simanini (Gutsevich, 1966) FM melanimon (Dyar, 1924) sinkiangensis (Hsiao, 1977) meprai (Martinez & Prosen, 1953) spencerii (Theobald, 1901) mercurator (Dyar, 1920) ssp. idahoensis (Theobald, 1903) milleri (Dyar, 1922) spilotus (Marks, 1963) mitchellae (Dyar, 1905) squamiger (Coquillett, 1902) montchadskyi (Dubitsky, 1968) sticticus (Meigen, 1838) nevadensis (Chapman & Barr, 1964) stigmaticus (Edwards, 1922) nigrinus (Eckstein, 1918) stimulans (Walker, 1848) nigripes (Zetterstedt, 1838) stramineus (Dubitzky, 1970) nigrithorax (Macquart, 1847) stricklandi Edwards, 1912

subalbirostris (Klein & Marks, 1960) thorntoni (Dyar & Knab, 1907) surcoufi (Theobald, 1912) triseriatus (Say, 1823) synchytus (Arnell, 1976) vargasi (Schick, 1970) taeniorhynchus (Wiedemann, 1821) zavortinki (Schick, 1970) tahoensis (Dyar, 1916) zoosophus (Dyar & Knab, 1918) theicter (Dyar, 1918) Subgenus uncertain theobaldi (Taylor, 1914) daliensis (Taylor, 1916) thibaulti (Dyar & Knab, 1910) tormentor (Dyar & Knab, 1906) Opifex Hutton, 1902 tortilis (Theobald, 1903) Subgenus Nothoskusea Dumbleton, 1962 trivittatus (Coquillett, 1902) chathamicus (Dumbleton, 1962) turneri (Marks, 1963) Subgenus Opifex upatensis (Anduze & Hecht, 1943) fu.scus Hutton, 1902 ventrovittis (Dyar, 1916) vigilax (Skuse, 1889) Paraedes Edwards, 1934 ssp. ludlowae (R. Blanchard, 1905) harraudi Edwards, 1934 ssp. vansomerenae (Mattingly, 1955) honneae (Mattingly, 1958) vittiger (Skuse, 1889) chrysoscuta (Theobald, 1910) washinoi (Lanzaro & Eldridge, 1992) collessi (Mattingly, 1958) zammitii (Theobald, 1903) menoni (Mattingly, 1958) Subgenus Protomacleaya Theobald, 1907 sensu auc- ostentatio (Leicester, 1908) torum pagei (Ludlow, 1911) aitkeni (Schick, 1970) thailandensis (Reinert, 1976) alboapicus (Schick, 1970) amahilis (Schick, 1970) Patmarksia Reinert, Harbach & Kitching, gen. argyrothorax (Bonne-Wepster & Bonne, 1920) nov. herlini (Schick, 1970) anggiensis (Bonne-Wepster, 1937), comb. nov. bertrami (Schick, 1970) argenteitarsis (Brug, 1932), comb. nov. braziliensis (Gordon & Evans, 1922) argyronotum (Belkin, 1962), comb. nov. brelandi (Zavortink, 1972) buxtoni (Belkin, 1962), comb. nov. buenaventura (Schick, 1970) clintoni (Taylor, 1946), comb. nov. burgeri (Zavortink, 1972) derooki (Brug, 1932), comb. nov. campana (Schick, 1970) dobodura (King & Hoogstraal, 1946), comb. casali (Schick, 1970) nov. chionotum (Zavortink, 1972) hollandia (King & Hoogstraal, 1946), comb. daryi (Schick, 1970) nov. diazi (Schick, 1970) mackerrasi (Taylor, 1927), comb. nov. gabriel (Schick, 1970) novalbitarsis (King & Hoogstraal, 1946), comb. galindoi (Schick, 1970) nov. hendersoni (Cockerell, 1918) palmarum (Edwards, 1924), comb. nov. heteropus (Dyar, 1921) papuensis (Taylor, 1914), comb. nov. homoeopus (Dyar, 1922) subalbitarsis (King & Hoogstraal, 1946), comb. idanus (Schick, 1970) nov. impostor (Schick, 1970) insolitus (Coquillett, 1906) Phagomyia Theobald, 1905, stat. nov., restored to knahi (Coquillett, 1906) genus rank kompi (Vargas & Downs, 1950) assamensis (Theobald, 1908), comb. nov. metoecopus (Dyar, 1925) cacharana (Barraud, 1923), comb. nov. niveoscutum (Zavortink, 1972) cogilli (Edwards, 1922), comb. nov. podographicus (Dyar & Knab, 1906) deccana (Barraud, 1923), comb. nov. sandrae (Zavortink, 1972) feegradei (Barraud, 1934), comb. nov. schicki (Zavortink, 1972) gubernatoris (Giles, 1901), comb. nov. schroederi (Schick, 1970) var. kotiensis (Barraud, 1934), comb. nov. sumidero (Schick, 1970) inquinata (Edwards, 1922), comb. nov. tehuantepec (Schick, 1970) iwi (Marks, 1955), comb. nov. terrens (Walker, 1856) khazani (Edwards, 1922), comb. nov.

kiangsiensis (Tung, 1955), comb. nov. longipalpus Randolph & ONeill, 1944 lophoventralis (Theobald, 1910), comb. nov. lutzii (Theobald, 1901) melanoptera (Giles, 1904), comb. nov. mathesoni Belkin & Heinemann, 1975 plumifera (King & Hoogstraal, 1946), comb. melanota Cerqueira, 1943 nov. mexicana (Bellardi, 1859) prominens (Barraud, 1923), comb. nov. pilosa Duret, 1971 stevensoni (Barraud, 1923), comb. nov. pseudoalbipes Duret, 1971 watasei (Yamada, 1921), comb. nov. pseudomelanota Barata & Cotrim, 1971 im totonaci Lassmann, 1951 Pseudoarmigeres Stone & Knight, 1956 varipes (Coquillett, 1904) albomarginatus (Newstead, 1907) Subgenus Psorophora argenteoventralis (Theobald, 1909) ciliata (Fabricius, 1794) ssp. dunni (Evans, 1928) cilipes (Fabricius, 1805) kummi (Edwards, 1930) holmhergii Lynch Arribaizaga, 1891 michaelikati (van Someren, 1946) howardii Coquillett, 1901 ssp. gurneri (van Someren, 1946) lineata (von Humboldt, 1819) natalensis (Edwards, 1930) ochripes (Macquart, 1850) pallescens Edwards, 1922 Pseudoskusea Theobald, 1907, stat. nov., restored pilipes to genus rank (Macquart, 1834) saeva Dyar & Knab, 1906 bancroftiana (Edwards, 1921), comb. nov. stonei Vargas, 1956 culiciformis (Theobald, 1905), comb. nov. Nomina dubia multiplex (Theobald, 1903), comb. nov. goeldii postspiraculosa (Dobrotworsky, 1961), comb. (Giles, 1904) marmorata nov. (Philippi, 1865) perterrens (Walker, 1856) Psorophora Robineau-Desvoidy, 1827 Subgenus Grabhamia Theobald, 1903 Rampamyia Reinert, Harbach & Kitching, gen. cingulata (Fabricius, 1805) ,nov. columbiae (Dyar & Knab, 1906) albilabris (Edwards, 1925), comb. nov. confinnis (Lynch Arribaizaga, 1891) notoscripta (Skuse, 1889), comb. nov. dimidiata Cerqueira, 1943 ssp. montana (Brug, 1939), comb. nov. discolor (Coquillett, 1903) quinquelineata (Edwards, 1922), comb. nov. infinis (Dyar & Knab, 1906) insularia (Dyar & Knab, 1906) Rhinoskusea jamaicensis Theobald, 1901 Edwards, 1929 longirostris leucocnemis Martini, 1931 (Leicester, 1908) pillaii paulli Paterson & Shannon, 1927 (Mattingly, 1958) pruinosa Martini, 1935 portonovoensis (Tewari & Hiriyan, 1992) pygmaea (Theobald, 1903) wardi (Reinert, 1976) santamarinai Broche, 2000 signipennis (Coquillett, 1904) Scutomyia Theobald, 1904 varinervis Edwards, 1922 albolineata Theobald, 1904 Subgenus Janthinosoma Lynch Arribaizaga, 1891 arhori cola (Knight & Rozeboom, 1946) albigenu (Peryassu, 1908) bambusicola Knight & Rozeboom, 1946 albipes (Theobald, 1907) boharti (Knight & Rozeboom, 1946) amazonica Cerqueira, 1960 hoogstraali (Knight & Rozeboom, 1946) champerico (Dyar & Knab, 1906) impatibilis (Walker, 1859) circumfiava Cerqueira, 1943 laffooni (Knight & Rozeboom, 1946) cyanescens (Coquillett, 1902) platylepida (Knight & Hull, 1951) discrucians (Walker, 1856) pseudalbolineata (Brug, 1939) ferox (von Humboldt, 1819) fiebrigi Edwards, 1922 Skusea Theobald, 1903 forceps Cerqueira, 1939 cartroni (Ventrillon, 1906) horrida (Dyar & Knab, 1908) lamhrechti (van Someren, 1971) johnstonii (Grabham, 1905) moucheti (Ravaonjanahary & Brunhes, 1977) lanei Shannon & Cerqueira, 1943 pembaensis (Theobald, 1901)

Stegomyia Theobald, 1901 hogsbackensis (Huang, 2004), comb. nov. aegypti (Linnaeus, 1762) hoguei (Belkin, 1962) ssp. formosus (Walker, 1848) horrescens (Edwards, 1935) africana Theobald, 1901 jostahae (Huang, 1988) agrihanensis (Bohart, 1957) katherinensis (Woodhill, 1949) albopicta (Skuse, 1895) keniensis (van Someren, 1946) alcasidi (Huang, 1972) kenyae (van Someren, 1946) alorensis (Bonne-Wepster & Brug, 1932) kesseli (Huang & Hitchcock, 1980) amaithea (de Meillon & Lavoipierre, 1944) kivuensis (Edwards, 1941) andrewsi (Edwards, 1926) krombeini (Huang, 1975) angusta (Edwards, 1935) lamberti (Ventrillon, 1904) annandalei Theobald, 1910 langata (van Someren, 1946) var. quadricincta (Barraud, 1923) ledgeri (Huang, 1981) aobae (Belkin, 1962) lilii Theobald, 1910 apicoargentea Theobald, 1909 luteocephala Newstead, 1907 bambusae (Edwards, 1935) maehleri (Bohart, 1957) blacklocki (Evans, 1925), comb. nov. malayensis (Colless, 1962) bromeliae Theobald, 1911 malikuli (Huang, 1973) hurnsi (Basio & Reisen, 1971) marshallensis (Stone & Bohart, 1944) calceata (Edwards, 1924) mascarensis (MacGregor, 1924) chaussieri (Edwards, 1923) masseyi (Edwards, 1923) chemulpoensis (Yamada, 1921) mattinglyorum (Huang, 1994) contigua (Edwards, 1936) maxgermaini (Huang, 1990) cooki (Belkin, 1962) mediopunctata Theobald, 1905 corneti (Huang, 1986) var. sureilensis (Barraud, 1934) cozi (Cornet, 1973) metallica (Edwards, 1912) craggi Barraud, 1923 mickevichae (Huang, 1988) cretina (Edwards, 1921) mpusiensis (Huang, 2004), comb. nov. daitensis (Miyagi & Toma, 1981) muroafcete (Huang, 1997) deboeri (Edwards, 1926) neoafricana (Cornet, Valade & Dieng, 1978) demeilloni (Edwards, 1936) neogalloisi (Chen & Chen, 2000) denderensis (Wolfs, 1949) neopandani (Bohart, 1957) dendrophila (Edwards, 1921) njomhiensis (Huang, 1997) desmotes Giles, 1904 novalbopicta (Barraud, 1931) dybasi (Bohart, 1957) opok (Corbet & van Someren, 1962) ealaensis (Huang, 2004), comb. nov. palauensis (Bohart, 1957) edwardsi Barraud, 1923 pandani (Stone, 1939) ethiopiensis (Huang, 2004), comb. nov. patriciae (Mattingly, 1954) flavopicta (Yamada, 1921) paullusi (Stone & Farner, 1945) ssp. downsi (Bohart & Ingram, 1946) pernotata Warner & Bohart, 1944) ssp. miyarai (Tanaka, Mizusawa & Saugstad, 1979) perplexa Leicester, 1908 fraseri Edwards, 1912 polynesiensis (Marks, 1951) futunae (Belkin, 1962) poweri Theobald, 1905 galloisi (Yamada, 1921) pseudalbopicta Borel, 1928 galloisiodes (Liu & Lu, 1984) pseudoafricana (Chwatt, 1949) gandaensis (Huang, 2004), comb. nov. pseudonigeria Theobald, 1910 gardnerii Ludlow, 1905 pseudoscutellaris Theobald, 1910 ssp. imitator Leicester, 1908 quasiscutellaris (Farner & Bohart, 1944) grantii Theobald, 1901 rhungkiangensis (Chang & Chang, 1974) guamensis (Farner & Bohart, 1944) riversi (Bohart & Ingram, 1946) gurneyi (Stone & Bohart, 1944) robinsoni (Belkin, 1962) hakanssoni (Knight & Hurlbut, 1949) rotana (Bohart & Ingram, 1946) hansfordi (Huang, 1997) rotumae (Belkin, 1962) hebridea (Edwards, 1926) ruwenzori (Haddow & van Someren, 1950) heischi (van Someren, 1951) saimedres (Huang, 1988) hensilli (Farner, 1945) saipanensis (Stone, 1945)

sampi (Huang, 2004), comb. nov. robertsi (Laffoon, 1946) schwetzi (Edwards, 1926) srilankensis (Reinert, 1977) scutellaris (Walker, 1858) stunga (Klein, 1973) scutoscripta (Bohart & Ingram, 1946) uniformis (Theobald, 1910) seampi (Huang, 1974) yusafi (Barraud, 1931) seatoi (Huang, 1969) Subgenus Neomacleaya Theobald, 1907 segermanae (Huang, 1997) adusta (Laffoon, 1946) sibirica (Danilov & Filippova, 1978) agrestis (Barraud, 1931) -S. simpsoni Theobald, 1905 andamanensis (Edwards, 1922) soleata (Edwards, 1924) assamensis Bhattacharyya, Tewari, Prakash, strelitziae (Muspratt, 1950) Mohapatra & Mahanta, 2004 subaihopicta (Barraud, 1931) atriisimilis (Tanaka & Mizusawa, 1973) subargentea (Edwards, 1925) atria (Barraud, 1928) tabu (Ramalingam & Belkin, 1965) campylostylus (Laffoon, 1946) tongae (Edwards, 1926) cauta (Barraud, 1928) tulagiensis (Edwards, 1926) clavata (Barraud, 1931) unilineata (Theobald, 1906) comata (Barraud, 1931) upolensis (Marks, 1957) comosa (Reinert, 1974) usambara (Mattingly, 1953) cretata (Delfinado, 1967 varuae (Belkin, 1962) cyrtolabis (Edwards, 1928) vinsoni (Mattingly, 1953) gibbosa (Delfinado, 1967) wadai (Tanaka, Mizusawa & Saugstad, 1979) harrisonica (Reinert, 1974) w-al bus Theobald, 1905 hispida (Delfinado, 1967) woodi (Edwards, 1922) incerta (Edwards, 1922) Nomina dubia indica (Theobald, 1907) ciliaris (Linnaeus, 1767) johnsoni (Laffoon, 1946) niveus (von Eichwald, 1837) johorensis (Reinert, 1974) rimandoi (Basio, 1971) komponga (Klein, 1973) lankaensis (Stone & Knight, 1958) latipennis Tanakaius Reinert, Harbach & Kitching, 2004 (Delfinado, 1967) leicesteri savoryi (Bohart, 1957) (Edwards, 1917) macrodixoa (Dyar & Shannon, 1925) togoi (Theobald, 1907) margarsen (Dyar & Shannon, 1925) neomacrodixoa (King & Hoogstraal, 1947) Udaya Thurman, 1954 nigrotarsis (Ludlow, 1908) argyrurus (Edwards, 1934) notahilis (Delfinado, 1967) lucaris Macdonald & Mattingly, 1960 nubicola (Laffoon, 1946) subsimilis (Barraud, 1927) panayensis (Ludlow, 1914) petroelephantus (Wijesundara, 1951) philippinensis Vansomerenis Reinert, Harbach & Kitching, gen. (Delfinado, 1968) nov. phnoma (Klein, 1973) prioekanensis (Brug, 1931) hancocki (van Someren, 1962), comb. nov. protuberans luteostriata (Robinson, 1950), comb. nov. (Delfinado, 1967) pseudodiurna pulchrithorax (Edwards, 1939), comb. nov. (Theobald, 1910) pseudomediofasciata (Theobald, 1910) pseudovarietas (Reinert, 1974) Verrallina Theobald, 1903 rami (Barraud, 1928) Subgenus Harhachius Reinert, 1999 rara (Delfinado, 1968) abdita (Barraud, 1931) sabahensis (Reinert, 1974) consonensis (Reinert, 1973) seculata (Menon, 1950) fragilis Leicester, 1908 singularis (Leicester, 1908) hamistylus (Laffoon, 1946) sohni (Reinert, 1974) indecorabilis Leicester, 1908 spermathecus (Wijesundara, 1951) nobukonis (Yamada, 1932) torosa (Delfinado, 1967) pahangi (Delfinado, 1968) uncus (Theobald, 1901) ramalingami (Reinert, 1974) vallistris (Barraud, 1928)

varietas (Leicester, 1908) APPENDIX 5 virilis Leicester, 1908 yerhuryi (Edwards, 1917) ABBREVIATIONS OF AEDINE GENERA Subgenus Verrallina Recommended two-letter abbreviations of formally azureosquamata (Bonne-Wepster, 1948) recognized genera of Aedini. bifoliata (King & Hoogstraal, 1947) butleri (Theobald, 1901) Abraedes Zavortink = Ab. carmenti (Edwards, 1924) Aedes Meigen = Ac. cuccioi (Belkin, 1962) Alanstonea Mattingly = As. cunninghami (Taylor, 1944) Albuginosus Reinert = Al. dux (Dyar & Shannon, 1925) Armigeres Theobald = Ar. embiensis (Huang, 1968) Ayurakitia Thurman = Ay. foliforinis (King & Hoogstraal, 1947) Aztecaedes Zavortink = Az. funerea (Theobald, 1903) Belkinius Reinert = Be. iriomotensis (Tanaka & Mizusawa, 1973) Bothaella Reinert = Bo. kiliertonis (Huang, 1968) Bruceharrisonius Reinert = Br. leilae (King & Hoogstraal, 1947) Christophersiomyia Barraud = Cr. lineata (Taylor, 1914) Collessius Reinert, Harbach & Kitching = Co. lugubris (Barraud, 1928) Dahiiana Reinert, Harbach & Kitching = Da. mccormicki (Belkin, 1962) Diceromyia Theobald = Di. milnensis (King & Hoogstraal, 1947) Dobrotworskyius Reinert, Harbach & Kintching = Db. muitifolium (King & Hoogstraal, 1947) Downsiomyia Vargas = Do. obsoleta (Huang, 1968) Edwardsaedes Belkin = Ed. parasimilis (King & Hoogstraal, 1947) Eretmapodites Theobald = Er. pipkini (Bohart, 1957) Finlaya Theobald = Fl. quadrifolium (Brug, 1934) Fredwardsius Reinert = Fr. quadrispinata (King & Hoogstraal, 1947) Georgecraigius Reinert, Harbach & Kitching = Ge. reesi (King & Hoogstraal, 1947) Geoskusea Edwards = Ge. sentania (King & Hoogstraal, 1947) Gilesius Reinert, Harbach & Kitching = Gi. similis (Theobald, 1910) Gymnometopa Coquillett = Gy. simpla (King & Hoogstraal, 1947) Haemagogus Williston = Hg. trispinata (King & Hoogstraal, 1947) Halaedes Belkin = Ha. vanapa (Huang, 1968) Heizmannia Ludlow = Hz. variabilis (Huang, 1968) Himalaius Reinert, Harbach & Kitching = Hi. Nomen dubium Howardina Theobald = Hw. perdita Leicester, 1908 Huaedes Huang = Hu. Hulecoeteomyia Theobald = Hi. Indusius Edwards = In. Zavortinkius Reinert, 1999 Isoaedes Reinert = Ia. brunhesi (Reinert, 1999) Jarneliius Reinert, Harbach & Kitching = Ja. hrygooi (Brunhes, 1971) Jihiienius Reinert, Harbach & Kitching = Ji. fulgens (Edwards, 1917) Kenknightia Reinert = Ke. geoffroyi (Reinert, 1999) Kompia Aitken = Ko. huangae (Reinert, 1999) Leptosomatomyia Theobald = Lp. interruptus (Reinert, 1999) Levua Stone & Bohart = Le. longipalpis (Grunberg, 1905) Lorrainea Belkin = Lo. •1. monetus (Edwards, 1935) Macleaya Theobald = Mc. mzooi (van Someren, 1962) Molpemyia Theobald = Mo. phillipi (van Someren, 1949) Mucidus Theobald = Mu. pollinctor (Graham, 1910) Neomelaniconion Newstead = Ne. Ochierotatus Lynch Arribaizaga = Oc. Zeugnomyia Leicester, 1908 Opifex Hutton = Op. aguilari Baisas & Feliciano, 1953 Paraedes Edwards = Pr. fajardoi Baisas & Feliciano, 1953 Patmarksia Reinert, Harbach & Kitching = Pm. gracilis Leicester, 1908 Phagomyia Theobald = Ph. lawtoni Baisas, 1946 Pseudarmigeres Stone & Knight = Pa.

Pseudoskusea Theobald = Pk. Tanakaius Reinert, Harbach & Kitching = Ta. Psorophora Robineau-Desvoidy = Ps. Udaya Thurman Ud. Rampamyia Reinert, Harbach & Kitching = Ra Vansomerenis Reinert, Harbach & Kitching = Va. Rhinoskusea Edwards = Rh. Verrallina Theobald = Ve. Scutomyia Theobald = Sc. Zavortinkius Reinert = Za. Skusea Theobald = Sk. Zeugnomyia Leicester = Ze. Stegomyia Theobald = St.