A Stable Classification of Tribe Aedini That Balances Utility with Current Knowledge of Evolutionary Relationships

RESEARCH ARTICLE Making Mosquito Taxonomy Useful: A Stable Classification of Tribe Aedini that Balances Utility with Current Knowledge of Evolutionary Relationships

Richard C. Wilkerson1*, Yvonne-Marie Linton1,2,3,4, Dina M. Fonseca5, Ted R. Schultz1, Dana C. Price5, Daniel A. Strickman6

1 Department of Entomology, National Museum of Natural History, Smithsonian Institution, Washington DC, United States of America, 2 Walter Reed Biosystematics Unit, Museum Support Center, Smithsonian Institution, Suitland, Maryland, United States of America, 3 Department of Entomology, Walter Reed Army Institute of Research, Silver Spring, Maryland, United States of America, 4 Faculty of Preventative Medicine and Biometrics, Uniformed Services University of the Health Sciences, Bethesda, Maryland, United States of America, 5 Entomology Department, School of Environmental and Biological Sciences, Rutgers University, New Brunswick, New Jersey, United States of America, 6 Global Health Program, Bill and Melinda Gates Foundation, Seattle, Washington, United States of America OPEN ACCESS * [email protected] Citation: Wilkerson RC, Linton Y-M, Fonseca DM, Schultz TR, Price DC, Strickman DA (2015) Making Mosquito Taxonomy Useful: A Stable Classification of Tribe Aedini that Balances Utility with Current Abstract Knowledge of Evolutionary Relationships. PLoS ONE The tribe Aedini (Family Culicidae) contains approximately one-quarter of the known spe- 10(7): e0133602. doi:10.1371/journal.pone.0133602 cies of mosquitoes, including vectors of deadly or debilitating disease agents. This tribe Editor: Zhong-Jian Liu, The National Orchid contains the genus Aedes, which is one of the three most familiar genera of mosquitoes. Conservation Center of China; The Orchid Conservation & Research Center of Shenzhen, During the past decade, Aedini has been the focus of a series of extensive morphology- CHINA based phylogenetic studies published by Reinert, Harbach, and Kitching (RH&K). Those Received: March 8, 2015 authors created 74 new, elevated or resurrected genera from what had been the single genus Aedes, almost tripling the number of genera in the entire family Culicidae. The Accepted: June 22, 2015 proposed classification is based on subjective assessments of the “number and nature of Published: July 30, 2015 the characters that support the branches” subtending particular monophyletic groups in Copyright: This is an open access article, free of all the results of cladistic analyses of a large set of morphological characters of representative copyright, and may be freely reproduced, distributed, species. To gauge the stability of RH&K’s generic groupings we reanalyzed their data with transmitted, modified, built upon, or otherwise used by anyone for any lawful purpose. The work is made unweighted parsimony jackknife and maximum-parsimony analyses, with and without available under the Creative Commons CC0 public ordering 14 of the characters as in RH&K. We found that their phylogeny was largely weakly domain dedication. supported and their taxonomic rankings failed priority and other useful taxon-naming crite- Data Availability Statement: Data analyzed in this ria. Consequently, we propose simplified aedine generic designations that 1) restore a study were originally generated and published by classification system that is useful for the operational community; 2) enhance the ability of Reinert, Harbach and Kitching (2009), Zool J Linn Soc 157: 700-794, and were retrieved manually. The taxonomists to accurately place new species into genera; 3) maintain the progress toward a nexus file dataset we used is publically available on natural classification based on monophyletic groups of species; and 4) correct the current the website of the Walter Reed Biosystematics Unit: classification system that is subject to instability as new species are described and existing (www.wrbu.org/DOCS/MQ0463.pdf). species more thoroughly defined. We do not challenge the phylogenetic hypotheses gener- Funding: The authors received no specific funding ated by the above-mentioned series of morphological studies. However, we reduce the for this work, except institutional support. This manuscript was prepared while YML held a National

PLOS ONE | DOI:10.1371/journal.pone.0133602 July 30, 2015 1/26 Stable Mosquito Classification

Research Council (NRC) Research Associateship ranks of the genera and subgenera of RH&K to subgenera or informal species groups, Award at the Walter Reed Army Institute of Research. respectively, to preserve stability as new data become available. The affiliated institutions had no role in study design, data collection and analysis, decision to publish, or preparation of the manuscript. The material to be published reflects the views of the authors and should not be construed to represent those of the US Department of the Army or the US Department of Introduction Defense. The Catalog of the Mosquitoes of the World, published in 1977 by Knight and Stone [1] and Competing Interests: The authors have declared – that no competing interests exist. its three supplements [2 5], which were based on an earlier 1959 catalog [6] and supplements [7–11], are the primary reference point for all modern mosquito systematics. These catalogs summarize the nomenclatorial organization of the entire Culicidae family and list all references that established those names. The family Culicidae includes 3,601 described species and sub- species (www.mosquitocatalog.org accessed 22 Jan., 2014), a number that steadily increases due to the intense medical and veterinary importance of mosquitoes. The family is organized into two subfamilies, the Anophelinae (482 species) and the Culicinae (3,119 species). Aedini, with 1,261 species, is the largest of the 11 tribes within Culicinae. The genus Aedes in the tribe Aedini is one of three best-known genera of mosquitoes (along with Culex and Anopheles) since many of its species are important vectors of arboviruses and pathogens (Table 1). Because of its medical importance, Aedes is recognized by thousands of researchers and operators con- cerned with mosquito control, public health, and veterinary health.

Formal Taxonomic Action This publication restores the generic structure of the tribe Aedini (Diptera: Culicidae) to its status prior to the year 2000. Specifically, all species in the tribe are placed in the following 10 genera: Aedes, Armigeres, Eretmapodites, Haemagogus, Heizmannia, Opifex, Psorophora, Udaya, Verrallina, and Zeugnomyia. Aedine genera created in and after 2000 are placed as subgenera of genus Aedes; subgenera created after 2000 are placed in informal species groupings (S1 Appendix, S2 Appendix). The classification proposed here, and further updates to culicid classification, will be reflected in the online taxonomic catalog at www.mosquitocatalog.org. The 1977 catalog [1] lists nine genera in the tribe Aedini (Aedes Meigen, 1818; Armigeres Theobald, 1901; Eretmapodites Theobald, 1901; Haemagogus Williston, 1896; Heizmannia Ludlow, 1905; Opifex Hutton, 1902; Psorophora Robineau-Desvoidy, 1827; Udaya Thurman, 1954; and Zeugnomyia Leicester, 1908). Of the 51 subgenera in Aedini, 38 subgenera with about 800 species were placed in the genus Aedes.An“Aedes Group” was mentioned by Edwards in 1932 [58] in a compendium of mosquito taxonomy that preceded the 1959 and 1977 catalogs. His Aedes Group included all of the above except Udaya, which had not yet been described, and Zeugnomyia, which was then considered to be in the Uranotaenia group. Tribe Aedini with all nine genera was first mentioned by Belkin in 1962 [27] in a two-volume work on mosquitoes of the South Pacific that also discussed mosquito systematics and biogeog- raphy in a world-wide context. Since Knight and Stone’s catalog and supplements, five new aedine subgenera were added (Isoaedes [59], Belkinius [60], Albuginosus [61], Kenknightia [62] and Zavortinkius [63]). Also during this period three subgenera of Aedes were elevated to genus, one in 1999 (Verrallina [64]) and two in 2000 (Ayurakitia [65] and Ochlerotatus [66]). Elevation of Verrallina and Ayurakitia to genus went largely unnoticed except by taxonomists. In contrast, because Ochler- otatus contained some well-known species, e.g. Ae.(Och.) sollicitans (Walker), a salt-marsh mosquito and important nuisance and disease vector (Table 1), there was a good deal of debate as to the merit of elevating it to a genus [67]. The classification that included Ochlerotatus as a

PLOS ONE | DOI:10.1371/journal.pone.0133602 July 30, 2015 2/26 Stable Mosquito Classification

Table 1. Name combinations used since 1977 for selected mosquitoes belonging to Tribe Aedini with associated human pathogens and invasive (established, spreading) or non-native (established, not spreading) behaviors. Names proposed herein are in bold typeface; previous name combinations follow in normal typeface.

SPECIES ASSOCIATED HUMAN PATHOGENS / INVASIVE AND NON-NATIVE SPECIES Aedes (Aedes) cinereus Meigen* EEEV[12], HJV[13], JCV[13], Aedes cinereus Meigen TAHV, WNV[14] Aedes (Aedimorphus) abnormalis (Theobald)* SFV, WSLV 'Aedes' ('Aedimorphus') abnormalis (Theobald) Aedimorphus abnormalis (Theobald) Aedes (Aedimorphus) cumminsii (Theobald)* RVFV[15], SPOV 'Aedes' ('Aedimorphus') cumminsii (Theobald) Aedimorphus cumminsii (Theobald) Aedes (Aedimorphus) dalzieli (Theobald)* BBKV[15], BOUV[15], CHIKV[15], 'Aedes' ('Aedimorphus') dalzieli (Theobald) KEDV, NDOV, PGAV, RVFV[15], WSLV, ZIKAV Aedimorphus dalzieli (Theobald) Aedes (Aedimorphus) dentatus (Theobald)* ORUV, RVFV[16], WSLV 'Aedes' ('Aedimorphus') dentatus (Theobald) Aedimorphus dentatus (Theobald) Aedes (Aedimorphus) fowleri (de Charmoy)* PGAV, RVFV+[17] 'Aedes' ('Aedimorphus') fowleri (de Charmoy) Aedimorphus fowleri (de Charmoy) Aedes (Aedimorphus) hirsutus (Theobald)* NRIV[18] 'Aedes' ('Aedimorphus') hirsutus (Theobald) Aedimorphus hirsutus (Theobald) Aedes (Aedimorphus) mediolineatus (Theobald)* WSLV 'Aedes' ('Aedimorphus') mediolineatus (Theobald) Aedimorphus mediolineatus (Theobald) Aedes (Aedimorphus) natrionius Edwards* UGSV 'Aedes' ('Aedimorphus') natronius Edwards Aedimorphus natronius (Edwards) Aedes (Aedimorphus) ochraceus (Theobald)* RVFV[15], WSLV[15] 'Aedes' ('Aedimorphus') ochraceus (Theobald) Aedimorphus ochraceus (Theobald) Aedes (Aedimorphus) vexans (Meigen)* TAHV, TVTV, WEE+[19], WNV[14,15], 'Aedes' ('Aedimorphus') vexans (Meigen) Banna[20], Chaoyang[21], Potosi[22] Aedimorphus vexans (Meigen) Aedes (Catageiomyia) argenteopunctatus (Theobald) NRIV, SFV Aedes (Aedimorphus) argenteopunctatus (Theobald)* 'Aedes' ('Aedimorphus') argenteopunctatus (Theobald) Catageiomyia argenteopunctata (Theobald) Aedes (Catageiomyia) minutus (Theobald) KEDV Aedes (Aedimorphus) minutus (Theobald)* 'Aedes' ('Aedimorphus') minutus (Theobald) Catageiomyia minuta (Theobald) Aedes (Catageiomyia) tarsalis (Newstead) KEDV, PGAV, WSLV Aedes (Aedimorphus) tarsalis (Newstead)* 'Aedes' ('Aedimorphus') tarsalis (Newstead) (Continued)

PLOS ONE | DOI:10.1371/journal.pone.0133602 July 30, 2015 3/26 Stable Mosquito Classification

Table 1. (Continued)

SPECIES ASSOCIATED HUMAN PATHOGENS / INVASIVE AND NON-NATIVE SPECIES Catageiomyia tarsalis (Newstead) Aedes (Diceromyia) furcifer (Edwards)* BOUV, CHIK[23], RVFV[15], Diceromyia furcifer (Edwards) YF[24] Aedes (Diceromyia) taylori Edwards* CHIK[23], YFV[24] Diceromyia taylori (Edwards)* Aedes (Downsiomyia) harinasutai Knight W. bancrofti[25] Aedes (Finlaya) harinasutai Knight* Downsiomyia harinasutai (Knight) Aedes (Downsiomyia) niveus (Ludlow) DENV, W. bancrofti[26] Aedes (Finlaya) niveus (Ludlow)* Downsiomyia nivea (Ludlow) Aedes (Finlaya) ﬁjiensis Marks* W. bancrofti[27] Finlaya ﬁjiensis (Marks) Aedes (Finlaya) kochi (Dönitz)* W. bancrofti[28] Ochlerotatus (Finlaya) kochi (Dönitz) Finlaya kochi (Dönitz) Aedes (Finlaya) poicilius (Theobald)* W. bancrofti[29] Ochlerotatus (Finlaya) poicilius (Theobald) Finlaya poicilia Theobald Aedes (Fredwardsius) vittatus (Bigot) BBKV, NRIV, PGAV, SFV Aedes (Stegomyia) vittatus (Bigot)* Fredwardsius vittatus (Bigot) Aedes (Georgecraigius) epactius Dyar & Knab WNV[14] Aedes (Ochlerotatus) epactius Dyar & Knab* Ochlerotatus (Ochlerotatus) epactius (Dyar & Knab) 'Ochlerotatus' ('Ochlerotatus') epactius (Dyar & Knab) Georgecraigius (Georgecraigius) epactius (Dyar & Knab) Aedes (Georgecraigius) atropalpus (Coquillett) Invasive[30], WNV[14] Aedes (Ochlerotatus) atropalpus (Coquillett)* Ochlerotatus (Ochlerotatus) atropalpus (Coquillett) 'Ochlerotatus' ('Ochlerotatus') atropalpus (Coquillett) Georgecraigius (Georgecraigius) atropalpus (Coquillett) Aedes (Hopkinsius) ingrami (Edwards) UGSV Aedes (Finlaya) ingrami Edwards* Ochlerotatus (Finlaya) ingrami (Edwards) 'Ochlerotatus' ('Finlaya') ingrami (Edwards) Hopkinsius (Hopkinsius) ingrami (Edwards) Aedes (Howardina) septemstriatus Dyar & Knab* WYOV Ochlerotatus (Howardina) septemstriatus (Dyar & Knab) Howardina septemstriata (Dyar & Knab) Aedes (Howardina) sexlineatus (Theobald)* WYOV Ochlerotatus (Howardina) sexlineatus (Theobald) Howardina sexlineata (Theobald) Aedes (Hulecoeteomyia) japonicus (Theobald) Invasive[30], CVV[31], WNV[14] Aedes (Finlaya) japonicus (Theobald)* Ochlerotatus (Finlaya) japonicus (Theobald) (Continued)

PLOS ONE | DOI:10.1371/journal.pone.0133602 July 30, 2015 4/26 Stable Mosquito Classification

Table 1. (Continued)

SPECIES ASSOCIATED HUMAN PATHOGENS / INVASIVE AND NON-NATIVE SPECIES 'Ochlerotatus' ('Finlaya') japonicus (Theobald) Hulecoeteomyia japonica (Theobald) Aedes (Hulecoeteomyia) koreicus (Edwards) Invasive[32] Aedes (Finlaya) koreicus (Edwards)* Ochlerotatus (Finlaya) koreicus Edwards 'Ochlerotatus' ('Finlaya') koreicus Edwards Hulecoeteomyia koreica (Edwards) Aedes (Neomelaniconion) mcintoshi Huang* NRIV[15], MIDV[33], RVFV[34], Neomelaniconion mcintoshi (Huang) WSLV[33] Aedes (Neomelaniconion) palpalis (Newstead)* RVFV[15], SFV Neomelaniconion palpale Newstead Aedes (Neomelaniconion) circumluteolus (Theobald)* BUNV, LEBV, PGAV, RVFV, SPON, WSLV Neomelaniconion circumluteolus (Theobald) Neomelaniconion circuluteolum (Theobald) Aedes (Neomelaniconion) lineatopennis (Ludlow)* WSLV Neomelaniconion lineatopenne (Ludlow) Aedes (Ochlerotatus) abserratus (Felt & Young)* JCV Ochlerotatus (Ochlerotatus) abserratus (Felt & Young) 'Ochlerotatus' ('Ochlerotatus') abserratus (Felt & Young) Ochlerotatus (subgenus unassigned) abserratus (Felt & Young) Ochlerotatus (Woodius) abserratus (Felt & Young) Aedes (Ochlerotatus) angustivittatus Dyar & Knab* ILHV[35], VEEV Ochlerotatus (Ochlerotatus) angustivittatus (Dyar & Knab) 'Ochlerotatus' ('Ochlerotatus') angustivittatus (Dyar & Knab) Aedes (Ochlerotatus) argyrothorax (Bonne-Wepster & WYOV Bonne)* Aedes (Protomacleaya) argyrothorax (Bonne-Wepster & Bonne) Ochlerotatus (Protomacleaya) argyrothorax (Bonne- Wepster & Bonne) 'Ochlerotatus' ('Protomacleaya') argyrothorax (Bonne- Wepster & Bonne) Aedes (Ochlerotatus) atlanticus Dyar & Knab* EEEV, EYEV, TENV, TVTV, WNV[14] Ochlerotatus (Ochlerotatus) atlanticus (Dyar & Knab) Ochlerotatus (subgenus unassigned) atlanticus (Dyar & Knab) Ochlerotatus (Protoculex) atlanticus (Dyar & Knab) Aedes (Ochlerotatus) bancroftianus (Edwards) BAMV Aedes (Pseudoskusea) bancroftianus Edwards* Ochlerotatus (Pseudoskusea) bancroftianus (Edwards) 'Ochlerotatus' ('Pseudoskusea') bancroftianus (Edwards) Pseudoskusea bancroftiana (Edwards) Aedes (Ochlerotatus) caballus (Theobald)* RVFV, WSLV Ochlerotatus (Ochlerotatus) caballus (Theobald) 'Ochlerotatus' ('Ochlerotatus') caballus (Theobald) Ochlerotatus (subgenus unassigned) caballus (Theobald) (Continued)

PLOS ONE | DOI:10.1371/journal.pone.0133602 July 30, 2015 5/26 Stable Mosquito Classification

Table 1. (Continued)

SPECIES ASSOCIATED HUMAN PATHOGENS / INVASIVE AND NON-NATIVE SPECIES Ochlerotatus (Juppius) caballus (Theobald) Aedes (Ochlerotatus) canadensis (Theobald)* EEEV[12], HJV[36], JCV[13], Ochlerotatus (Ochlerotatus) canadensis (Theobald) RVFV+[37], WNV[19] 'Ochlerotatus' ('Ochlerotatus') canadensis (Theobald) Ochlerotatus (subgenus unassigned) canadensis (Theobald) Ochlerotatus (Culicada) canadensis (Theobald) Aedes (Ochlerotatus) cantans (Meigen)* TAHV, WNV Ochlerotatus (Ochlerotatus) cantans (Meigen) 'Ochlerotatus' ('Ochlerotatus') cantans (Meigen) Ochlerotatus (subgenus unassigned) cantans (Meigen) Ochlerotatus (Woodius) cantans (Meigen) Aedes (Ochlerotatus) cantator (Coquillett)* CVV[31], EEEV[36], HJV[36], JCV, Ochlerotatus (Ochlerotatus) cantator (Coquillett) RVFV+[37], WNV[14] 'Ochlerotatus' ('Ochlerotatus') cantator (Coquillett) Ochlerotatus (subgenus unassigned) cantator (Coquillett) Aedes (Ochlerotatus) caspius (Pallas)* TAHV, WNV[38] Ochlerotatus (Ochlerotatus) caspius (Pallas) 'Ochlerotatus' ('Ochlerotatus') caspius (Pallas) Ochlerotatus (subgenus unassigned) caspius (Pallas) Aedes (Ochlerotatus) communis (de Geer)* LACV, JCV, TAHV Ochlerotatus (Ochlerotatus) communis (de Geer) Ochlerotatus (subgenus unassigned) communis de Geer Aedes (Ochlerotatus) condolescens Dyar & Knab* WNV[14] 'Ochlerotatus' ('Ochlerotatus') condolescens (Dyar & Knab) Ochlerotatus (Ochlerotatus) condolescens (Dyar & Knab) Aedes (Ochlerotatus) detritus Haliday* TAHV[39] Ochlerotatus (Ochlerotatus) detritus (Haliday) 'Ochlerotatus' ('Ochlerotatus') detritus (Haliday) Ochlerotatus (subgenus unassigned) detritus (Haliday) Aedes (Ochlerotatus) diantaeus Howard, Dyar & Knab* TAHV Aedes (Ochlerotatus) diantaeus Howard, Dyar & Knab* Ochlerotatus (Ochlerotatus) diantaeus (Howard, Dyar & Knab) 'Ochlerotatus' ('Ochlerotatus') diantaeus (Howard, Dyar & Knab) Ochlerotatus (subgenus unassigned) diantaeus (Howard, Dyar & Knab) Ochlerotatus (Woodius) diantaeus (Howard, Dyar & Knab) Aedes (Ochlerotatus) dorsalis (Meigen)* WEE[40], WNV[14], Banna[20], Ochlerotatus (Ochlerotatus) dorsalis (Meigen) Liaoning[41] Ochlerotatus (subgenus unassigned) dorsalis (Meigen) Aedes (Ochlerotatus) dupreei (Coquillett)* WNV[14] Ochlerotatus (Ochlerotatus) dupreei (Coquillett) 'Ochlerotatus') dupreei (Coquillett) Ochlerotatus (Protoculex) dupreei (Coquillett) (Continued)

PLOS ONE | DOI:10.1371/journal.pone.0133602 July 30, 2015 6/26 Stable Mosquito Classification

Table 1. (Continued)

SPECIES ASSOCIATED HUMAN PATHOGENS / INVASIVE AND NON-NATIVE SPECIES Aedes (Ochlerotatus) excrucians (Walker)* RVFV+[37] Ochlerotatus (Ochlerotatus) excrucians (Walker) 'Ochlerotatus' ('Ochlerotatus') excrucians (Walker) Ochlerotatus (subgenus unassigned) excrucians (Walker) Aedes (Ochlerotatus) fitchii (Felt & Young)* WNV[14] Ochlerotatus (Ochlerotatus) fitchii (Felt & Young) 'Ochlerotatus' ('Ochlerotatus') fitchii (Felt & Young) Ochlerotatus (subgenus unassigned) fitchii (Felt & Young) Aedes (Ochlerotatus) fulvus (Wiedemann)* EEEV, WNV, WYOV Ochlerotatus (Ochlerotatus) fulvus (Wiedemann) Ochlerotatus (subgenus unassigned) fulvus (Wiedemann) Ochlerotatus (Chrysoconops) fulvus (Wiedemann) Aedes (Ochlerotatus) grossbecki Dyar & Knab* WNV[14] Ochlerotatus (Ochlerotatus) grossbecki (Dyar & Knab) 'Ochlerotatus' ('Ochlerotatus') grossbecki (Dyar & Knab) Ochlerotatus (subgenus unassigned) grossbecki (Dyar & Knab) Aedes (Ochlerotatus) infirmatus Dyar & Knab* EEEV[42], Keystone[42], Ochlerotatus (Ochlerotatus) infirmatus (Dyar & Knab) Tensaw[42], TVTV, WNV[14] Ochlerotatus (subgenus unassigned) infirmatus (Dyar & Knab) Aedes (Ochlerotatus) melanimon Dyar* SLEV, WEEV[19], WNV[14] Ochlerotatus (Ochlerotatus) melanimon (Dyar) 'Ochlerotatus' ('Ochlerotatus') melanimon (Dyar) Ochlerotatus (subgenus unassigned) melanimon (Dyar) Aedes (Ochlerotatus) mitchellae (Dyar)* EEEV, TENV Ochlerotatus (Ochlerotatus) mitchellae (Dyar) 'Ochlerotatus' ('Ochlerotatus') mitchellae (Dyar) Ochlerotatus (Culicelsa) mitchellae (Dyar) Aedes (Ochlerotatus) nigromaculis (Ludlow)* WNV[14] Ochlerotatus (Ochlerotatus) nigromaculis (Ludlow) 'Ochlerotatus' ('Ochlerotatus') nigromaculis (Ludlow) Ochlerotatus (Culicelsa) nigromaculis (Ludlow) Aedes (Ochlerotatus) normanensis (Taylor)* BAMV, GGV, MVEV, RRV, SINV Ochlerotatus (Ochlerotatus) normanensis (Taylor) 'Ochlerotatus' ('Ochlerotatus') normanensis (Taylor) Ochlerotatus (subgenus unassigned) normanensis (Taylor) Aedes (Ochlerotatus) provocans (Walker)* WNV[14] Ochlerotatus (Ochlerotatus) provocans (Walker) Ochlerotatus (Rusticoidus) provocans (Walker) Aedes (Ochlerotatus) punctor (Kirby)* BATV Ochlerotatus (Ochlerotatus) punctor (Kirby) 'Ochlerotatus' ('Ochlerotatus') punctor (Kirby) Ochlerotatus (subgenus unassigned) punctor (Kirby) Aedes (Ochlerotatus) scapularis (Rondani)* ILHV[43], SLEV, VEEV, WYOV, YFV[44], Ochlerotatus (Ochlerotatus) scapularis (Rondani) W. bancrofti[44] Ochlerotatus (subgenus unassigned) scapularis (Rondani) (Continued)

PLOS ONE | DOI:10.1371/journal.pone.0133602 July 30, 2015 7/26 Stable Mosquito Classification

Table 1. (Continued)

SPECIES ASSOCIATED HUMAN PATHOGENS / INVASIVE AND NON-NATIVE SPECIES Aedes (Ochlerotatus) serratus (Theobald)* SLEV, VEEV, WYOV Ochlerotatus (Ochlerotatus) serratus (Theobald) 'Ochlerotatus' ('Ochlerotatus') serratus (Theobald) Ochlerotatus (Protoculex) serratus (Theobald) Aedes (Ochlerotatus) sollicitans (Walker)* CVV[31], EEEV, RVFV+[37], Ochlerotatus (Ochlerotatus) sollicitans (Walker) VEEV, WNV[14] Ochlerotatus (subgenus unassigned) sollicitans (Walker) Ochlerotatus (Culicelsa) sollicitans (Walker) Aedes (Ochlerotatus) squamiger (Coquillett)* WNV[14] Ochlerotatus (Ochlerotatus) squamiger (Coquillett) 'Ochlerotatus' ('Ochlerotatus') squamiger (Coquillett) Ochlerotatus (subgenus unassigned) squamiger (Coquillett) Aedes (Ochlerotatus) sticticus (Meigen)* EEEV, JCV, TAHV, WNV[14] Ochlerotatus (Ochlerotatus) sticticus (Meigen) 'Ochlerotatus' ('Ochlerotatus') sticticus (Meigen) Ochlerotatus (subgenus unassigned) sticticus (Meigen) Aedes (Ochlerotatus) stimulans (Walker)* JCV, WNV[14] Ochlerotatus (Ochlerotatus) stimulans (Walker) 'Ochlerotatus' ('Ochlerotatus') stimulans (Walker) Ochlerotatus (subgenus unassigned) stimulans (Walker) Aedes (Ochlerotatus) taeniorhynchus (Wiedemann)* CVV, EEEV, EYEV, ORIV, RVFV+[37], Ochlerotatus (Ochlerotatus) taeniorhynchus (Wiedemann) TENV, TVTV, VEEV, WNV[14], WYOV 'Ochlerotatus' ('Ochlerotatus') taeniorhynchus (Wiedemann) Ochlerotatus (Culicelsa) taeniorhynchus (Wiedemann) Aedes (Ochlerotatus) thelcter Dyar* VEEV Ochlerotatus (Ochlerotatus) thelcter (Dyar) 'Ochlerotatus' ('Ochlerotatus') thelcter (Dyar) Aedes (Ochlerotatus) trivittatus (Coquillett)* EEEV[12], LACV, TVTV, WNV[14] 'Ochlerotatus' ('Ochlerotatus') trivittatus (Coquillett) Ochlerotatus (Ochlerotatus) trivittatus (Coquillett) Aedes (Ochlerotatus) vigilax (Skuse)* Non-native[30], BAMV, EHV, GGV, KOKV, RRV, SINV Ochlerotatus (Ochlerotatus) vigilax (Skuse) 'Ochlerotatus' ('Ochlerotatus') vigilax (Skuse) Ochlerotatus (Empihals) vigilax (Skuse) Aedes (Polyleptiomyia) albocephalus (Theobald) WNV[45] Aedes (Aedimorphus) albocephalus (Theobald)* 'Aedes' ('Aedimorphus') albocephalus (Theobald) Polyleptiomyia albocephala (Theobald) Aedes (Protomacleaya) triseriatus (Say)* CVV[46], EEEV[12], JCV++, LACV, 'Ochlerotatus' ('Protomacleaya') triseriatus (Say) RVFV+[37], WNV[14], Potosi[46] Aedes (Rampamyia) notoscriptus (Skuse) Invasive[30] Aedes (Finlaya) notoscriptus (Skuse)* Ochlerotatus (Finlaya) notoscriptus (Skuse) 'Ochlerotatus' ('Finlaya') notoscriptus (Skuse) (Continued)

PLOS ONE | DOI:10.1371/journal.pone.0133602 July 30, 2015 8/26 Stable Mosquito Classification

Table 1. (Continued)

SPECIES ASSOCIATED HUMAN PATHOGENS / INVASIVE AND NON-NATIVE SPECIES Rampamyia notoscripta (Skuse) Aedes (Skusea) pembaensis Theobald BUNV, TAHV Aedes (Skusea) pembaensis Theobald* Skusea pembaensis (Theobald) Aedes (Stegomyia) aegypti (Linnaeus)* Invasive[30], CHIK, DENV, ORUV, VEEV, WNV, YFV Stegomyia aegypti (Linnaeus) Stegomyia (Stegomyia) aegypti (Linnaeus) Aedes (Stegomyia) africanus (Theobald)* BBKV, BOUV, CHIK, RVFV[47], YFV, ZIKAV Stegomyia africana Theobald Stegomyia (subgenus unassigned) africana Theobald Aedes (Stegomyia) albopictus (Skuse)* Invasive[30], CHIK[48], CVV[46], Stegomyia albopicta (Skuse) DENV, EEEV[49], LACV[46], Stegomyia (subgenus unassigned) albopicta (Skuse) WNV[50], YFV[51] Aedes (Stegomyia) bromeliae (Theobald)* YFV[52] Stegomyia bromeliae Theobald Stegomyia (Mukwaya) bromeliae Theobald Aedes (Stegomyia) luteocephalus (Newstead)* CHIK[23], DEN[47], YFV[47], ZIKAV Stegomyia luteocephala Newstead Stegomyia (subgenus unassigned) luteocephala Newstead Aedes (Stegomyia) neoafricanus Cornet, Valade & NRIV Dieng* Stegomyia neoafricana (Cornet, Valade & Dieng) Stegomyia (subgenus unassigned) neoafricana (Cornet, Valade & Dieng) Aedes (Stegomyia) opok Corbet & van Someren* YFV Stegomyia opok (Corbet & van Someren) Stegomyia (subgenus unassigned) opok (Corbet & van Someren) Aedes (Stegomyia) polynesiensis Marks* W. bancrofti[27,53] Stegomyia polynesiensis (Marks) Stegomyia (subgenus unassigned) polynesiensis (Marks) Aedes (Stegomyia) scutellaris (Walker)* DEN[27], W. bancrofti[27] Stegomyia scutellaris (Walker) Stegomyia (subgenus unassigned) scutellaris (Walker) Aedes (Stegomyia) simpsoni (Theobald)* BBKV, NRIV+++, YFV Stegomyia simpsoni Theobald Aedes (Mukwaya) simpsoni (Theobald) Aedes (Tanakius) togoi (Theobald) Non-native[30], JEV[54], Aedes (Finlaya) togoi (Theobald)* Brugia malayi[54], W. bancrofti[54], Ochlerotatus (Finlaya) togoi (Theobald) D. immitis[54] Tanakius togoi (Theobald) Aedes (Zavortinkius) longipalpis (Gruenberg) UGSV Aedes (Finlaya) longipalpis (Gruenberg)* Ochlerotatus (Finlaya) longipalpis (Gruenberg) Zavortinkius longipalpis (Gruenberg) Armigeres (Armigeres) obturbans (Walker)* GETV[39], Kadipiro[39] (Continued)

PLOS ONE | DOI:10.1371/journal.pone.0133602 July 30, 2015 9/26 Stable Mosquito Classification

Table 1. (Continued)

SPECIES ASSOCIATED HUMAN PATHOGENS / INVASIVE AND NON-NATIVE SPECIES Armigeres (Armigeres) subalbatus (Coquillett)* JEV[55], W. bancrofti[54] Haemagogus (Haemagogus) janthinomys Dyar* YFV[56] Haemagogus (Haemagogus) mesodentatus Komp & YFV[55] Kumm* Haemagogus (Haemagogus) spegazzinii Brethes* YFV[55] Haemogogus (Conopostegus) leucocelanus (Dyar & WYOV, YFV Shannon)* Psorophora (Janthinsoma) cyanescens (Coquillett)* VEEV Psorophora (Janthinsoma) ferox (von Humboldt)* CVV[31], EEEV[12], ORIV, SLEV, WNV[14], WYOV Psorophora (Grabhamia) columbiae (Dyar & Knab)* WNV[14], Potosi[22] Psorophora (Grabhamia) conﬁnnis (Lynch Arribálzaga) CVV, TENV, VEEV * Psorophora (Grabhamia) discolor (Coquillett)* VEEV Psorophora (Janthinosoma) ferox (von Humboldt)* WNV[14] Psorophora (Janthinosoma) albipes (Theobald)* WYOV Psorophora (Psorophora) ciliata (Fabricius)* TENV[22], VEEV, WNV[14] Psorophora (Psorophora) cilipes (Fabricius)* VEEV Psorophora (Psorophora) howardii Coquillett* WNV[14]

*Name combination as in Knight & Stone (1977) [1] + Experimental infection ++ Virus detected in eggs +++ Virus detected in males Virus-speciesassociationslistedinboldtypefacearethoselistedontheCDCArbovirusCatalog(2015)[57]. Virus names and abbreviations: BAMV—Barmah Forest; BATV—Batai; BBKV—Babanki; BOUV—Bouboui; BUNV—Bunyamwera; CHIKV—Chikungunya; CVV—Cache Valley; DENV—Dengue; EEEV—Eastern Equine Encephalitis; EHV—Edge Hill; EYEV—Everglades; GETV—Getah; GGV—Gan Gan; HJV—Highland J; ILHV—Ilheus; JCV—James Canyon; KEDV—Kedougou; LACV—La Crosse; LEBV—Lebombo; MVEV— Murray Valley River; NDOV—Nyando; NRIV—Ngari; ORIV—Oriboca; ORUV—Orungo; PGAV—Pongola; RRV—Ross River; RVFV—Rift Valley Fever; SFV—Semiliki Forest; SINV—Sindbis; SLEV—St Louis Encephalitis; SPOV—Spondweri; TAHV—Tahyna; TENV—Tensaw; TVTV—Trivittatus; UGSV—Uganda S; VEEV—Venezuelan Equine Encephalitis; WNV—West Nile; WSLV—Wesselbron; WYOV—Wyeomyia; YFV —Yellow Fever; ZIKAV—Zika.

doi:10.1371/journal.pone.0133602.t001

genus was followed by some workers but ignored by others [68,69]. The recognition of these additional genera and subgenera in Aedini was based on traditional taxonomic judgments of similarities and relationships rather than on quantitative cladistic analysis. In a series of papers published from 2004 through 2009 [70–73], Reinert, Harbach, and Kitching (henceforth RH&K) made many changes to the nomenclature of the tribe Aedini. The RH&K reclassifications [70–73] were based on comprehensive morphological studies of all life stages of the tribe. Their papers report many entirely new morphological observations, especially on the female genitalia. The authors were not able to examine every species in the tribe, but instead chose those species that were available to them and that they considered representative of previously established groups, usually choosing the type species from a genus or subgenus. Based on their results, RH&K identified monophyletic groups and declared new

PLOS ONE | DOI:10.1371/journal.pone.0133602 July 30, 2015 10 / 26 Stable Mosquito Classification

Fig 1. Graph comparing number of aedine mosquito genera and subgenera from 1977 to present. doi:10.1371/journal.pone.0133602.g001

classifications that were further refined in each publication. The series of papers established 74 genera (S1 Fig, Fig 1) from what was formerly genus Aedes, essentially tripling the number of genera in the family Culicidae. Most of the decisions on classification, particularly in the 2009 paper [73], were based on quantitative cladistic analyses. These decisions were primarily anchored to an objectively defined measure of "Groups present/Contradicted" (GC) [74]. Those groups were identified by pseudo-replication, or jackknifing, using symmetric resampling in an implied-weights parsimony analysis in the computer program TNT [75]. When a grouping was supported by a GC threshold 40, it was considered cladistically significant [73]. However, the authors departed from this quantitative scheme by recognizing clades with GC<40 when the “number and nature of the characters that support the branches” were sub- jectively considered significant (Fig 2). RH&K did not propose diagnostic characters for the new genera, though they did assign many species in Aedini to genus or subgenus based on accepted groupings and their own analyses of characters. Exceptions were: genus Ochlerotatus, 138 species unassigned to subgenus; ‘Ochlerotatus’ (‘Finlaya’) sensu auctorum (= a name used by subsequent authors in a sense different from that established by the original author [76]), 28 species; ‘Ochlerotatus’ (‘Protomacleaya’) sensu auctorum, 39 species; and genus Stegomyia, 104 species unassigned to subgenus. Although phylogenetic studies of DNA sequences should ultimately be helpful in determin- ing taxonomic relationships between groups within Aedini, the few studies to date have not been able to test the validity of the reclassification proposed by RH&K based on morphological

PLOS ONE | DOI:10.1371/journal.pone.0133602 July 30, 2015 11 / 26 Stable Mosquito Classification

Fig 2. Tree derived from the single most parsimonious cladogram in Reinert, Harbach & Kitching 2009 [73]. Branches with GC40 are indicated by dark circles. doi:10.1371/journal.pone.0133602.g002

PLOS ONE | DOI:10.1371/journal.pone.0133602 July 30, 2015 12 / 26 Stable Mosquito Classification

characters. DNA barcoding studies have examined 217 aedine species in China [77], India [78], and Afghanistan [79]. Of note, barcoding involves comparison of short sequences of the COI mitochondrial gene, producing distance measures that are only useful for delineating and differentiating species-level taxa, not for estimating their relationships. Combining COI and COII mitochondrial genes from 17 aedine species in western Africa [80] failed to distinguish them as a group from three non-aedine species and also failed to show significant groupings of species. Restriction mapping of rDNA [81] was similarly unsuccessful for a study of 12 culicine species representing six genera. Analysis of the white gene [82] successfully distinguished three aedine species from nine others, but the sample size was too small to examine relationships among the aedine species. The Linnaean system of nomenclature that has served biology since 1758 was formalized in the International Code of Zoological Nomenclature (ICZN), a basic set of rules designed to promote stability and communication [76]. Classifications governed by the ICZN are meant to be “natural,” a term that since the translation of Hennig’s “Phylogenetic Systematics” [83]has been defined as strictly monophyletic, meaning “including all descendants from a common ancestor.” Wiley [84,85], one of the early theorists of phylogenetic systematics, asserted that an annotated Linnaean classification can take more than one form as long as each is logically con- sistent with a phylogeny, and that “...minimum taxonomic decisions will be made whenever possible to construct a classification or to modify an existing classification.” Wiley and Lieber- mann [86] also stated that “the ranks of well-known clades will be retained whenever possible.” Although explicitly "tree-based" classification systems have been suggested as alternatives to Linnaean classification (e.g., numerical prefix schemes [87] and the PhyloCode [88]), these alternative schemes require a well-supported phylogeny based on multiple lines of evidence, a condition not yet reached in Aedini as discussed above. Regardless of the system, some accommodations are necessary for serving the sometimes non-overlapping requirements of taxonomy, information retrieval, and communication [86]. Recently, Vences and colleagues [89] discussed the relationship between nomenclatorial utility and phylogenetic accuracy in an expansion on previous treatments of the subject [84–86]. They acknowledged that taxonomic instability “...can become a serious problem consuming public and private resources...”. To address this and other issues, they proposed several levels of taxon naming criteria (TNCs), the priority criteria being monophyly, clade stability, and phenotypic diagnosability. Accessory criteria included, among others, the medical or economic significance of a taxon. These authors emphasized that “...except for monophyly, the priority TNCs are not proposed as mandatory requirements of a Linnaean taxon but as yardsticks to allow for an informed choice among various clades in a tree...” and warn that “...taxa of unstable monophyly or poor diagnosability reduce the information content and hence the utility of the Linnaean system.” After reviewing the history of the classification of tribe Aedini and previous morphological and molecular phylogenetic analyses, we reanalyzed RH&K data and performed a critical assessment of their methods and results in the context of Wiley [84,85], Wiley and Lieberman [86]’s, “conventions for annotated Linnaean classifications”, and also Vences et al. [89]’s, “taxon naming criteria”. To preserve information about morphological evolution in mosquitoes, we suggest alternative methods for reflecting RH&K’s findings in a classification that retains the stability necessary for effective information retrieval and communication.

Materials and Methods We reanalyzed RH&K’s data in the program TNT [75,90], not for the purpose of producing a classification, but rather to test the robustness of RH&K’s[73] generic groups (Fig 3; S1 Tree,

PLOS ONE | DOI:10.1371/journal.pone.0133602 July 30, 2015 13 / 26 Stable Mosquito Classification

PLOS ONE | DOI:10.1371/journal.pone.0133602 July 30, 2015 14 / 26 Stable Mosquito Classification

Fig 3. Strict consensus tree derived from the reanalysis of the Aedini morphology data set using unweighted parsimony with 14 ordered characters. Dark circle = nodes supported in [73] and herein; triangle = supported in RH&K [73] but not herein; square = supported herein but not in RH&K [73]. doi:10.1371/journal.pone.0133602.g003

S2 Tree, S3 Tree, S4 Tree) under alternative, but similar, cladistic analyses. We assumed that if proposed relationships among genera are not robust when subjected to closely related analytical methods, they are unlikely to be robust to the introduction of new data. To gauge the stability of RH&K’s[73] generic groupings to standard parsimony-based analytical methods, we reanalyzed their data with two alternative methods: (i) unweighted parsimony jackknife analyses, identifying clades and their GC (Groups present/Contradicted) support values [74] on the resulting jackknife tree, and (ii) unweighted maximum-parsimony analyses, identifying clades occurring on the strict consensus of the resulting most parsimonious trees. For each of these analytical methods, we conducted two sets of multiple analyses, one in which all characters were unordered and one in which 14 characters were ordered, as was the case in a subset of the original analyses [73]. We performed all analyses in TNT v.1.1 [75,90], although in some cases we carried out tree-bisection and reconnection (TBR) branch-swapping on accumulated trees in PAUPÃ [91]. Jackknife analyses. We conducted unweighted parsimony jackknife analyses consisting of 1,000 pseudoreplicates, each pseudoreplicate consisting of 10 separate tree searches utilizing TBR branch-swapping on a random-taxon-addition Wagner tree and limited to finding 10 most-parsimonious trees. Support for clades was measured with the GC (Groups present/Con- tradicted) criterion [74] based on the TNT default tree output, which displays a tree consisting of all groups with positive GC values (S1 Tree, S2 Tree). Following RH&K’s criterion, we report nodes with GC 40. Maximum parsimony analyses. We conducted multiple, progressive, unweighted parsimony analyses in which resulting trees were saved, accumulated trees were subjected to TBR branch- swapping, and duplicate trees were discarded. These analyses employed varying combinations of TNT "new technology" algorithms, including the parsimony ratchet (5 to 100,000 iterations), sectorial searching (RSS and CSS), tree drifting (10–1000 cycles), and tree fusing (5–100 rounds), in which accumulating trees from prior searches were carried over into subsequent searches. Multiple driven searches were also conducted, including (i) 1,000 initial random- taxon-addition sequences with five rounds of ratcheting and three rounds of tree-fusing and requiring that minimum-length trees be found 10 times; and (ii) combined sectorial, ratchet, tree-drifting, and tree-fusion searches requiring the consensus tree to stabilize four times (S3 Tree, S4 Tree). GC support criterion. GC is calculated as the difference between the frequency with which a given group is retrieved (e.g., in the jackknife replicates) and the frequency with which its most frequent contradictory group is retrieved [74]. Absolute frequencies (the usual method of counting frequencies in jackknife or bootstrap analyses) do not distinguish between a group with a frequency of 60% that is uncontradicted by an alternative grouping (GC = 60) and a group with a frequency of 60% that is contradicted by an alternative grouping that may occur, e.g., 20% of the time, which would produce GC = 40 [74]. Ordered characters. Following RH&K, the 14 ordered characters (out of a total of 336 characters) are all measurement or ratio characters, ordered as suggested by Thiele [92] (characters 13, 14, 17, 20, 62, 67, 72, 84, 107, 138, 161, 273, 280, 318; for TNT characters 12, 13, 16, 19, 61, 66, 71, 83, 106, 137, 160, 272, 279, 317).

PLOS ONE | DOI:10.1371/journal.pone.0133602 July 30, 2015 15 / 26 Stable Mosquito Classification

Results A series of 20 maximum-parsimony searches in which all characters were unordered converged multiple times on the same set of 832 most-parsimonious trees of unweighted length = 6,609 (S4 Tree). A series of 51 maximum-parsimony searches in which 14 characters were ordered produced 4,000 most-parsimonious trees of unweighted length = 6,654 (S3 Tree), but searches continued to find additional trees, so it is likely that not all of the most-parsimonious trees were found. Finding all most-parsimonious trees, however, is not necessary for calculating the strict consensus [93]. The majority of tree islands were likely visited and the strict consensus is likely stable because all 51 searches, including the driven consensus-stabilization searches, employed the parsimony ratchet [94]. At best, any additional most-parsimonious trees would have no effect on the consensus and, at worst, they would further erode its resolution. Regardless of the specific analysis used (unweighted parsimony or strict consensus, ordered or unordered: see S1 Tree, S2 Tree, S3 Tree, S4 Tree), 48 supported clades, including all 10 pre- 2000 Aedine genera, were supported by GC values 40 (Fig 3). These included 45 of the 46 clades proposed by RH&K, with the exception of the proposed genus Catageiomyia that split into two unsupported clades in our analysis (Fig 3). In addition, our analysis supports three clades corresponding to the proposed genera Danielsia, Hopkinsius and Petermattinglyius (Fig 3), which were not supported in the analysis of RH&K (Fig 1 in [73]). Significantly, our analyses split Aedimorphus into four distinct clades, none of which were supported by GC values 40 (Fig 3). Numerous instances of species groupings unsupported according to RH&K’s criteria but nonetheless ultimately elevated to generic status [73] were also not supported in our analyses (S1 Tree, S2 Tree, S3 Tree, S4 Tree, Fig 3). Most importantly, we found minimal support for the phylogenetic relationships among taxa (i.e., very low GC or jackknife support at branches), underscoring the low phylogenetic resolution of the character set used to assign ranks among the taxa. Detailed examples are discussed fully in the next section.

Discussion As we would expect, the implied-weight parsimony method used by RH&K [73] resulted in a more resolved phylogeny (Fig 2) than the standard equally weighted parsimony strategy we employed (Fig 3). In particular, and as admitted by the authors [73], the high level of precision of the floating-point arithmetic used to calculate fit in the TNT implementation of the implied- weights method is unlikely to find more than a single optimal cladogram. A single-solution, highly resolved cladogram may serve a valuable purpose in posing bold hypotheses maximally vulnerable to refutation by the addition of new data (i.e., taxa or characters) [95]. However, the potential instability that makes monophyletic groups on optimal trees ideal for the generation of bold hypotheses can make those same groups poor candidates upon which to base stable classifications. Because they recognized this problem, RH&K [73] utilized the support criterion of "Groups present/Contradicted" (GC) 40, based on jackknifing with symmetric resampling, to judge clades on their implied-weights parsimony tree (Fig 2). Although we applied this same criterion (i.e., GC 40) in our alternative unweighted parsimony analyses, we found negligible support for the basal branches uniting the terminal taxa, i.e., for higher-level relationships between supraspecific groups (Fig 3). Without a better understanding of the relationships among terminal taxa, application of the principle of equivalent ranking advocated by RH&K is unfeasible. In light of these results and using Vences et al. [89] as a guide, we make a point-by-point evaluation to assess whether the classification of Aedini as proposed by RH&K [73] meets the taxonomic naming criteria (TNCs) described. Of 11 TNCs discussed by Vences et al. [89] the

PLOS ONE | DOI:10.1371/journal.pone.0133602 July 30, 2015 16 / 26 Stable Mosquito Classification

following seven are applicable to our argument (quotes are from Vences et al. [89] unless otherwise noted).

TNC 1: Monophyly “The only strict TNC, [...]: All supraspecific taxa must be monophyletic units in the respective species tree (although they might be paraphyletic in any gene tree). Monophyly of a taxon should be assessed by an explicit phylogenetic analysis with adequately dense taxon sampling.” Monophyly of family Culicidae has been demonstrated [96–98] and Tribe Aedini is monophyletic based on morphological characters [99], egg structure, and physiology [67]. RH&K propose and at least partially demonstrate monophyly for many aedine generic level taxa [73]. However, questions remain about branch support and subjectivity, particularly the use of the GC value as the primary determinant of branch support and the arbitrary use of GC40 as the cut-off for significance. Of the 74 terminal clades raised to genera by RH&K [73], 36 are supported according to the GC40 criterion. Elevation of the remaining 38 unsupported clades (GC<40) to generic status is not further explained or justified except to reference the “nature of the characters that support the branches.” RH&K [73] indicate three measures of support on each branch of their most-parsimonious implied-weights tree (Fig 1 in [73]): (i) the absolute (i.e., non-GC) frequency from symmetric resampling (a form of jackknifing that accounts for character weights; see [74] and [90]), (ii) the GC frequency from symmetric resampling, and (iii) the absolute (i.e., non-GC) frequency based on typical (i.e., non-symmetric) jackknife resampling. On some branches, these support measures take on zero (measure i) or negative (measures ii and iii) values, indicating no support or contradictory support for groups that do not appear on the most-parsimonious tree. Except for one case (the clade Juppius + Lepidokenon), in discussing their choices to elevate groups of species to genus level, RH&K only consider measure ii and ignore the other two measures. Critically, support (GC40) uniting pairs of genera (i.e. putative sister taxa) was limited to two nodes in RH&K’s 2009 analysis [73](Downsiomyia–Haemagogus and Elpeytonius-Cata- geiomyia), and more basal support (GC40) for relationships between multiple genera occurs only twice (a clade with four genera (Sallumia, Geoskusea, Levua, Rhinoskusea, and the entire clade from Elpeytonius to Stegomyia). However, when GC support values are taken into account, their tree is essentially a polytomy with regard to relationships between species groups (Fig 1 in [73]). Our simplified RH&K tree (Fig 2) does not show the internal relationships within Mucidus, Ochlerotatus, Petermattinglyius, and Stegomyia as proposed by RH&K (Fig 1 in [73]). Genus Mucidus had two well-supported subgenera and genus Ochlerotatus was divided into 14 subgenera, nine of which were supported (Pseudoskusea, Woodius, Culicada, Gilesia, Ochlerotatus, Protoculex, Chrysoconops, Pholeomyia, Rusticoidus). The branch leading to Ochlerotatus subgenera Juppius and Leptokeneon only has single species scored thus it was not possible to judge their support. Interspersed with the named Ochlerotatus subgenera were 124 (of 198) species unassigned to subgenus (Appendix 2 in [73]). Lack of consistency in naming within the Ochler- otatus clade was evident. For example, it was unclear why subgenus Culicelsa was elevated from synonymy [72] while the genus Ochlerotatus species pair andersoni and nivalis, with similarly poor support, was not given equivalent subgeneric rank [73]. Genus Petermattinglyius was split into two subgenera even though only the nominotypical subgenus was supported by GC40. Genus Stegomyia, containing several medically important species, emerged as two poorly supported clades. Eight well-supported subgenera were created for 30 species of Stego- myia, leaving the remaining 100 species unassigned to subgenus. We also noted that genus

PLOS ONE | DOI:10.1371/journal.pone.0133602 July 30, 2015 17 / 26 Stable Mosquito Classification

Aedimorphus, was not divided into subgenera by RH&K even though it contains four branches with GC40 support [73]. The above hypotheses of monophyly are accepted by us as the best currently available. Given our own results and a reassessment of those presented in RH&K [73], we follow Vences et al. [89], who proposed that subgenera or species group names be used “before the phylogeny is fully resolved” or “when important future changes to classification are to be expected.” Since the monophyly of groups within Aedini is only partially supported, we argue it serves little purpose to subdivide the well-known taxon Aedes into many additional generic level groups. It is preferable at this time to preserve the groups as subgenera or informal groups.

TNC 2: Clade Stability “Those clades selected for naming as taxa in a phylogeny should be supported by as many different independent data sets and analysis methods as possible, and not strongly contradicted by any of them (strong / significant contradictions require a biological explanation to be put forward).” This criterion is not met by RH&K since only one data set and one analytical approach (implied-weights parsimony) was considered. Our analyses of the same data set with a closely related method, jackknife unweighted parsimony jackknifing, produced significant support (i.e., by RH&K's criterion of GC40) for just over half the terminal clades assumed to be equivalent-rank genera by RH&K. This lack of support for terminal species-group clades, as well as the non-significant support for internal nodes in both RH&K's and our analyses, demonstrates the weakness of the phylogenetic signal in the dataset used by RH&K to infer relationships among taxa upon which they base their taxonomic ranks.

TNC 3: Phenotypic Diagnosability “A taxon to which a Linnaean rank is assigned should be diagnosable and identifiable pheno- typically. Preference should be on diagnostic characters that are unequivocally synapomorphic, externally visible in as many sexes and life-history stages of the organism as possible, and rec- ognizable also by non-specialists;.....” For this we also follow Wiley & Liebermann’s[86] description of diagnosis which is: “Diagnoses in revisionary work have a different function than diagnoses used for conveying phylogenetic characters of monophyletic groups.” None of the new or resurrected generic level taxa [70–73] were accompanied by diagnoses or keys. They are characterized instead by lists of homoplasious polythetic characters that are not easily retrievable from the publications [70–73].

TNC 8: Manageability “If equally stable and diagnosable clades are alternatively available for naming as taxa, choose those that contain a number of taxa manageable for the human mind. Avoid over splitting and deliberately creating monotypic taxa.” The avoidance of over splitting applies to diagnosable clades and multiple data sets, which as discussed above fundamentally does not apply. However, we cite TNC 8 because we argue that, based on the available dataset, the classification has been over split and most names are difficult to remember and manage.

PLOS ONE | DOI:10.1371/journal.pone.0133602 July 30, 2015 18 / 26 Stable Mosquito Classification

TNC 9: Hall of Fame “Take particular care with taxa that are of high public interest beyond taxonomy and where communication is thus particularly important. The more prominent a rank the more carefully should any change be applied. Intermediate ranks or unranked taxon names can be used pref- erentially when Clade Stability and Phenotypic Diagnosability TNCs are not sufficiently met.” We show 101 high-profile mosquito species in tribe Aedini, 85 formerly in genus Aedes (Table 1). Given the importance of the relationship of classification to communication and information retrieval, this alone is reason to use intermediate ranks or informal groups to reflect hypotheses of relationships, at least until the classification is declared stable.

TNC 10: Nomenclatural Stability “If equally stable and diagnosable clades are alternatively available for naming as taxa, avoid a classification in which unstable names .... are resurrected from synonymy.” Also, “Minimum taxonomic decisions will be made, whenever possible, to construct a classification or to modify an existing classification.” Although this TNC is for more than one alternative clade we apply it to the excessive changes proposed for genus Aedes between 2000 and 2009. S1 Fig is a summary of currently valid generic-level names with a chronology of changes from 1977–2015. The dominant start- ing point for the chronology was the 1977 catalog [1]. As a result of Reinert [66] and RH&K [70–73] there have been 118 status and / or rank changes in the original genus Aedes (elevation from synonymy, change from subgenus to genus and vice versa, and new names). Several taxonomic changes were proposed and later amended or reversed in subsequent publications e.g., Kenknightia (3 changes), Levua (4 changes with a reversal), Pseudoskusea (3 changes with a reversal), and Rhinoskusea (3 changes with a reversal) (S1 Fig). These changes underscore the instability of this proposed classification.

TNC 11: Community Consensus “If equally stable and diagnosable clades are alternatively available for naming as taxa, choose a classification which is favored by the majority of taxonomists and, if applicable, other biolo- gists, e.g., because it conserves the traditional content and definition of prominent taxa, ...” We measured community consensus regarding the proposed new classifications of Aedini by RH&K [70–73] by using the number of “hits” in the “Web of Science,” searching “topic” AND “title” on 13 September, 2014. The time reference was from 2004 (elevation of genus Ste- gomyia) to present. We chose a few important species to determine which names researchers were using. Search terms and hits are as follow: “Aedes aegypti” (n = 6,300) versus “Stegomyia aegypti” (n = 191); “Aedes albopictus” (n = 2,042) versus “Stegomyia albopicta” (n = 23); “Aedes japonicus” (n = 102) versus “Ochlerotatus japonicus” (n = 72) and “Hulecoeteomyia japonica” (n = 0). The “traditional” use of Aedes for this small but important sample of species is obvi- ously preferred by the majority of workers. It is likely that “Ochlerotatus japonicus”, a recent introduction to North America [100], received more use because it was adopted in a revision of the influential Darsie & Ward key to North American mosquitoes [101]. RH&K’s work, published from 2004 through 2009 [70–73], caused a dramatic change in the structure of Aedini, as outlined above. The elevation of Ochlerotatus to genus level in 2000 [66] removed many familiar species from Aedes, which in North America involved most of the floodwater species that formed a functional unit from the standpoint of operational mosquito control. In that sense, Ochlerotatus was useful from a functional perspective, even if it meant that morphological keys could not be used to separate them at a generic level (i.e., identifica- tions were performed at a species level and then assigned to either Ochlerotatus or Aedes

PLOS ONE | DOI:10.1371/journal.pone.0133602 July 30, 2015 19 / 26 Stable Mosquito Classification

retrospectively [67]). The establishment of 74 additional genera by RH&K [70–73](S1 Fig, Fig 1) removed that functionality. RH&K stated that they were attempting to make divisions within Aedini comparable in taxonomic significance by elevating each monophyletic group to genus rank. They used, but did not define or reference, the “principle of equivalent rank” as one of the bases of this decision. While Hennig [83] introduces and discusses many criteria that should be used in the absolute ranking of higher taxa, such as assigning equal rank to sister taxa, he does not specifically define or highlight a principle of equivalent rank. Application of the “principle of equivalent rank” sensu RH&K resulted in many new taxon groups being recognized by those authors as equivalent to the small number of traditionally recognized genera, and therefore raised by them to genus status. The most significant outcomes of this process were the near trebling of the number of total genera and the reduction of the number of species in genus Aedes from over 800 to just 12. These proposed changes have caused extensive debate about translating phylogenies into classifications [102–105]. The attempt to align aedine classification with cladistically defined monophyletic groups was probably motivated by a desire to have a natural classification. Such a classification has the potential to make information retrieval easier, in that identifying a particular specimen or pop- ulation to genus would associate it with biologically significant characteristics shared by species within that genus. Such a classification could be helpful not only by predicting likely characters of unfamiliar species, but also by serving as a powerful tool for educating people about the structure of the larger groups. Unfortunately, in our experience, many of these advantages have not resulted from the new classification because most of the genera cannot be identified except by combinations of character states, i.e., they do not have simple, unreversed diagnostic character states. This problem not only inhibits the assignment of new species to the genera, it also tends to force entomologists to identify specimens to species before assigning them to a genus. In contrast, the traditional genus Aedes has morphological and biological characters held in common by its species [67], so that identification of a specimen to Aedes is informative. The large number of species in the genus is not in itself a reason to divide it, since it may be the result of an evolutionary radiation of a cohesive genetic architecture that is particularly success- ful under current conditions (for examples refer to [106]). Hindrance of communication and information retrieval resulting from the reclassification of Aedini is illustrated by the successive renaming of Aedes (Finlaya) japonicus, a recent invasive species in the US and elsewhere [100]. It was first changed to Ochlerotatus japonicus [66] and then Hulecoeteomyia japonica [71](Table 1). The two most globally important arboviral vectors Aedes (Stegomyia) aegypti and Ae.(Stegomyia) albopictus were renamed Stegomyia aegypti and Stegomyia albopicta, respectively [70](Table 1). Another example is the invasive Australasian species Aedes notoscriptus (Skuse), recently found in California (Kenn Fujioka, Pers. Comm.). During the past decade, this species has been referred to as Aedes (Finlaya) notoscriptus [1], Ochlerotatus (Finlaya) notoscriptus [70], and Rampamyia notoscripta [71] (Table 1). Successive name changes during a short period of time are confusing and significantly interfere with critical information retrieval from the literature (Table 1). In conclusion, we formally restore the generic classification of Aedini to its status prior to the publications of RH&K (S1 Appendix), update the subgenera and informal species groups in genus Aedes in accordance with their findings (S1 Appendix, S2 Appendix), and propose updated two and three-letter abbreviations for genera and subgenera, respectively (S3 Appen- dix). We argue that this action will stabilize the Aedini classification and maximize its useful- ness to the operational community, while highlighting the need to strive towards a natural system where group membership will provide insights into ecological, evolutionary, and epidemiological criteria. Basing subgenera and informal species groups on the results of RH&K's morphological research incorporates the progress that they have made in defining clades. In

PLOS ONE | DOI:10.1371/journal.pone.0133602 July 30, 2015 20 / 26 Stable Mosquito Classification

taking these actions we restore the utility of the generic designations within Aedini while safe- guarding advances in our understanding of morphological evolution in mosquitoes. Specifically, we propose the following: 1. Return to a linear classification that retains names as they were commonly used prior to the year 2000. This arbitrarily retains the recent elevation of the morphologically diagnosable genus Verrallina (S1 Appendix, S1 Fig). 2. Reduction in rank of all aedine genera created in and after 2000 to subgenera of Aedes (S1 Appendix, S1 Fig). Reduction of all subgenera designated by RH&K to informal species- group status (S2 Appendix). This respects the taxonomic conclusions of RH&K [73]. 3. Acceptance of a revised list of species (S1 Appendix; abbreviations in S3 Appendix) with appropriate endings, authors, and dates to reflect membership in the genus Aedes.

Supporting Information S1 Appendix. Tribe Aedini generic, subgeneric and species name combinations proposed herein. (PDF) S2 Appendix. Informal group names proposed here for RH&K [73] post-1999 subgenera. (PDF) S3 Appendix. Mosquito generic (two letters) and subgeneric (three letters) abbreviations. Based on: Reinert JF (2009) List of abbreviations for currently valid generic-level taxa in family Culicidae (Diptera). European Mosquito Bulletin 27: 68–76. (PDF) S1 Fig. Timeline illustration of changes in the classification of Aedini from 1977 to present. Valid generic level names and graphical timeline of nomenclature combination changes in Aedine taxa pre-2000 to date. Genera are indicated in blue, subgenera in green and synonomy (Syn.) in red. Author abbreviations include: D&S = Dyar and Shannon; H&R = Harbach & Rattanarithikul; R-D = Robineau-Desvoidy RH&K = Reinert, Harbach & Kitching; S&P = Shevchenko & Prudkina. Generic abbreviations follow those in Reinert, Harbach & Kitching (2009) and herein as listed in S3 Appendix. ÃNyctomyia is a replacement name for the pre- occupied genus name Nyx (Harbach et al., Zootaxa, 3683(2), 159–177 (2013). (PDF) S1 Tree. Unweighted parsimony jackknife analysis of the Aedini morphology dataset with 14 characters ordered. (PDF) S2 Tree. Unweighted parsimony jackknife analysis of the Aedini morphology dataset with 14 characters unordered. (PDF) S3 Tree. Unweighted parsimony analysis of the Aedini morphology dataset identifying clades occurring on the strict consensus of the resulting most parsimonious trees with 14 characters ordered. (PDF)

PLOS ONE | DOI:10.1371/journal.pone.0133602 July 30, 2015 21 / 26 Stable Mosquito Classification

S4 Tree. Unweighted parsimony analysis of the Aedini morphology dataset identifying clades occurring on the strict consensus of the resulting most parsimonious trees with 14 characters unordered. (PDF)

Acknowledgments We thank Dave Swofford, Ed Wiley, Torsten Dikow, Rudy Meiswinkel and Art Borkent for enlightened and informative discussions on technical, basic, and applied aspects of this study that significantly improved the manuscript. We also thank Judith Stoffer for excellent graphics support. This research was performed under a Memorandum of Understanding between the Walter Reed Army Institute of Research and the Smithsonian Institution, with institutional support provided by both organizations. This manuscript was prepared whilst YML held a National Research Council (NRC) Research Associateship Award at the Walter Reed Army Institute of Research. The material to be published reflects the views of the authors and should not be construed to represent those of the US Department of the Army or the US Department of Defense.

Author Contributions Conceived and designed the experiments: TRS RCW YML. Performed the experiments: TRS. Analyzed the data: TRS RCW YML. Wrote the paper: RCW YML DMF TRS DAS DCP. Initi- ated this study: RCW DCP DAS YML.

References 1. Knight KL, Stone A (1977) A catalog of the mosquitoes of the world (Diptera: Culicidae). College Park, Maryland: Entomological Society of America. 611 p. 2. Knight KL (1978) Supplement to a catalog of the mosquitoes of the world (Diptera: Culicidae). College Park, Maryland: Entomological Society of America. 107 p. 3. Ward RA (1984) Second supplement to "A catalog of the mosquitoes of the world" (Diptera: Culici- dae). Mosq Syst 16: 227–270. 4. Gaffigan TV, Ward RA (1985) Index to the second supplement to " A catalog of the mosquitoes of the world", with corrections and additions (Diptera: Culicidae). Mosq Syst 17: 52–63. 5. Ward RA (1992) Third supplement to "A catalog of the mosquitoes of the world" (Diptera: Culicidae). Mosq Syst 24: 177–230. 6. Stone A, Knight KL, Starcke H (1959) A synoptic catalog of the mosquitoes of the world (Diptera, Culi- cidae). College Park, Maryland: Entomological Society of America. 358 p. 7. Stone A (1961) A synoptic catalog of the mosquitoes of the world, supplement I (Diptera: Culicidae). Proc Entomol Soc Wash 63: 29–52. 8. Stone A (1963) A synoptic catalog of the mosquitoes of the world, suppl. II (Diptera: Culicidae). Proc Entomol Soc Wash 65: 117–140. 9. Stone A (1967) A synoptic catalog of the mosquitoes of the world, suppl. III (Diptera: Culicidae). Proc Entomol Soc Wash 69: 197–224. 10. Stone A (1970) A synoptic catalog of the mosquitoes of the world, suppl. IV (Diptera: Culicidae). Proc Entomol Soc Wash 72: 137–171. 11. Stone A (1971) Corrections to mosquito catalog supplements III and IV. Proc Entomol Soc Wash 73: 180. 12. Armstrong PM, Andreadis TG (2010) Eastern equine encephalitis virus in mosquitoes and their role as bridge vectors. Emerg Inf Dis 16: 1869–1874. PMID: 21122215. PMCID: PMC3294553. doi: 10. 3201/eid1612.100640 13. Andreadis T, Capotosto P, Shope R, Tirrell S (1994) Mosquito and arbovirus surveillance in Connecti- cut, 1991–1992. J Am Mosq Control Assoc 10: 556–564. PMID: 7707064.

PLOS ONE | DOI:10.1371/journal.pone.0133602 July 30, 2015 22 / 26 Stable Mosquito Classification

14. CDC (2012) Mosquito species in which West Nile virus has been detected, United States,1999–2012 (reported to ArboNET) (http://www.cdc.gov/westnile/resources/pdfs/MosquitoSpecies1999-2012.pdf, accessed 02/20/2015). Atlanta, GA: Centers for Disease Control and Prevention. 15. Fontenille D, Traore-Lamizana M, Diallo M, Thonnon J, Digoutte J, Zeller H (1998) New vectors of Rift Valley fever in West Africa. Emerg Infect Dis 4: 289–293. PMID: 9621201. PMCID: PMC2640145. doi: 10.3201/eid0402.980218 16. McIntosh B (1972) Rift Valley fever. 1. Vector studies in the field. J S Afr Vet Assoc 43: 391–395. PMID: 4670767. 17. Turrell M, Faran M, Cornet M, Bailey C (1988) Vector competence of Sénégalese Aedes fowleri (Dip- tera: Culicidae) for Rift Valley fever virus. J Med Entomol 25: 262–266. PMID: 3404545. doi: 10.1093/ jmedent/25.4.262 18. Gordon SW, Tammariello RF, Linthicum KJ, Dohm DJ, Digoutte JP, Calvo-Wilson MA (1992) Arbovi- rus isolations from mosquitoes collected during 1988 in the Senegal River basin. Amer J Trop Med Hyg 47: 742–748. PMID: 1361722. 19. Turrell MJ, Faran ME, Cornet M, Bailey CL (2005) Vector competence of Sénégalese Aedes fowleri (Diptera: Culicidae) for Rift Valley fever virus. J Med Entomol 25: 262–266. PMID: 3404545. doi: 10. 1093/jmedent/25.4.262 20. Liu H, Li M, Zhai Y, Meng W, Sun X, Cao Y, et al. (2010) Banna virus, China, 1987–2007. Emerg Infect Dis 16: 514–517. PMID: 20202434. PMCID: PMC3322026. doi: 10.3201/eid1603.091160 21. Wang Z, An S, Wang Y, Han Y, Guo J (2009) A new virus of Flavivirus: Chaoyang virus isolated in Liaoning province. Chin Pub Health 25: 769–772. 22. Wozniak A, Dowda H, Tolson M, Karabatsos N, Vaughan D, Turner P, et al. (2001) Arbovirus surveillance in South Carolina, 1996–98. J Amer Mosq Contr Assoc 17: 73–78. PMID: 11345423. 23. Diallo M, Thonnon J, Traore-Lamizana M, Fontenille D (1999) Vectors of Chikungunya virus in Sene- gal: current data and transmission cycles. Amer J Trop Med Hyg 60: 281–286. PMID: 10072152. 24. Germain M, Francy DB, Monath TP, Ferrara L, Bryan J, Salaun JJ, et al. (1980) Yellow Fever in The Gambiae, 1978–1979: Entomological aspects and epidemiological correlations. Amer J Trop Med Hyg 29: 929–940. PMID: 7435794. 25. Gould DJ, Bailey CL, Vongpradist S (1982) Implication of forest mosquitoes in transmission of Wuchereria bancroft in Thailand. Mosq News 42: 560–564. 26. Harinasuta C, Sucharit S, Deesin T, Surathin K, Vutikes S (1970) Bancroftian filariasis in Thailand, a new endemic area. Southeast Asia J Trop Med Pub Hlth 1: 233–245. 27. Belkin JN (1962) The mosquitoes of the South Pacific (Diptera, Culicidae): University of California Press. 608 p. 28. Marks EN (1947) Studies of Queensland mosquitoes. Part I. The Aedes (Finlaya) kochi group with descriptions of new species from Queensland, Bougainville and Fiji. Univ Queensland Papers Dept Biol 2: 1–66. 29. Suguri S, Cabrera BD, Shibuya T, Harada M, Valeza FS, Nagata T, et al. (1985) Vector mosquitoes of Wuchereria bancrofti at Bicol region in the Philippines. 1. Transmission capability. Jap J Exp Med 55: 61–65. PMID: 3900489. 30. Juliano SA, Lounibos LP (2005) Ecology of invasive mosquitoes: effects on resident species and on human health. Ecol Lett 8: 558–574. PMCID: PMC1920178. doi: 10.1111/j.1461-0248.2005.00755.x PMID: 17637849 31. Andreadis TG, Armstrong PM, Anderson JF, Main AJ (2014) Spatial-temporal analysis of Cache Val- ley Virus (Bunyaviridae: Orthobunyavirus) infection in anopheline and culicine mosquitoes (Diptera: Culicidae) in the Northeastern United States, 1997–2012. Vector Borne Zoonotic Dis 14: 763–773. PMID: 25325321 [PubMed—in process] PMCID: PMC4208611 [Available on 2015-10-01]. doi: 10. 1089/vbz.2014.1669 32. Versteirt V, De Clercq EM, Fonseca DM, Pecor J, Schaffner F, Coosemans M, et al. (2012) Bionomics of the established exotic Mosquito species Aedes koreicus in Belgium, Europe. J Med Entomol 49: 1226–1232. PMID: 23270149. doi: 10.1603/me11170 33. McIntosh BM (1971) The aedine subgenus Neomelaniconion Newstead (Culicidae, Diptera) in south- ern Africa with description of two new species. J Ent Soc S Africa: 319–333. 34. McIntosh BM, Jupp PG, dos Santos I, Barnard BJH (1980) Vector studies on Rift Valley fever virus in South Africa. S Afr Med J 58: 127–132. PMID: 6105722. 35. Rodaniche ED, Galindo P (1963) Ecological observations of Ilheus virus in vicinity of Almirante, Republic of Panama. Amer J Trop Med Hyg 12: 924–928. PMID: 14072452.

PLOS ONE | DOI:10.1371/journal.pone.0133602 July 30, 2015 23 / 26 Stable Mosquito Classification

36. Andreadis T, Anderson J, Tirrell-Peck S (1998) Multiple isolations of eastern equine encephalitis and highlands J viruses from mosquitoes (Diptera: Culicidae) during a 1996 epizootic in southeastern Connecticut. J Med Entomol 35: 296–302. PMID: 9615549. doi: 10.1093/jmedent/35.3.296 37. Gargan TP, Clark GG, Dohm DJ, Turell MJ, Bailey CL (1988) Vector potential of selected North Ameri- can mosquito species for Rift Valley fever virus. Amer J Trop Med Hyg 38: 440–446. PMID: 2895591. 38. Ergunay K, Gunay F, Oter K, Kasap OE, Orsten S, Akkutay AZ, et al. (2013) Arboviral surveillance of field-collected mosquitoes reveals circulation of West Nile Virus Lineage 1 strains in Eastern Thrace, Turkey. Vector Borne Zoonotic Dis 13: 744–752. PMID: 23919608. doi: 10.1089/vbz.2012.1288 39. Liu H, Gao X, Liang G (2011) Newly recognized mosquito-associated viruses in mainland China, in the last two decades. Virol J 8: 68. PMID: 21314994. PMCID: PMC3048566. doi: 10.1186/1743- 422X-8-68 40. Carpenter SJ, LaCasse WJ (1955) Mosquitoes of North America (North of Mexico). Univ of California Press. 495 p. 41. Tao S, Zhang H, Yang D, Wang H, Liu Q, Zhang Y, et al. (2003) Investigation of arboviruses in Lan- cang river down-stream area in Yunnan province. Chin J Exp Clin Virol 17: 322–326. PMID: 15340542. 42. Wellings FM, Lewis AL, Pierce LV (1972) Agents encountered during arboviral ecological studies: Tampa Bay area, Florida, 1963–1970. Amer Soc Trop Med Hyg 21: 201–213. PMID: 4400812. 43. Pauvolid-Corrêa A, Kenney JL, Couto-Lima D, Campos ZM, Schatzmayr HG, Nogueira RM, et al. (2013) Ilheus virus isolation in the Pantanal, west-central Brazil. PLoS Negl Trop Dis 7: e2318. PMID: 23875051. PMCID: PMC3715421. doi: 10.1371/journal.pntd.0002318 44. Arnell JH (1976) Mosquito studies (Diptera, Culicidae). XXXIII. A revision of the Scapularis Group of Aedes (Ochlerotatus). Cont Amer Entomol Inst 13: 1–144. 45. Hubálek Z, Halouzka J (1999) West Nile fever–a reemerging mosquito-borne viral disease in Europe. Emerg Infect Dis 5: 643–650. PMCID: PMC2627720. doi: 10.3201/eid0505.990505 PMID: 10511520 46. Mitchell CJ, Haramis LD, Karabatsos N, Smith GC, Starwalt VJ (1998) Isolation of La Crosse, Cache Valley, and Potosi viruses from Aedes mosquitoes (Diptera: Culicidae) collected at used-tire sites in Illinois during 1994–1995. J Med Entomol 35: 573–577. PMID: 9701947. doi: 10.1093/jmedent/35.4. 573 47. Huang Y-M (1990) The subgenus Stegomyia of Aedes in the Afrotropical region. 1. The africanus group of species (Diptera: Culicidae). Contr Amer Entomol Inst 26: 3–90. 48. Pagès F, Peyrefitte CN, Mve MT, Jarjaval F, Brisse S, Iteman I, et al. (2009) Aedes albopictus mosquito: The main vector of the 2007 Chikungunya outbreak in Gabon. PLoS ONE 4. PMID: 19259263. doi: 10.1371/journal.pone.0004691 49. Niebylski ML, Mutebi JP, Craig GBJ, Mulrennan JAJ, Hopkins RS (1992) Eastern equine encephalitis virus associated with Aedes albopictus Florida, 1991. MMWR Morb Mortal Wkly Rep 41: 121. PMID: 1736088. 50. Hurlbut H (1956) West Nile virus infection in arthropods. Am J Trop Med Hyg 5: 76–85. PMID: 13292654 51. Miller BR, Mitchell CJ, Ballinger ME (1989) Replication, tissue tropisms and transmission of yellow fever virus in Aedes albopictus. Trans R Soc Trop Med Hyg 83: 252–255. PMID: 2609379. doi: 10. 1016/0035-9203(89)90667-6 52. Huang Y-M (1986) Aedes (Stegomyia) bromeliae (Diptera: Culicidae), The yellow fever virus vector in east Africa. J Med Entomol 23: 196–200. PMID: 3701804. doi: 10.1093/jmedent/23.2.196 53. Iyengar MOT (1954) Distribution of filariasis in the South Pacific region. South Pacific Commission Technical Paper 66. pp. 52. 54. Tanaka K, Mizusawa K, Saugstad ES (1979) A revision of the adult and larval mosquitoes of Japan (including the Ryukyu Archipelago and the Ogasawara Islands) and Korea (Diptera: Culicidae). Contr Amer Entomol Inst 16: 987. 55. Clements AN (2012) The biology of mosquitoes. Volume 3. Transmission of viruses and interactions with bacteria. Wallingford, England: CABI. 571 p. 56. Arnell JH (1973) Mosquito studies (Diptera, Culicidae). XXXII. A revision of the genus Haemagogus. Contr Amer Entomol Inst 10: 1–174. 57. CDC (2015) Arbovirus Catalog (https://wwwn.cdc.gov/Arbocat/Default.aspx/, accessed 02/2015). Atlanta GA: Centers for Disease Control and Prevention. 58. Edwards FW (1932) Genera Insectorum. Diptera. Fam. Culicidae. Bruxelles, Belgium: Desmet-Ver- teneuil, Imprimeur-Editeur. 258 p.

PLOS ONE | DOI:10.1371/journal.pone.0133602 July 30, 2015 24 / 26 Stable Mosquito Classification

59. Reinert JF (1979) A description of Isoaedes, A new subgenus of Aedes Meigen, and its type-species, Ae.(Isa.) cavaticus new species (Diptera: Culicidae). Mosq Syst 11: 144–162. 60. Reinert JF (1982) Belkinius, a new subgenus of Aedes and a description of Ae.(Blk.) aurotaeniatus Edwards (Diptera: Culicidae). Mosq Syst 14: 101–121. 61. Reinert JF (1987) Albuginosus, a new subgenus of Aedes Meigen (Diptera: Culicidae) described from the Afrotropical region. Mosq Syst 18: 307–326. 62. Reinert JF (1990) Medical entomology studies- XVII. Biosystematics of Kenknightia, A new subgenus of the mosquito genus Aedes Meigen from the oriental region (Diptera: Culicidae). Contr Amer Ento- mol Inst 26: 1–119. 63. Reinert JF (1999) Descriptions of Zavortinkius, A new subgenus of Aedes, and the eleven included species from the Afrotropical region (Diptera: Culicidae). Contr Amer Entomol Inst 31: 1–105. 64. Reinert JF (1999) Restoration of Verrallina to generic rank in tribe Aedini (Diptera: Culicidae) and descriptions of the genus and three included subgenera. Contr Amer Entomol Inst 31: 1–83. 65. Reinert JF (2000) Restoration of Ayurakitia to generic rank in tribe Aedini and a revised definition of the genus. J Am Mosq Control Assoc 16: 57–65. PMID: 10901627 66. Reinert JF (2000) New classification for the composite genus Aedes (Diptera: Culicidae: Aedini), elevation of subgenus Ochlerotatus to generic rank, reclassification of the other subgenera, and notes on certain subgenera and species. J Am Mosq Control Assoc 16: 175–188. PMID: 11081644. 67. Savage HM, Strickman D (2004) The genus and subgenus categories within Culicidae and placement of Ochlerotatus as a subgenus of Aedes. J Am Mosq Control Assoc 20: 208–214. PMID: 15264635. 68. Black W (2004) Learning to use Ochlerotatus is just the beginning. J Am Mosq Control Assoc 20: 215–216. PMID: 15264636. 69. Edman JD (2005) Journal policy on names of aedine mosquito genera and subgenera. J Med Entomol 42: 511. PMID: 16119537. 70. Reinert JF, Harbach RE, Kitching IJ (2004) Phylogeny and classification of Aedini (Diptera: Culicidae), based on morphological characters of all life stages. Zool J Linn Soc 142: 289–368. doi: 10.1111/j. 1096-3642.2004.00144.x 71. Reinert JF, Harbach RE, Kitching IJ (2006) Phylogeny and classification of Finlaya and allied taxa (Diptera: Culicidae: Aedini) based on morphological data from all life stages. Zool J Linn Soc 148: 1– 101. doi: 10.1111/j.1096-3642.2006.00254.x 72. Reinert JF, Harbach RE, Kitching IJ (2008) Phylogeny and classification of Ochlerotatus and allied taxa (Diptera: Culicidae: Aedini) based on morphological data from all life stages. Zool J Linn Soc 153: 29–114. doi: 10.1111/j.1096-3642.2008.00382.x 73. Reinert JF, Harbach RE, Kitching IJ (2009) Phylogeny and classification of tribe Aedini (Diptera: Culi- cidae). Zool J Linn Soc 157: 700–794. doi: 10.1111/j.1096-3642.2009.00570.x 74. Goloboff PA, Farris JS, Kallersjo M, Oxelman B, Ramirez MJ, Szumik A (2003) Improvements to resampling measures of group support. Cladistics 19: 324–332. 75. Goloboff PA (2014) Extended implied weighting. Cladistics 30: 260–272. doi: 10.1111/cla.12047 76. ICZN (1999) International Code of Zoological Nomenclature. London, UK: International Trust for Zoo- logical Nomenclature. 306 p. 77. Wang G, Li C, Guo X, Xing D, Dong Y, Wang Z, et al. (2012) Identifying the main mosquito species in China based on DNA barcoding. PLoS ONE 7: e47051. PMID: 23071708. doi: 10.1371/journal.pone. 0047051 78. Kumar NP, Rajavel AR, Natarjan R, Jambulingam P (2007) DNA barcodes can distinguish species of Indian mosquitoes (Diptera: Culicidae). J Med Entomol 44: 1–7. PMID: 17294914. doi: 10.1093/ jmedent/41.5.01 79. Ashfaq M, Heber PDN, Mirza JH, Khan AM, Zafar Y, Mirza MS (2014) Analyzing mosquito (Diptera: Culicidae) diversity in Pakistan by DNA barcoding. PLoS ONE 9: e97268. PMID: 24827460. PMCID: PMC4036727. doi: 10.1371/journal.pone.0097268 80. Cook S, Diallo M, Sall AA, Cooper A, Holmes EC (2005) Mitochondrial markers for molecular identification of Aedes mosquitoes (Diptera: Culicidae) involved in transmission of arboviral disease in West Africa. J Med Entomol 42: 19–28. PMID: 15691004. doi: 10/1093/jmedent/42.1.19 81. Kumar A, Black WC, Rai KS (1998) An estimate of phylogenetic relationships among culicine mosquitoes using a restriction map of the rDNA cistron. Insect Mol Biol 7: 367–373. PMID: 9723874. doi: 10. 1046/j.1365-2583.1998.740367.x 82. Besansky NJ, Fahey GT (1997) Utility of the white gene in estimating phylogenetic relationships among mosquitoes (Diptera: Culicidae). Mol Biol Evol 14: 442–454. PMID: 9100374. 83. Hennig W (1966) Phylogenetic Systematics. Annu Rev Entomol 10: 97–116.

PLOS ONE | DOI:10.1371/journal.pone.0133602 July 30, 2015 25 / 26 Stable Mosquito Classification

84. Wiley EO (1979) An annotated Linnaean hierarchy, with comments on natural taxa and competing systems. Syst Zool 28: 308–337. doi: 10.1093/sysbio/28.3.308 85. Wiley EO (1981) Phylogenetics. The Theory of Phylogenetic Systematics. New York: Wiley-Inter- science. 439 p. 86. Wiley EO, Liebermann BS (2011) Phylogenetic Systematics. Theory and Practice of Phylogenetic Systematics. Hoboken, New Jersey: Wiley-Blackwell. 87. Hennig W (1981) Insect Phylogeny. New York: John Wiley and Sons. 88. de Queiroz K, Gauthier J (1992) Phylogenetic Taxonomy. Annu Rev Ecol Syst 23: 449–480. doi: 10. 1146/annurev.es.23.110192.002313 89. Vences M, Guayasamin JM, Miralles A, de la Riva I (2013) To name or not to name: Criteria to promote economy of change in Linnaean classification schemes. Zootaxa 3636: 201–244. PMID: 26042291. doi: 10.11646/zootaxa.3636.2.1 90. Goloboff PA, Farris JS, Nixon KC (2008) TNT, a free program for phylogenetic analysis. Cladistics 24: 774–786. doi: 10.1111/j.1096-0031.2008.00217.x 91. Swofford DL (2002) PAUP*. Phylogenetic Analysis Using Parsimony (*and Other Methods). Sunder- land, Massachusetts: Sinauer Associates. 92. Thiele K (1993) The Holy Grail of the perfect character: the cladistic treatment of morphometric data. Cladistics 9: 275–304. doi: 10.1111/j.1096-0031.1993.tb00226.x 93. Farris JS, Albert VA, Källersjo D, Lipscomb D, Kluge AG (1996) Parsimony jackknifing outperforms neighbor-joining. Cladistics 12: 99–124. doi: 10.1006/clad.1996.0008 94. Nixon KC (1999) The parsimony ratchet, a new method for rapid parsimony analysis. Cladistics 15: 407–414. doi: 10.1006/clad.1999.0121 95. Kluge AG (1997) Testbility and the refutation and corroboration of cladistic hyphotheses. Cladistics 13: 81–96. doi: 10.1111/j.1096-0031.1997.tb00242.x 96. Wood DM, Borkent A (1989) Phylogeny and classification of the Nematocera. In: McAlpine JF, Wood DM, editors. Manual of Nearctic Diptera. Hull, Quebec, Canada: Research Branch Agriculture Canada Monograph No. 32, Canadian Government Publishing Centre. pp. 1333–1370. 97. Miller BR, Crabtree MB, Savage HM (1997) Phylogenetic relationships of the Culicomorpha inferred from 18S and 5.8S ribosomal DNA sequences (Diptera: Nematocera). Insect Mol Biol 6: 105–114. PMID: 9099574. doi: 10.1111/j.1365-2583.1997.tb00078.x 98. Harbach RE, Kitching IJ (1998) Phylogeny and classification of the Culicidae (Diptera). Syst Entomol 23: 327–370. doi: 10.1046/j.1365-3113.1998.00072.x 99. Harbach RE (2007) The Culicidae (Diptera): a review of taxonomy, classification and phylogeny. Lin- naeus Tercentenary: Progress in Invertebrate Taxonomy 1668: 591–638. 100. Kaufman MG, Fonseca DM (2014) Invasion biology of Aedes japonicus japonicus. Annu Rev Entomol 59: 31–49. PMCID: PMC4106299. doi: 10.1146/annurev-ento-011613-162012 PMID: 24397520 101. Darsie RF, Ward RA (2005) Identification and geographical distribution of the mosquitos of North America, north of Mexico Gainesville Florida: University Press of Florida. xiv, 383 p. 102. Marcondes CB (2007) Taxonomic changes: disprove or accept them. Trends Parasitol 23: 302–303. doi: 10.1016/j.pt.2007.04.011 PMID: 17481955 103. Polaszek A (2006) Two worlds colliding: resistance to changes in the scientific names of animals— Aedes vs Stegomyia. Trends Parasitol 22: 8–9. PMID: 16300998. doi: 10.1016/j.pt.2005.11.003 104. Polaszek A (2007) Response to Marcondes: Accepting name changes. Trends Parasitol 23: 303. doi: 10.1016/j.pt.2007.04.010 105. Savage HM (2005) Classification of mosquitoes in tribe Aedini (Diptera: Culicidae): Paraphylyphobia, and classification versus cladistic analysis. J Med Entomol 42: 923–927. PMID: 16465729 doi: 10. 1093/jmedent/42.6.923 106. Glaubrechet M (2010) Evolution in action: Case studies in adaptive radiation, speciation and the origin of biodiversity. New York, NY, USA: Springer. 592 p.

PLOS ONE | DOI:10.1371/journal.pone.0133602 July 30, 2015 26 / 26 Appendix S1. Tribe Aedini generic, subgeneric and species names proposed herein. As appropriate, species endings are written to agree in gender with their respective genera and author parentheses are provided for species described in a genus different than in the current classification.

Genus Aedes

Subgenus Abraedes Zavortink, 1970

papago Zavortink, 1970

Subgenus Acartomyia Theobald, 1903

mariae (Sergent & Sergent, 1903) phoeniciae Coluzzi & Sabatini, 1968 zammitii (Theobald, 1903)

Subgenus Aedes Meigen, 1818

alexandrei Gornostaeva, 2005 cinereus Meigen, 1818 dahuricus Danilov, 1987 dmitryi Gornostaeva, 2005 esoensis Yamada, 1921 geminus Peus, 1970 mubiensis Luh & Shih, 1958 nataliae Gornostaeva, 2005 rossicus Dolbeskin, Gorickaja & Mitrofanova, 1930 sasai Tanaka, Mizusawa & Saugstad, 1975 valeryi Gornostaeva, 2005 yamadai Sasa, Kano & Takahasi, 1950

Subgenus Aedimorphus Theobald, 1903

abnormalis (Theobald, 1909) abnormalis kabwachensis Edwards, 1941 aerarius McIntosh, 1975 albodorsalis Fontenille & Brunhes, 1985 alboscutellatus (Theobald, 1905) alboventralis (Theobald, 1910) argenteoscutellatus Carter & Wijesundara, 1948 bambiotai Geoffroy, 1987 bancoi Geoffroy, 1987 bevisi (Edwards, 1915) caecus (Theobald, 1901) caliginosus (Graham, 1910) centropunctatus (Theobald, 1913) culicinus Edwards, 1922 cumminsii (Theobald, 1903) cumminsii mediopunctatus (Theobald, 1909) dalzieli (Theobald, 1910) davidi Basio, 1971 dentatus (Theobald, 1904) domesticus (Theobald, 1901) durbanensis (Theobald, 1903) durbanensis angolae Ribeiro & Ramos, 1974 ebogoensis Rickenbach & Ferrara, 1965 eritreae Lewis, 1942 fowleri (de Charmoy, 1908) gibbinsi Edwards, 1935 gouldi Reinert, 1972 grjebinei Hamon, Taufflieb & Maillot, 1957 hirsutus (Theobald, 1901) hirsutus adenensis Edwards, 1941 holocinctus Edwards, 1941 jamesi (Edwards, 1914) karooensis Muspratt, 1961 leesoni Edwards, 1932 leptolabis Edwards, 1936 leucarthrius (Speiser, 1909) longiseta Edwards, 1936 lowisii (Theobald, 1910) mansouri Qutubuddin, 1959 masoalensis Fontenille & Brunhes, 1985 mathioti Fontenille & Brunhes, 1985 mattinglyi Hamon & Rickenbach, 1954 mediolineatus (Theobald, 1901) natronius Edwards, 1932 nigricephalus (Theobald, 1901) nigrostriatus (Barraud, 1927) oakleyi Stone, 1939 ochraceus (Theobald, 1901) orbitae Edwards, 1922 ovazzai Hamon & Adam, 1959 pachyurus Edwards, 1936 pallidostriatus (Theobald, 1907) pampangensis (Ludlow, 1905) pipersalatus (Giles, 1902) pubescens Edwards, 1925 punctifemoris (Ludlow, 1921) quasiunivittatus (Theobald, 1901) rickenbachi Hamon & Adam, 1959 semlikiensis van Someren, 1950 senyavinensis Knight & Hurlbut, 1949 stenoetrus (Theobald, 1907) stenoscutus (Edwards, 1912) subdentatus Edwards, 1936 syntheticus Barraud, 1928 taeniorhynchoides (Christophers, 1911) tauffliebi Rickenbach & Ferrara, 1965 tricholabis Edwards, 1941 tricholabis bwamba (van Someren, 1950) trimaculatus (Theobald, 1905) trukensis Bohart, 1957 vexans (Meigen, 1830) vexans arabiensis (Patton, 1905) vexans nipponii (Theobald, 1907) vexans nocturnus (Theobald, 1903) wigglesworthi Edwards, 1941

Subgenus Alanstonea Mattingly, 1960

brevitibia (Edwards, 1914) treubi (de Meijere, 1910)

Subgenus Albuginosus Reinert, 1987

capensis Edwards, 1924 gilliesi van Someren, 1962 haworthi Edwards, 1923 kapretwae Edwards, 1941 kennethi Muspratt, 1956 marshallii (Theobald, 1901) ngong van Someren, 1950 stokesi Evans, 1929 teesdalei van Someren, 1954

Subgenus Ayurakitia Thurman, 1954

griffithi (Thurman, 1954) peytoni Reinert, 1972

Subgenus Aztecaedes Zavortink, 1972

ramirezi Vargas & Downs, 1950

Subgenus Belkinius Reinert, 1982

aurotaeniatus Edwards, 1922

Subgenus Bifidistylus Reinert, Harbach & Kitching, 2009

boneti Gil Collado, 1936 boneti kumbae Chwatt, 1948 lamborni Edwards, 1923

Subgenus Borichinda Harbach & Rattanarithikul, 2007

cavernicolus (Rattanarithikul & Harbach, 2007)

Subgenus Bothaella Reinert, 1973

alongi Galliard & Ngu, 1947 brownscutumus Dong, Zhu & Dong, 1999 eldridgei Reinert, 1973 helenae Reinert, 1973 kleini Reinert, 1973 manhi (Harbach & Cook, 2010)

Subgenus Bruceharrisonius Reinert, 2003

alektorovi Stackelberg, 1943 aureostriatus (Doleschall, 1857) christophersi Edwards, 1922 doonii Wattal, Bhatia & Kalra, 1958 greenii (Theobald, 1903) hurlbuti Lien, 1967 okinawanus Bohart, 1946 taiwanus Lien, 1968

Subgenus Cancraedes Edwards, 1929

cancricomes Edwards, 1922 curtipes Edwards, 1915 indonesiae Mattingly, 1958 kohkutensis Mattingly, 1958 mamoedjoensis Mattingly, 1958 masculinus Mattingly, 1958 palawanicus Mattingly, 1958 penghuensis Lien, 1968 simplex (Theobald, 1903) thurmanae Mattingly, 1958

Subgenus Catageiomyia Theobald, 1903

adami Geoffroy, 1971 argenteopunctatus (Theobald, 1901) bedfordi Edwards, 1936 chamboni Cornet, 1968 dialloi Hamon & Brengues, 1965 falabreguesi Hamon, 1957 filicis Ingram & de Meillon, 1927 grenieri Hamon, Service, Adam & Taufflieb, 1961 hopkinsi Edwards, 1936 insolens Edwards, 1936 irritans (Theobald, 1901) lokojoensis Service, 1959 lottei Hamon & Brengues, 1965 microstictus Edwards, 1936 minutus (Theobald, 1901) mixtus Edwards, 1936 mutilus Edwards, 1936 nyounae Hamon & Adam, 1959 phyllolabis Edwards, 1929 pseudotarsalis van Someren, 1946 punctothoracis (Theobald, 1909) reali Hamon & Adam, 1959 smithburni van Someren, 1950 tarsalis (Newstead, 1907) veeniae McIntosh, 1975 wendyae Service, 1959 yangambiensis de Meillon & Lavoipierre, 1944 yvonneae Edwards, 1941

Subgenus Catatassomyia Dyar & Shannon, 1925

meronephada (Dyar & Shannon, 1925)

Subgenus Christophersiomyia Barraud, 1923

annulirostris (Theobald, 1905) chionodes Belkin, 1962 gombakensis Mattingly, 1959 ibis Barraud, 1931 thomsoni (Theobald, 1905)

Subgenus Coetzeemyia Huang, Mathis & Wilkerson

fryeri (Theobald, 1912)

Subgenus Collessius Reinert, Harbach & Kitching, 2006

banksi Edwards, 1922 elsiae (Barraud, 1923) elsiae vicarius Lien, 1968 hatorii Yamada, 1921 macdougalli Edwards, 1922 macfarlanei (Edwards, 1914) ningheensis Lei, 1989 pseudotaeniatus (Giles, 1901) shortti (Barraud, 1923) tonkinensis Galliard & Ngu, 1947

Subgenus Cornetius Huang, 2005

cozi Cornet, 1973

Subgenus Dahliana Reinert, Harbach & Kitching, 2006

echinus (Edwards, 1920) geniculatus (Olivier, 1791) gilcolladoi (Sánchez-Covisa Villa, Rodríguez Rodríguez & Guillén Llera, 1985)

Subgenus Danielsia Theobald, 1904

albotaeniatus (Leicester, 1904) harperi Knight, 1948 lepchanus (Barraud, 1923)

Subgenus Dendroskusea Edwards, 1929

kanarensis Edwards, 1934 micropterus (Giles, 1901) periskelatus (Giles, 1902) ramachandrai Reuben, 1967 reginae Edwards, 1922

Subgenus Diceromyia Theobald, 1911

adersi (Edwards, 1917) bananea Wolfs, 1958 cordellieri Huang, 1986 coulangesi Rodhain & Boutonnier, 1983 fascipalpis (Edwards, 1912) flavicollis Edwards, 1928 furcifer (Edwards, 1913) grassei Doucet, 1951 madagascarensis van Someren, 1949 mefouensis Ferrara, 1974 sylvaticus Brunhes, 1983 taylori Edwards, 1936 tiptoni Grjebine, 1953 zethus de Meillon & Lavoipierre, 1944

Subgenus Dobrotworskyius Reinert, Harbach & Kitching, 2006

alboannulatus (Macquart, 1850) milsoni (Taylor, 1915) occidentalis (Skuse, 1889) rubrithorax (Macquart, 1850) rupestris Dobrotworsky, 1959 subbasalis Dobrotworsky, 1962 tubbutiensis Dobrotworsky, 1959

Subgenus Downsiomyia Vargas, 1950

albolateralis (Theobald, 1908) alboniveus Barraud, 1934 axitiosus Kulasekera, Knight & Harbach, 1990 dorseyi Knight, 1946 ganapathi Colless, 1958 harinasutai Knight, 1978 idjenensis Brug, 1934 inermis Colless, 1958 lacteus Knight, 1946 laoagensis Knight, 1946 leonis Colless, 1958 litoreus Colless, 1958 mikrokopion Knight & Harrison, 1988 mjobergi (Edwards, 1926) mohani Knight, 1969 nipponicus LaCasse & Yamaguti, 1948 nippononiveus Sasa & Nakahashi, 1952 nishikawai Tanaka, Mizusawa & Saugstad, 1979 niveoides Barraud, 1934 niveus (Ludlow, 1903) novoniveus Barraud, 1934 omorii Lien, 1968 pexus Colless, 1958 pseudoniveus (Theobald, 1905) saperoi Knight, 1946 shehzadae Qutubuddin, 1972 sinensis Chow, 1950 subniveus Edwards, 1922 vanus Colless, 1958 watteni Lien, 1968

Subgenus Edwardsaedes Belkin, 1962

bekkui Mogi, 1977 imprimens (Walker, 1861) pingpaensis Chang, 1965

Subgenus Elpeytonius Reinert, Harbach & Kitching, 2009

apicoannulatus (Edwards, 1912) simulans (Newstead & Carter, 1911)

Subgenus Finlaya Theobald, 1903

alocasicola Marks, 1947 ananae Knight & Laffoon, 1946 avistylus Brug, 1939 bougainvillensis Marks, 1947 burnetti Belkin, 1962 croceus Knight & Laffoon, 1946 dobrotworskyi Marks, 1958 fijiensis Marks, 1947 flavipennis Giles, 1904 franclemonti Belkin, 1962 freycinetiae Laird, 1957 fuscipalpis Belkin, 1962 fuscitarsis Belkin, 1962 gahnicola Marks, 1947 gani Bonne-Wepster, 1940 gressitti Bohart, 1957 hollingsheadi Belkin, 1962 horotoi Taylor, 1972 hui Bohart, 1957 josephinae Marks, 1958 knighti Stone & Bohart, 1944 kochi (Dönitz, 1901) lewelleni Starkey & Webb, 1946 luteus (Ludlow, 1905) maffii Taylor & Tenorio, 1974 medleri Knight & Laffoon, 1946 neogeorgianus Belkin, 1962 oceanicus Belkin, 1962 poicilius (Theobald, 1903) samoanus (Grünberg, 1913) schlosseri Belkin, 1962 solomonis Stone & Bohart, 1944 sorsogonensis Bañez & Jueco, 1966 stonei Knight & Laffoon, 1946 tutuilae Ramalingam & Belkin, 1965 wallacei Edwards, 1926

Subgenus Fredwardsius Reinert, 2000

vittatus (Bigot, 1861)

Subgenus Georgecraigius Reinert, Harbach & Kitching, 2006

atropalpus (Coquillett, 1902) epactius Dyar & Knab, 1908 fluviatilis (Lutz, 1904)

Subgenus Geoskusea Edwards, 1929

baisasi Knight & Hull, 1951 becki Belkin, 1962 daggyi Stone & Bohart, 1944 fimbripes Edwards, 1924 kabaenensis Brug, 1939 longiforceps Edwards, 1929 lunulatus King & Hoogstraal, 1946 perryi Belkin, 1962 timorensis (Miyagi, Toma & Lien, 2004) tonsus Edwards, 1924

Subgenus Gilesius Reinert, Harbach & Kitching, 2006

alius Lien, 1968 pulchriventer (Giles, 1901)

Subgenus Gymnometopa Coquillett, 1905

mediovittatus (Coquillett, 1906)

Subgenus Halaedes Belkin, 1962

ashworthi Edwards, 1921 australis (Erichson, 1842) wardangensis Brust, Ballard, Driver, Hartley, Galway & Curran, 1998

Subgenus Himalaius Reinert, Harbach & Kitching, 2006

gilli (Barraud, 1924) simlensis Edwards, 1922

Subgenus Hopkinsius Reinert, Harbach & Kitching, 2008

albocinctus (Barraud, 1924) barnardi Edwards, 1924 embuensis Edwards, 1930 ingrami Edwards, 1930 nyasae Edwards, 1930 seoulensis Yamada, 1921 wellmanii (Theobald, 1905)

Subgenus Howardina Theobald, 1903

albonotatus (Coquillett, 1906) allotecnon Kumm, Komp & Ruiz, 1940 arborealis Bonne-Wepster & Bonne, 1920 argyrites Dyar & Núñez Tovar, 1927 aureolineatus Berlin, 1969 aurites (Theobald, 1907) aurivittatus Cerqueira, 1943 bahamensis Berlin, 1969 brevis Berlin, 1969 brevivittatus Berlin, 1969 busckii (Coquillett, 1906) cozumelensis Díaz Nájera, 1966 ecuadoriensis Berlin, 1969 eleanorae Berlin, 1969 fulvithorax (Lutz, 1904) grabhami Berlin, 1969 guatemala Berlin, 1969 guerrero Berlin, 1969 inaequalis (Grabham, 1907) ioliota Dyar & Knab, 1913 leei Berlin, 1969 lorraineae Berlin, 1969 marinkellei Berlin, 1969 martinezi Berlin, 1969 osornoi Berlin, 1969 pseudodominicii Komp, 1936 quadrivittatus (Coquillett, 1902) septemstriatus Dyar & Knab, 1907 sexlineatus (Theobald, 1901) spinosus Berlin, 1969 stenei Thompson, 1956 vanemdeni Martini, 1931 walkeri (Theobald, 1901) whitmorei Dunn, 1918

Subgenus Huaedes Huang, 1968

medialis (Brug, 1932) variepictus (King & Hoogstraal, 1946) wauensis Huang, 1968

Subgenus Hulecoeteomyia Theobald, 1904

chrysolineatus (Theobald, 1907) formosensis Yamada, 1921 harveyi (Barraud, 1923) japonicus (Theobald, 1901) japonicus amamiensis Tanaka, Mizusawa & Saugstad, 1979 japonicus shintienensis Tsai & Lien, 1950 japonicus yaeyamensis Tanaka, Mizusawa & Saugstad, 1979 jugraensis (Leicester, 1908) koreicus (Edwards, 1917) nigrorhynchus Brug, 1931 pallirostris Edwards, 1922 reinerti Rattanarithikul & Harrison, 1988 rizali (Banks, 1906) saxicola Edwards, 1922 sherki Knight, 1948 yunnanensis (Gaschen, 1934)

Subgenus Indusius Edwards, 1934

pulverulentus Edwards, 1922

Subgenus Isoaedes Reinert, 1979

cavaticus Reinert, 1979

Subgenus Jarnellius Reinert, Harbach & Kitching, 2006

deserticola Zavortink, 1969 laguna Arnell & Nielsen, 1972 monticola Belkin & McDonald, 1957 sierrensis (Ludlow, 1905) varipalpus (Coquillett, 1902)

Subgenus Jihlienius Reinert, Harbach & Kitching, 2006

chungi Lien, 1968 gonguoensis Gong & Lu, 1986 unicinctus Edwards, 1922

Subgenus Kenknightia Reinert, 1990

dissimilierodes Dong, Zhou & Dong, 2002 dissimilis (Leicester, 1908) gaffigani Reinert, 1990 harbachi Reinert, 1990 karwari (Barraud, 1924) lerozeboomi Reinert, 1990 leucomeres (Giles, 1904) litwakae Reinert, 1990 luzonensis Rozeboom, 1946 paradissimilis Rozeboom, 1946 pecori Reinert, 1990 wilkersoni Reinert, 1990

Subgenus Kompia Aitken, 1941

purpureipes Aitken, 1941

Subgenus Leptosomatomyia Theobald, 1905

aurimargo Edwards, 1922

Subgenus Levua Stone & Bohart, 1944

geoskusea Amos, 1944

Subgenus Lewnielsenius Reinert, Harbach & Kitching, 2006

muelleri Dyar, 1920

Subgenus Lorrainea Belkin, 1962

amesii (Ludlow, 1903) celebicus Mattingly, 1959 dasyorrhus King & Hoogstraal, 1946 fumidus Edwards, 1928 lamelliferus Bohart & Ingram, 1946

Subgenus Luius Reinert, Harbach & Kitching, 2008

fengi Edwards, 1935

Subgenus Macleaya Theobald, 1903

calabyi Marks, 1963 elchoensis Taylor, 1929 humeralis Edwards, 1922 littlechildi Taylor, 1933 macmillani Marks, 1964 moloiensis Taylor, 1929 spinosipes Edwards, 1922 stoneorum Marks, 1977 tremulus (Theobald, 1903) tulliae (Taylor, 1929) wattensis Taylor, 1929

Subgenus Molpemyia Theobald, 1910

auridorsum Edwards, 1922 pecuniosus Edwards, 1922 purpureus (Theobald, 1910)

Subgenus Mucidus Theobald, 1901

alternans (Westwood, 1835) aurantius (Theobald, 1907) aurantius chrysogaster (Taylor, 1927) ferinus Knight, 1947 grahamii (Theobald, 1909) laniger (Wiedemann, 1820) lucianus Muspratt, 1959 mucidus (Karsch, 1887) nigerrimus (Theobald, 1913) nigrescens (Edwards, 1929) quadripunctis (Ludlow, 1910) quasiferinus Mattingly, 1961 scatophagoides (Theobald, 1901) sudanensis (Theobald, 1908) tonkingi Gebert, 1948

Subgenus Neomelaniconion Newstead, 1907

albicosta (Edwards, 1913) albiradius (le Goff, Boussès & Brunhes, 2007) albothorax (Theobald, 1907) aurovenatus Worth, 1960 belleci (le Goff, Boussès & Brunhes, 2007) bequaerti Wolfs, 1947 bergerardi Pajot & Geoffroy, 1971 bolense Edwards, 1936 carteri Edwards, 1936 circumluteolum (Theobald, 1908) crassiforceps Edwards, 1927 ellinorae Edwards, 1941 flavimargo Edwards, 1941 fontenillei (le Goff, Boussès & Brunhes, 2007) fuscinervis (Edwards, 1914) jamoti Hamon & Rickenbach, 1954 lineatopennis (Ludlow, 1905) lineatopennis aureum (Gutsevich, 1955) luridus McIntosh, 1971 luteolateralis (Theobald, 1901) mcintoshi Huang, 1985 monotrichus Edwards, 1936 nigropterum (le Goff, Boussès & Brunhes, 2007) palpale Newstead, 1907 pogonurus Edwards, 1936 punctocostalis (Theobald, 1909) sylvaticum (le Goff, Boussès & Brunhes, 2007) taeniarostris (Theobald, 1909) unidentatum McIntosh, 1971

Subgenus Nyctomyia Harbach, 2013

pholeocolus (Linton & Harbach, 2013) biunguiculatus (Harbach, 2014)

Subgenus Ochlerotatus Lynch Arribalzaga, 1891

aboriginis Dyar, 1917 abserratus (Felt & Young, 1904) aculeatus (Theobald, 1903) aenigmaticus Cerqueira & Costa, 1946 akkeshiensis Tanaka, 1998 albescens Edwards, 1921 albifasciatus (Macquart, 1838) albineus Séguy, 1923 aloponotum Dyar, 1917 ambreensis Rodhain & Boutonnier, 1983 andersoni Edwards, 1926 angustivittatus Dyar & Knab, 1907 annulipes (Meigen, 1830) antipodeus (Edwards, 1920) atactavittatus Arnell, 1976 atlanticus Dyar & Knab, 1906 auratus Grabham, 1906 aurifer (Coquillett, 1903) bancroftianus Edwards, 1921 behningi Martini, 1926 bejaranoi Martinez, Carcavallo & Prosen, 1960 berlandi Séguy, 1921 bicristatus Thurman & Winkler, 1950 bimaculatus (Coquillett, 1902) biskraensis Brunhes, 1999 bogotanus (Arnell, 1976) breedensis Muspratt, 1953 burjaticus (Kuchartshuk, 1973) burpengaryensis (Theobald, 1905) caballus (Theobald, 1912) cacozelus Marks, 1963 calcariae Marks, 1957 calumnior Belkin, Heinemann & Page, 1970 campestris Dyar & Knab, 1907 camptorhynchus (Thomson, 1869) canadensis (Theobald, 1901) canadensis mathesoni Middlekauff cantans (Meigen, 1818) cantator (Coquillett, 1903) caspius (Pallas, 1771) caspius hargreavesi Edwards, 1920 caspius meirae Ribeiro, Ramos, Capela & Pires, 1980 cataphylla Dyar, 1916 chelli (Edwards, 1915) churchillensis Ellis & Brust, 1973 clelandi (Taylor, 1914) clivis Lanzaro & Eldridge, 1992 coluzzii Rioux, Guilvard & Pasteur, 1998 comitatus Arnell, 1976 communis (de Geer, 1776) condolescens Dyar & Knab, 1907 continentalis Dobrotworsky, 1960 crinifer (Theobald, 1903) culiciformis (Theobald, 1905) cunabulanus Edwards, 1924 cyprioides Danilov & Stupin, 1982 cyprius Ludlow, 1920 dahlae (Nielsen, 2009) decticus Howard, Dyar & Knab, 1917 deficiens Arnell, 1976 detritus Haliday, 1833 diantaeus Howard, Dyar & Knab, 1913 dorsalis (Meigen, 1830) dufouri Hamon, 1953 duplex Martini, 1926 dupreei (Coquillett, 1904) dzeta Séguy, 1924 edgari Stone & Rosen, 1952 eidsvoldensis Mackerras, 1927 eucephalaeus Dyar, 1918 euedes Howard, Dyar & Knab, 1913 euiris Dyar, 1922 euplocamus Dyar & Knab, 1906 excrucians (Walker, 1856) explorator Marks, 1964 fitchii (Felt & Young, 1904) flavescens (Müller, 1764) flavifrons (Skuse, 1889) fulvus (Wiedemann, 1828) fulvus pallens Ross, 1943 grossbecki Dyar & Knab, 1906 gutzevichi Dubitsky & Deshevykh, 1978 hakusanensis Yamaguti & Tamaboko, 1954 harrisoni Muspratt, 1953 hastatus Dyar, 1922 hesperonotius Marks, 1959 hexodontus Dyar, 1916 hodgkini Marks, 1959 hokkaidensis Tanaka, Mizusawa & Saugstad, 1979 hungaricus Mihalyi, 1955 imperfectus Dobrotworsky, 1962 impiger (Walker, 1848) impiger daisetsuzanus Tanaka, Mizusawa & Saugstad, 1979 implicatus Vockeroth, 1954 incomptus Arnell, 1976 increpitus Dyar, 1916 inexpectatus Bonne-Wepster, 1948 infirmatus Dyar & Knab, 1906 intermedius Danilov & Gornostaeva, 1987 intrudens Dyar, 1919 jacobinae Serafim & Davis, 1933 jorgi Carpintero & Leguizamón, 2000 juppi McIntosh, 1973 kasachstanicus Gutsevich, 1962 krymmontanus Alekseev, 1989 lasaensis Meng, 1962 lasaensis gyirongensis Ma, 1982 lepidonotus Edwards, 1920 lepidus Cerqueira & Paraense, 1945 leucomelas (Meigen, 1804) linesi Marks, 1964 longifilamentus Su & Zhang, 1988 luteifemur Edwards, 1926 macintoshi Marks, 1959 martineti Senevet, 1937 mcdonaldi Belkin, 1962 melanimon Dyar, 1924 meprai Martinez & Prosen, 1953 mercurator Dyar, 1920 milleri Dyar, 1922 mitchellae (Dyar, 1905) montchadskyi Dubitsky, 1968 multiplex (Theobald, 1903) nevadensis Chapman & Barr, 1964 nigrinus (Eckstein, 1918) nigripes (Zetterstedt, 1838) nigrithorax (Macquart, 1847) nigrocanus Martini, 1927 nigromaculis (Ludlow, 1906) niphadopsis Dyar & Knab, 1918 nivalis Edwards, 1926 normanensis (Taylor, 1915) nubilus Theobald, 1903 obturbator Dyar & Knab, 1907 oligopistus Dyar, 1918 patersoni Shannon & del Ponte, 1928 pectinatus Arnell, 1976 pennai Antunes & Lane, 1938 perkinsi Marks, 1949 pertinax Grabham, 1906 phaecasiatus Marks, 1964 phaeonotus Arnell, 1976 pionips Dyar, 1919 postspiraculosus Dobrotworsky, 1961 procax (Skuse, 1889) provocans (Walker, 1848) pseudonormanensis Marks, 1949 pulcritarsis (Rondani, 1872) pulcritarsis asiaticus Edwards, 1926 pullatus (Coquillett, 1904) punctodes Dyar, 1922 punctor (Kirby, 1837) purpuraceus Brug, 1932 purpureifemur Marks, 1959 purpuriventris Edwards, 1926 quasirusticus Torres Cañamares, 1951 ratcliffei Marks, 1959 raymondi del Ponte, Castro & García, 1951 refiki Medschid, 1928 rempeli Vockeroth, 1954 rhyacophilus da Costa Lima, 1933 riparioides Su & Zhang, 1987 riparius Dyar & Knab, 1907 rusticus (Rossi, 1790) rusticus subtrichurus Martini, 1927 rusticus trichurus (Dyar, 1904) sagax (Skuse, 1889) sapiens Marks, 1964 scapularis (Rondani, 1848) schizopinax Dyar, 1929 schtakelbergi Shingarev, 1928 scutellalbum Boshell-Manrique, 1939 sedaensis Lei, 1989 sergievi Danilov, Markovich & Proskuryakova, 1978 serratus (Theobald, 1901) shannoni Vargas & Downs, 1950 silvestris (Dobrotworsky, 1961) simanini Gutsevich, 1966 sinkiangensis Hsiao, 1977 sollicitans (Walker, 1856) spencerii (Theobald, 1901) spencerii idahoensis (Theobald, 1903) spilotus Marks, 1963 squamiger (Coquillett, 1902) sticticus (Meigen, 1838) stigmaticus Edwards, 1922 stimulans (Walker, 1848) stramineus Dubitzky, 1970 stricklandi (Edwards, 1912) subalbirostris Klein & Marks, 1960 subdiversus Martini, 1926 surcoufi (Theobald, 1912) synchytus Arnell, 1976 taeniorhynchus (Wiedemann, 1821) tahoensis Dyar, 1916 thelcter Dyar, 1918 theobaldi (Taylor, 1914) thibaulti Dyar & Knab, 1910 tormentor Dyar & Knab, 1906 tortilis (Theobald, 1903) trivittatus (Coquillett, 1902) turneri Marks, 1963 upatensis Anduze & Hecht, 1943 ventrovittis Dyar, 1916 vigilax (Skuse, 1889) vigilax ludlowae (R. Blanchard) vigilax vansomerenae Mattingly vittiger (Skuse, 1889) washinoi Lanzaro & Eldridge, 1992

Subgenus Paraedes Edwards, 1934

barraudi (Edwards, 1934) bonneae Mattingly, 1958 chrysoscuta (Theobald, 1910) collessi Mattingly, 1958 menoni Mattingly, 1958 ostentatio (Leicester, 1908) pagei (Ludlow, 1911) thailandensis Reinert, 1976

Subgenus Patmarksia Reinert, Harbach & Kitching, 2006

anggiensis Bonne-Wepster, 1937 argenteitarsis Brug, 1932 argyronotum Belkin, 1962 buxtoni Belkin, 1962 clintoni Taylor, 1946 derooki Brug, 1932 dobodurus King & Hoogstraal, 1946 hollandius King & Hoogstraal, 1946 mackerrasi Taylor, 1927 novalbitarsis King & Hoogstraal, 1946 palmarum Edwards, 1924 papuensis (Taylor, 1914) subalbitarsis King & Hoogstraal, 1946

Subgenus Petermattinglyius Reinert, Harbach & Kitching, 2009

franciscoi Mattingly, 1959 iyengari Edwards, 1923 punctipes Edwards, 1921 scanloni Reinert, 1970 whartoni Mattingly, 1965

Subgenus Phagomyia Theobald, 1905

assamensis (Theobald, 1908) cacharanus (Barraud, 1923) cogilli Edwards, 1922 deccanus (Barraud, 1923) feegradei Barraud, 1934 gubernatoris (Giles, 1901) gubernatoris kotiensis Barraud, 1934 inquinatus Edwards, 1922 iwi Marks, 1955 khazani Edwards, 1922 kiangsiensis Tung, 1955 lophoventralis (Theobald, 1910) melanopterus (Giles, 1904) plumiferus King & Hoogstraal, 1946 prominens (Barraud, 1923) stevensoni (Barraud, 1923) watasei Yamada, 1921

Subgenus Polyleptiomyia Theobald, 1905

albocephalus (Theobald, 1903) gandarai da Cunha Ramos, Capela & Ribeiro, 1995

Subgenus Pseudoarmigeres Stone & Knight, 1956

albomarginatus (Newstead, 1907) argenteoventralis (Theobald, 1909) argenteoventralis dunni (Evans, 1928) kummi Edwards, 1930 michaelikati van Someren, 1946 michaelikati gurneri van Someren, 1946 natalensis Edwards, 1930

Subgenus Rampamyia Reinert, Harbach & Kitching, 2006

albilabris Edwards, 1925 notoscriptus (Skuse, 1889) notoscriptus montanus Brug, 1939 quinquelineatus Edwards, 1922

Subgenus Rhinoskusea Edwards, 1929

longirostris (Leicester, 1908) pillaii Mattingly, 1958 portonovoensis Tewari & Hiriyan, 1992 wardi Reinert, 1976

Subgenus Sallumia Reinert, Harbach & Kitching, 2008

hortator Dyar & Knab, 1907 perventor Cerqueira & Costa, 1946

Subgenus Scutomyia Theobald, 1904

albolineatus (Theobald, 1904) arboricola Knight & Rozeboom, 1946 bambusicola Knight & Rozeboom, 1946 boharti Knight & Rozeboom, 1946 hoogstraali Knight & Rozeboom, 1946 impatibilis (Walker, 1860) laffooni Knight & Rozeboom, 1946 platylepidus Knight & Hull, 1951 pseudalbolineatus Brug, 1939

Subgenus Skusea Theobald, 1903

cartroni (Ventrillon, 1906) lambrechti van Someren, 1971 moucheti Ravaonjanahary & Brunhes, 1977 pembaensis Theobald, 1901

Subgenus Stegomyia Theobald, 1901

aegypti (Linnaeus, 1762) aegypti formosus (Walker, 1848) africanus (Theobald, 1901) agrihanensis Bohart, 1957 albopictus (Skuse, 1895) alcasidi Huang, 1972 alorensis Bonne-Wepster & Brug, 1932 amaltheus de Meillon & Lavoipierre, 1944 andrewsi Edwards, 1926 angustus Edwards, 1935 annandalei (Theobald, 1910) aobae Belkin, 1962 apicoargenteus (Theobald, 1909) bambusae Edwards, 1935 blacklocki Evans, 1925 bromeliae (Theobald, 1911) burnsi Basio & Reisen, 1971 calceatus Edwards, 1924 chaussieri Edwards, 1923 chemulpoensis Yamada, 1921 contiguus Edwards, 1936 cooki Belkin, 1962 corneti Huang, 1986 craggi (Barraud, 1923) cretinus Edwards, 1921 daitensis Miyagi & Toma, 1981 deboeri Edwards, 1926 demeilloni Edwards, 1936 denderensis Wolfs, 1949 dendrophilus Edwards, 1921 desmotes (Giles, 1904) dybasi Bohart, 1957 ealaensis Huang, 2004 edwardsi (Barraud, 1923) ethiopiensis Huang, 2004 flavopictus Yamada, 1921 flavopictus downsi Bohart & Ingram, 1946 flavopictus miyarai Tanaka, Mizusawa & Saugstad, 1979 fraseri (Edwards, 1912) futunae Belkin, 1962 galloisi Yamada, 1921 galloisiodes Liu & Lu, 1984 gandaensis Huang, 2004 gardnerii (Ludlow, 1905) gardnerii imitator (Leicester, 1908) grantii (Theobald, 1901) guamensis Farner & Bohart, 1944 gurneyi Stone & Bohart, 1944 hakanssoni Knight & Hurlbut, 1949 hansfordi Huang, 1997 hebrideus Edwards, 1926 heischi van Someren, 1951 hensilli Farner, 1945 hogsbackensis Huang, 2004 hoguei Belkin, 1962 horrescens Edwards, 1935 josiahae Huang, 1988 katherinensis Woodhill, 1949 keniensis van Someren, 1946 kenyae van Someren, 1946 kesseli Huang & Hitchcock, 1980 kivuensis Edwards, 1941 krombeini Huang, 1975 langata van Someren, 1946 ledgeri Huang, 1981 lilii (Theobald, 1910) luteocephalus (Newstead, 1907) maehleri Bohart, 1957 malayensis Colless, 1962 malikuli Huang, 1973 marshallensis Stone & Bohart, 1944 mascarensis MacGregor, 1924 masseyi Edwards, 1923 mattinglyorum Huang, 1994 maxgermaini Huang, 1990 mediopunctatus (Theobald, 1905) mediopunctatus sureilensis Barraud, 1934 metallicus (Edwards, 1912) mickevichae Huang, 1988 mpusiensis Huang, 2004 muroafcete Huang, 1997 neoafricanus Cornet, Valade & Dieng, 1978 neogalloisi Chen & Chen, 2000 neopandani Bohart, 1957 njombiensis Huang, 1997 novalbopictus Barraud, 1931 opok Corbet & van Someren, 1962 palauensis Bohart, 1957 pandani Stone, 1939 patriciae Mattingly, 1954 paullusi Stone & Farner, 1945 pernotatus Farner & Bohart, 1944 perplexus (Leicester, 1908) polynesiensis Marks, 1951 poweri (Theobald, 1905) pseudalbopictus (Borel, 1928) pseudoafricanus Chwatt, 1949 pseudonigeria (Theobald, 1910) pseudoscutellaris (Theobald, 1910) quasiscutellaris Farner & Bohart, 1944 rhungkiangensis Chang & Chang, 1974 riversi Bohart & Ingram, 1946 robinsoni Belkin, 1962 rotanus Bohart & Ingram, 1946 rotumae Belkin, 1962 ruwenzori Haddow & van Someren, 1950 saimedres Huang, 1988 saipanensis Stone, 1945 sampi Huang, 2004 schwetzi Edwards, 1926 scutellaris (Walker, 1858) scutoscriptus Bohart & Ingram, 1946 seampi Huang, 1974 seatoi Huang, 1969 segermanae Huang, 1997 sibiricus Danilov & Filippova, 1978 simpsoni (Theobald, 1905) soleatus Edwards, 1924 strelitziae Muspratt, 1950 subalbopictus Barraud, 1931 subargenteus Edwards, 1925 tabu Ramalingam & Belkin, 1965 tongae Edwards, 1926 tulagiensis Edwards, 1926 unilineatus (Theobald, 1906) upolensis Marks, 1957 usambara Mattingly, 1953 varuae Belkin, 1962 vinsoni Mattingly, 1953 w-albus (Theobald, 1905) wadai Tanaka, Mizusawa & Saugstad, 1979 woodi Edwards, 1922

Subgenus Tanakaius Reinert, Harbach & Kitching, 2004

savoryi Bohart, 1957 togoi (Theobald, 1907)

Subgenus Tewarius Reinert, 2006

agastyai Tewari & Hiriyan, 1992 nummatus Edwards, 1923 pseudonummatus Reinert, 1973 reubenae Tewari & Hiriyan, 1992

Subgenus Vansomerenis Reinert, Harbach & Kitching, 2006

luteostriatus Robinson, 1950 pulchrithorax Edwards, 1939 hancocki van Someren, 1962

Subgenus Zavortinkius Reinert, 1999

brunhesi Reinert, 1999 brygooi Brunhes, 1971 fulgens (Edwards, 1917) geoffroyi Reinert, 1999 huangae Reinert, 1999 interruptus Reinert, 1999 longipalpis (Grünberg, 1905) monetus Edwards, 1935 mzooi van Someren, 1962 phillipi van Someren, 1949 pollinctor (Graham, 1910)

Subgenus uncertain

aitkeni Schick, 1970 alboapicus Schick, 1970 alticolaa Bonne-Wepster, 1948 amabilis Schick, 1970 argyrothorax Bonne-Wepster & Bonne, 1920 auronitens Edwards, 1922 australiensis (Theobald, 1910) berlini Schick, 1970 bertrami Schick, 1970 biocellatus (Taylor, 1915) braziliensis Gordon & Evans, 1922 brelandi Zavortink, 1972 britteni Marks & Hodgkin, 1958 buenaventura Schick, 1970 burgeri Zavortink, 1972 campana Schick, 1970 candidoscutellum Marks, 1947 casali Schick, 1970 chionotum Zavortink, 1972 crossi Lien, 1967 daliensis (Taylor, 1916) daryi Schick, 1970 diazi Schick, 1970 eatoni (Edwards, 1916) gabriel Schick, 1970 galindoi Schick, 1970 gracilelineatus Bonne-Wepster, 1937 hendersoni Cockerell, 1918 heteropus Dyar, 1921 homoeopus Dyar, 1922 idanus Schick, 1970 impostor Schick, 1970 insolitus (Coquillett, 1906) keefei King & Hoogstraal, 1946 knabi (Coquillett, 1906) kompi Vargas & Downs, 1950 koreicoides Sasa, Kano & Hayashi, 1950 lauriei (Carter, 1920) mallochi Taylor, 1944 metoecopus Dyar, 1925 monocellatus Marks, 1948 niveoscutum Zavortink, 1972 oreophilus (Edwards, 1916) peipingensis Feng, 1938 plagosus Marks, 1959 podographicus Dyar & Knab, 1906 quasirubithorax (Theobald, 1910) roai Belkin, 1962 rubiginosus Belkin, 1962 sandrae Zavortink, 1972 schicki Zavortink, 1972 schroederi Schick, 1970 sintoni (Barraud, 1924) stanleyi Peters, 1963 subauridorsum Marks, 1948 suffusus Edwards, 1922 sumidero Schick, 1970 tehuantepec Schick, 1970 terrens (Walker, 1856) thorntoni Dyar & Knab, 1907 toxopeusi Bonne-Wepster, 1948 triseriatus (Say, 1823) tsiliensis King & Hoogstraal, 1946 vargasi Schick, 1970 versicolor (Barraud, 1924) wasselli Marks, 1947 zavortinki Schick, 1970 zoosophus Dyar & Knab, 1918

Not placed by Sathe & Girhe, 2002

kolhapuriensis Sathe & Girhe, 2002 panchgangee Sathe & Girhe, 2002 sangitee Sathe & Girhe, 2002 sangiti Girhe & Sathe, 2001

Genus Armigeres

Armigeres Theobald, 1901

alkatirii Toma, Miyagi & Syafruddin, 1995 apoensis Bohart & Farner, 1944 aureolineatus (Leicester, 1908) azurini Basio, 1971 baisasi Stone & Thurman, 1958 bhayungi Thurman & Thurman, 1958 breinli (Taylor, 1914) candelabrifer Brug, 1939 confusus Edwards, 1915 conjungens Edwards, 1914 denbesteni Brug, 1925 durhami (Edwards, 1917) ejercitoi Baisas, 1935 fimbriatus Edwards, 1930 foliatus Brug, 1931 giveni Edwards, 1926 hybridus Edwards, 1914 joloensis (Ludlow, 1904) jugraensis (Leicester, 1908) kesseli Ramalingam, 1987 kinabaluensis Ramalingam, 1972 kuchingensis Edwards, 1915 lacuum Edwards, 1922 laoensis Toma & Miyagi, 2003 maiae (Edwards, 1917) malayi (Theobald, 1901) manalangi Baisas, 1935 maximus Edwards, 1922 milnensis Lee, 1944 moultoni Edwards, 1914 obturbans (Walker, 1859) pallithorax Dong, Zhou & Dong, 2004 papuensis Peters, 1963 setifer Delfinado, 1966 seticoxitus Luh & Li, 1981 subalbatus (Coquillett, 1898) subalbatus chrysocorporis Hsieh & Liao, 1956 theobaldi Barraud, 1934 yunnanensis Dong, Zhou & Dong, 1995

Leicesteria Theobald, 1904

annulipalpis (Theobald, 1910) annulitarsis (Leicester, 1908) balteatus Macdonald, 1960 cingulatus (Leicester, 1908) dentatus Barraud, 1927 digitatus (Edwards, 1914) dolichocephalus (Leicester, 1908) flavus (Leicester, 1908) inchoatus Barraud, 1927 lepidocoxitus Dong, Zhou & Dong, 1995 longipalpis (Leicester, 1904) magnus (Theobald, 1908) menglaensis Dong, Zhou & Dong, 2002 omissus (Edwards, 1914) pectinatus (Edwards, 1914) pendulus (Edwards, 1914) traubi Macdonald, 1960 vimoli Thurman & Thurman, 1958

Genus Eretmapodites

adami Ferrara & Eouzan, 1974 angolensis da Cunha Ramos & Ribeiro, 1992 argyrurus Edwards, 1936 brenguesi Rickenbach & Lombrici, 1975 brottesi Rickenbach, 1967 caillardi Rickenbach, Ferrara & Eouzan, 1968 chrysogaster Graham, 1909 corbeti Hamon, 1962 dracaenae Edwards, 1916 dundo da Cunha Ramos & Ribeiro, 1992 eouzani Rickenbach & Lombrici, 1974 ferrarai Rickenbach & Eouzan, 1970 forcipulatus Edwards, 1936 germaini Rickenbach & Eouzan, 1970 gilletti van Someren, 1949 grahami Edwards, 1911 grenieri Hamon & van Someren, 1961 haddowi van Someren, 1949 hamoni Grjebine, 1972 harperi van Someren, 1949 hightoni van Someren, 1947 inornatus Newstead, 1907 intermedius Edwards, 1936 jani Rickenbach & Lombrici, 1976 lacani Rickenbach & Eouzan, 1970 leucopous Graham, 1909 mahaffyi van Someren, 1949 marcellei Adam & Hamon, 1959 mattinglyi Hamon & van Someren, 1961 melanopous Graham, 1909 mortiauxi Cunha Ramos & Ribeiro, 1990 oedipodeios Graham, 1909 parvipluma Edwards, 1941 pauliani Grjebine, 1950 penicillatus Edwards, 1941 plioleucus Edwards, 1941 plioleucus brevis Edwards, 1941 productus Edwards, 1941 quinquevittatus Theobald, 1901† ravissei Rickenbach & Eouzan, 1970 rickenbachi Ferrara & Eouzan, 1974 salauni Rickenbach, Ferrara & Eouzan, 1968 semisimplicipes Edwards, 1914 silvestris Ingram & de Meillon, 1927 silvestris conchobius Edwards, 1941 subsimplicipes Edwards, 1914 tendeiroi da Cunha Ramos, Ribeiro & Machado, 1992 tonsus Edwards, 1941 vansomereni Hamon, 1962 wansoni Edwards, 1941 wansoni douceti Adam & Hamon, 1959

Genus Haemagogus

Subgenus Conopostegus Dyar, 1925

clarki (Galindo, Carpenter & Trapido, 1953) leucocelaenus (Dyar & Shannon, 1924) leucophoebus (Galindo, Carpenter & Trapido, 1953) leucotaeniatus (Komp, 1938)

Subgenus Haemagogus Williston, 1896

acutisentis Arnell, 1973 aeritinctus Galindo & Trapido, 1967 albomaculatus Theobald, 1903 anastasionis Dyar, 1921 andinus Osorno-Mesa, 1944 argyromeris Dyar & Ludlow, 1921 baresi Cerqueira, 1960 boshelli Osorno-Mesa, 1944 capricornii Lutz, 1904 celeste Dyar & Nuñez Tovar, 1927 chalcospilans Dyar, 1921 chrysochlorus Arnell, 1973 equinus Theobald, 1903 iridicolor Dyar, 1921 janthinomys Dyar, 1921 lucifer (Howard, Dyar & Knab, 1913) mesodentatus Komp & Kumm, 1938 nebulosus Arnell, 1973 panarchys Dyar, 1921 regalis Dyar & Knab, 1906 soperi Levi-Castillo, 1955 spegazzinii Brethès, 1912 splendens Williston, 1896 tropicalis Cerqueira & Antunes, 1938

Genus Heizmannia

Subgenus Heizmannia Ludlow, 1905

aurea Brug, 1932 aureochaeta (Leicester, 1908) carteri Amerasinghe, 1993 chandi Edwards, 1922 chengi Lien, 1968 communis (Leicester, 1908) complex (Theobald, 1910) covelli Barraud, 1929 demeilloni Mattingly, 1970 funerea (Leicester, 1908) greenii (Theobald, 1905) heterospina Gong & Lu, 1986 himalayensis Edwards, 1922 indica (Theobald, 1905) kana Tanaka, Mizusawa & Saugstad, 1979 kanhsienensis Tung, 1955 lii Wu, 1936 macdonaldi Mattingly, 1957 mattinglyi Thurman, 1959 menglianensis Lu & Gong, 1986 menglianeroides Dong, Dong & Zhou, 2003 persimilis Mattingly, 1970 propinqua Mattingly, 1970 proxima Mattingly, 1970 reidi Mattingly, 1957 ruiliensis Dong, Zhou & Wang, 1997 scanloni Mattingly, 1970 scintillans Ludlow, 1905 taiwanensis Lien, 1968 tengchongensis Dong, Wang & Zhou, 2002 thelmae Mattingly, 1970 viridis Barraud, 1929

Subgenus Mattinglyia Lien, 1968

achaetae (Leicester, 1908) catesi (Lien, 1968) discrepans (Edwards, 1922) lui Gong & Li, 1999 occidentayunnana (Gong & Lu, 1991) tripunctata (Theobald, 1908)

Genus Opifex

Subgenus Nothoskusea Dumbleton, 1962 chathamicus (Dubleton, 1962)

Subgenus Opifex Hutton, 1902 fuscus Hutton, 1902

Genus Psorophora

Subgenus Grabhamia Theobald, 1903

cingulata (Fabricius, 1805) columbiae (Dyar & Knab, 1906) confinnis (Lynch Arribálzaga, 1891) dimidiata Cerqueira, 1943 discolor (Coquillett, 1903) infinis (Dyar & Knab, 1906) insularia (Dyar & Knab, 1906) jamaicensis Theobald, 1901 leucocnemis Martini, 1931 paulli Paterson & Shannon, 1927 pruinosa Martini, 1935 pygmaea (Theobald, 1903) santamarinai Broche, 2000 signipennis (Coquillett, 1904) varinervis Edwards, 1922

Subgenus Janthinosoma Lynch Arribálzaga, 1891

albigenu (Peryassú, 1908) albipes (Theobald, 1907) amazonica Cerqueira, 1960 champerico (Dyar & Knab, 1906) circumflava Cerqueira, 1943 cyanescens (Coquillett, 1902) discrucians (Walker, 1856) ferox (von Humboldt, 1819) fiebrigi Edwards, 1922 forceps Cerqueira, 1939 horrida (Dyar & Knab, 1908) johnstonii (Grabham, 1905) lanei Shannon & Cerqueira, 1943 longipalpus Randolph & O'Neill, 1944 lutzii (Theobald, 1901) mathesoni Belkin & Heinemann, 1975 melanota Cerqueira, 1943 mexicana (Bellardi, 1859) pilosa Duret, 1971 pseudoalbipes Duret, 1971 pseudomelanota Barata & Cotrim, 1971 totonaci Lassmann, 1951 varipes (Coquillett, 1904)

Subgenus Psorophora Robineau-Desvoidy, 1827

ciliata (Fabricius, 1794) cilipes (Fabricius, 1805) holmbergii Lynch Arribálzaga, 1891 howardii Coquillett, 1901 lineata (von Humboldt, 1819) ochripes (Macquart, 1850) pallescens Edwards, 1922 pilipes (Macquart, 1834) saeva Dyar & Knab, 1906 stonei Vargas, 1956

Nomina dubia goeldii (Giles, 1904) marmorata (Philippi, 1865) perterrens (Walker, 1856)

Genus Udaya Thurman, 1954

argyrurus (Edwards, 1934) lucaris Macdonald & Mattingly, 1960 subsimilis (Barraud, 1927)

Genus Verrallina

Subgenus Harbachius Reinert, 1999

abdita (Barraud, 1931) consonensis (Reinert, 1973) fragilis Leicester, 1908 hamistylus (Laffoon, 1946) indecorabilis Leicester, 1908 nobukonis (Yamada, 1932) pahangi (Delfinado, 1968) ramalingami (Reinert, 1974) robertsi (Laffoon, 1946) srilankensis (Reinert, 1977) stunga (Klein, 1973) uniformis (Theobald, 1910) yusafi (Barraud, 1931)

Subgenus Neomacleaya Theobald, 1907

adusta (Laffoon, 1946) agrestis (Barraud, 1931) andamanensis (Edwards, 1922) assamensis Bhattacharyya, Tewari, Prakash, Mohapatra & Mahanta, 2004 atriisimilis (Tanaka & Mizusawa, 1973) atria (Barraud, 1928) campylostylus (Laffoon, 1946) cauta (Barraud, 1928) clavata (Barraud, 1931) comata (Barraud, 1931) comosa (Reinert, 1974) cretata (Delfinado, 1967) cyrtolabis (Edwards, 1928) gibbosa (Delfinado, 1967) harrisonica (Reinert, 1974) hispida (Delfinado, 1967) incertus (Edwards, 1922) indica (Theobald, 1907) johnsoni (Laffoon, 1946) johorensis (Reinert, 1974) komponga (Klein, 1973) lankaensis (Stone & Knight, 1958) latipennis (Delfinado, 1967) leicesteri (Edwards, 1917) macrodixoa (Dyar & Shannon, 1925) margarsen (Dyar & Shannon, 1925) neomacrodixoa (King & Hoogstraal, 1947) nigrotarsis (Ludlow, 1908) notabilis (Delfinado, 1967) nubicola (Laffoon, 1946) panayensis (Ludlow, 1914) petroelephantus (Wijesundara, 1951) philippinensis (Delfinado, 1968) phnoma (Klein, 1973) prioekanensis (Brug, 1931) protuberans (Delfinado, 1967) pseudodiurna (Theobald, 1910) pseudomediofasciata (Theobald, 1910) pseudovarietas (Reinert, 1974) rami (Barraud, 1928) rara (Delfinado, 1968) sabahensis (Reinert, 1974) seculata (Menon, 1950) singularis (Leicester, 1908) sohni (Reinert, 1974) spermathecus (Wijesundara, 1951) torosa (Delfinado, 1967) uncus (Theobald, 190l) vallistris (Barraud, 1928) varietas (Leicester, 1908) virilis Leicester, 1908 yerburyi (Edwards, 1917)

Subgenus Verrallina Theobald, 1903

azureosquamata (Bonne-Wepster, 1948) bifoliata (King & Hoogstraal, 1947) butleri (Theobald, 1901) carmenti (Edwards, 1924) cuccioi (Belkin, 1962) cunninghami (Taylor, 1944) dux (Dyar & Shannon, 1925) embiensis (Huang, 1968) foliformis (King & Hoogstraal, 1947) funerea (Theobald, 1903) iriomotensis (Tanaka & Mizusawa, 1973) killertonis (Huang, 1968) leilae (King & Hoogstraal, 1947) lineata (Taylor, 1914) lugubris (Barraud, 1928) mccormicki (Belkin, 1962) milnensis (King & Hoogstraal, 1947) multifolium (King & Hoogstraal, 1947) obsoleta (Huang, 1968) parasimilis (King & Hoogstraal, 1947) pipkini (Bohart, 1957) quadrifolium (Brug, 1934) quadrispinata (King & Hoogstraal, 1947) reesi (King & Hoogstraal, 1947) sentania (King & Hoogstraal, 1947) similis (Theobald, 1910) simpla (King & Hoogstraal, 1947) trispinata (King & Hoogstraal, 1947) vanapa (Huang, 1968) variabilis (Huang, 1968)

Nomen dubium perdita Leicester, 1908

Genus Zeugnomyia Leicester, 1908

aguilari Baisas & Feliciano, 1953 fajardoi Baisas & Feliciano, 1953 gracilis Leicester, 1908 lawtoni Baisas, 1946 Table XX. Proposed Informal species groups listed as subgenera in Reinert et al. (2009)

Reinert et al. (2009) Herein: genus / subgenus Suggested informal group Species prefix or previous group name Collessius (Alloeomyia) Aedes (Collessius) Alloeomyia banksi Edwards pseudotaeniatus (Giles) tonkinensis Galliard & Ngu Collessius Collessius Aedes Collessius Collessius elsiae (Barraud) elsiae vicarious Lien ( ) ( ) hatorii Yamada macdougalli Edwards macfarlanei (Edwards) ningheensis Lei shortti (Barraud) Georgecraigius Aedes (Georgecraigius Georgecraigius atropalpus (Coquillet) (Georgecraigius) ) epactius Dyar & Knab Georgecraigius Aedes (Georgecraigius) Horsfallius fluviatilis (Lutz) (Horsfallius) Hopkinsius (Hopkinsius) Aedes (Hopkinsius) Hopkinsius barnardi Edwards embuensis Edwards ingrami Edwards nyasae Edwards wellmanii (Theobald) Hopkinsius (Yamada) Aedes (Hopkinsius) Yamada albocinctus (Barraud) seoulensis Yamada Macleaya Aedes (Macleaya) Chaetocruiomyia calabyi Marks (Chaetocruiomyia) elchoensis Taylor humeralis Edwards macmillani Marks moloiensis Taylor spinosipes Edwards tulliae (Taylor) wattensis Taylor Macleaya (Macleaya) Aedes (Macleaya) Macleaya littlechildi Taylor stoneorum Marks tremulus (Theobald) Mucidus (Mucidus) Aedes (Mucidus) Mucidus alternans (Westwood) ferinus Knight grahamii (Theobald) laniger (Wiedemann) lucianus Muspratt mucidus (Karsch) nigerrimus (Theobald) quasiferinus Mattingly scatophagoides (Theobald) sudanensis (Theobald) tonkingi Gebert Mucidus (Pardomyia) Aedes (Mucidus) Pardomyia aurantius (Theobald) aurantius chrysogaster (Taylor) nigrescens

quadripunctis (Ludlow) Ochlerotatus (Buvirilia) Aedes (Ochlerotatus) Buvirilia edgari(Edwards) Stone & Rosen Ochlerotatus Aedes (Ochlerotatus) Chrysoconops bimaculatus (Coquillett) (Crysoconops) fulvus (Wiedemann) fulvus pallens (Ross) jorgi Carpintero & Lequizamón pennai Antunes & Lane stigmaticus Edwards Ochlerotatus (Culicada) Aedes (Ochlerotatus) Culicada canadensis (Theobald) canadensis mathesoni Middlekauff Ochlerotatus (Culicelsa) Aedes (Ochlerotatus) Culicelsa mitchellae (Dyar) nigromaculis (Ludlow) sollicitans (Walker) taeniorhynchus (Wiedemann) Ochlerotatus (Empihals) Aedes (Ochlerotatus) Empihals vigilax (Skuse) vigilax ludlowae (R. Blanchard) vigilax vansomerenae Mattingly Ochlerotatus (Gilesia) Aedes (Ochlerotatus) Gilesia aculeatus (Theobald) mcdonaldi Belkin purpuraceus Brug Ochlerotatus (Juppius) Aedes (Ochlerotatus) Juppius caballus (Theobald) chelli (Edwards) juppi McIntosh Ochlerotatus Aedes (Ochlerotatus) Lepidokeneon spilotus Marks (Lepidokeneon) stricklandi (Edwards) turneri Marks Ochlerotatus Aedes (Ochlerotatus) Ochlerotatus angustivittatus Dyar & (Ochlerotatus) Knab atactavittatus Arnell auratus Grabham bogotanus (Arnell) comitatus Arnell condolescens Dyar & Knab crinifer (Theobald) deficiens Arnell euplocamus Dyar & Knab incomptus Arnell infirmatus Dyar & Knab meprai Martinez & Prosen obturbator Dyar & Knab patersoni Shannon & del Ponte pectinatus Arnell phaeonotus (Arnell raymondi del Ponte, Castro & Garcia ) rhyacophilus da Costa Lima scapularis (Rondani) synchytus Arnell thelcter Dyar tortilis (Theobald) trivittatus (Coquillett) Ochlerotatus (Pholeomyia) Aedes (Ochlerotatus) Pholeomyia calcariae Marks clelandi (Taylor) flavifrons (Skuse) purpuriventris Edwards Ochlerotatus (Protoculex) Aedes (Ochlerotatus) Protoculex aenigmaticus Cerqueira & Costa atlanticus Dyar & Knab dupreei (Coquillett) eucephalaeus Dyar hastatus Dyar nubilis Theobald oligopistus Dyar pertinax Grabham serratus (Theobald) tormentor Dyar & Knab Ochlerotatus Aedes (Ochlerotatus) Pseudoskusea bancroftianus Edwards (Pseudoskusea) culiciformis (Theobald) multiplex (Theobald) postspiraculosus Dobrotworsky Ochlerotatus (Rusticoidus) Aedes (Ochlerotatus) Rusticoidus albescens Edwards bicristatus Thurman & Winkler krymmontanus Alekseev lepidonotus Edwards provocans (Walker) quasirusticus Torres Cañamares refiki Medschid rusticus (Rossi) rusticus subtrichurus Martini subdiversus Martini Ochlerotatus (Woodius) Aedes (Ochlerotatus) Woodius diantaeus Howard, Dyar & Knab intrudens Dyar Petermattinglyius Aedes (Petermattinglyius) Aglaonotus whartoni Mattingly (Aglaonotus) Petermattinglyius Aedes (Petermattinglyius) Petermattinglyius franciscoi Mattingly (Petermattinglyius) iyengari Edwards punctipes Edwards scanloni Reinert Stegomyia (Actinothrix) Aedes (Stegomyia) Actinothrix edwardsi (Barraud) robinsoni Belkin seampi Huang tulagiensis Edwards Stegomyia (Bohartius) Aedes (Stegomyia) Pandani Subgroup of agrihanensis Bohart Bohart (1957) neopandani Bohart pandani Stone rotanus Bohart & Ingram saipanensis Stone Stegomyia (Heteraspidion) Aedes (Stegomyia) Annandalei Subgroup of annandalei (Theobald) Huang (1977) craggi (Barraud) Stegomyia (Huangmyia) Aedes (Stegomyia) Mediopunctatus Subgroup malikuli Huang of Huang (1977) mediopunctatus (Theobald) mediopunctatus sureilensis Barraud perplexus (Leicester) Stegomyia (Mukwaya) Aedes (Stegomyia) Simpsoni Group of Huang bromeliae (Theobald) (2004) gandaensis Huang josiahae Huang kivuensis Edwards lilii (Theobald) sampi Huang simpsoni (Theobald) strelitziae Muspratt subargenteus Edwards woodi Edwards Stegomyia (Stegomyia) Aedes (Stegomyia) Aegypti Group of Huang aegypti (Linnaeus) (2004) aegypti formsus (Walker) Stegomyia (Xyele) Aedes (Stegomyia) Desmotes Subgroup of desmotes (Giles) Huang (1977) Stegomyia (Zoromorphus) Aedes (Stegomyia) Zoromorphus futunae Belkin

S3 Appendix. Updated mosquito generic (two letters) and subgeneric (three letters) abbreviations. Based on: Reinert JF (2009) List of abbreviations for currently valid generic-level taxa in family Culicidae (Diptera). European Mosquito Bulletin 27: 68-76.

* = Used here for the first time

Subfamily Anophelinae

Genus Anopheles Meigen = An. Subgenus Anopheles Meigen = Ano. Subgenus Baimaia Harbach, Rattanarithikul & Harrison = Bmi. Subgenus Cellia Theobald= Cel. Subgenus Kerteszia Theobald = Ker. Subgenus Lophopodomyia Antunes = Lph. Subgenus Nyssorhynchus Blanchard = Nys. Subgenus Stethomyia Theobald = Ste. Genus Bironella Theobald = Bi. Genus Chagasia Cruz = Ch.

Subfamily Culicinae

Tribe Aedeomyiini

Genus Aedeomyia Theobald = Ad. Subgenus Aedeomyia Theobald = Ady. Subgenus Lepiothauma Enderlein = Lpi.

Tribe Aedini

Genus Aedes Meigen = Ae. Subgenus Abraedes Zavortink = Abr. Subgenus Acartomyia Theobald = Acy. Subgenus Aedimorphus Theobald = Adm. Subgenus Alanstonea Mattingly = Ala. Subgenus Albuginosus Reinert = Alb. Subgenus Ayurakitia Thurman = Ayu. Subgenus Aztecaedes Zavortink = Azt. Subgenus Belkinius Reinert = Blk. Subgenus Bifidistylus Reinert, Harbach & Kitching = Bif.* Subgenus Borichinda Harbach & Rattanarithikul = Bor.* Subgenus Bothaella Reinert = Bot. Subgenus Bruceharrisonius Reinert = Brh. Subgenus Cancraedes Edwards = Can. Subgenus Catageiomyia Theobald = Cat.* Subgenus Catatassomyia Dyar & Shannon = Cts.* Subgenus Christophersiomyia Barraud = Chr. Subgenus Collessius Reinert, Harbach & Kitching = Col. Subgenus Cornetius Huang = Cor. Subgenus Dahliana Reinert, Harbach & Kitching = Dah.* Subgenus Danielsia Theobald = Dan.* Subgenus Dendroskusea Edwards = Dsk.* Subgenus Diceromyia Theobald = Dic. Subgenus Dobrotworskyius Reinert, Harbach & Kitching = Dob.* Subgenus Downsiomyia Vargas = Dow.* Subgenus Edwardsaedes Belkin = Edw. Subgenus Elpeytonius Reinert, Harbach & Kitching = Elp.* Subgenus Finlaya Theobald = Fin. Subgenus Fredwardsius Reinert = Fre. Subgenus Georgecraigius Reinert, Harbach & Kitching = Grg.* Subgenus Geoskusea Edwards = Geo. Subgenus Gilesius Reinert, Harbach & Kitching = Gil.* Subgenus Gymnometopa Coquillett = Gym. Subgenus Halaedes Belkin = Hal. Subgenus Himalaius Reinert, Harbach & Kitching = Him.* Subgenus Hopkinsius Reinert, Harbach & Kitching = Hop.* Subgenus Howardina Theobald = How. Subgenus Huaedes Huang = Hua. Subgenus Hulecoeteomyia Theobald = Hul.* Subgenus Indusius Edwards = Ind. Subgenus Isoaedes Reinert = Isa. Subgenus Jarnellius Reinert, Harbach & Kitching = Jar.* Subgenus Jihlienius Reinert, Harbach & Kitching = Jih.* Subgenus Kenknightia Reinert = Ken. Subgenus Kompia Aitken = Kom. Subgenus Leptosomatomyia Theobald = Lep. Subgenus Levua Stone & Bohart = Lev. Subgenus Lewnielsenius Reinert, Harbach & Kitching = Lew. Subgenus Lorrainea Belkin = Lor. Subgenus Luius Reinert, Harbach & Kitching = Lui.* Subgenus Macleaya Theobald = Mac. Subgenus Molpemyia Theobald = Mol. Subgenus Mucidus Theobald = Muc. Subgenus Neomelaniconion Newstead = Neo. Subgenus Nyctomyia Harbach & Linton = Nyc. Subgenus Ochlerotatus Lynch Arribálzaga = Och. Subgenus Paraedes Edwards = Par. Subgenus Patmarksia Reinert, Harbach & Kitching = Pat.* Subgenus Petermattinglyius Reinert, Harbach & Kitching = Pet.* Subgenus Phagomyia Theobald = Phg.* Subgenus Polyleptiomyia Theobald = Pmt.* Subgenus Pseudarmigeres Stone & Knight = Psa. Subgenus Rampamyia Reinert, Harbach & Kitching = Ram.* Subgenus Rhinoskusea Edwards = Rhi. Subgenus Sallumia Reinert, Harbach & Kitching = Sal. Subgenus Scutomyia Theobald = Sct. Subgenus Skusea Theobald = Sku. Subgenus Stegomyia Theobald = Stg. Subgenus Tanakaius Reinert, Harbach & Kitching = Tan.* Subgenus Tewarius Reinert = Tew.* Subgenus Vansomerenis Reinert, Harbach & Kitching = Van.

Genus Armigeres Theobald = Ar. Subgenus Armigeres Theobald = Arm. Subgenus Leicesteria Theobald = Lei.

Genus Eretmapodites Theobald = Er.

Genus Haemagogus Williston = Hg. Subgenus Conopostegus Dyar = Con. Subgenus Haemagogus Williston = Hag.

Genus Heizmannia Ludlow = Hz. Subgenus Heizmannia Ludlow = Hez. Subgenus Mattinglyia Lien = Mat.

Genus Opifex Hutton = Op. Subgenus Nothoskusea Dumbleton = Not. Subgenus Opifex Hutton = Opi.

Genus Psorophora Robineau-Desvoidy = Ps. Subgenus Grabhamia Theobald = Gra. Subgenus Janthinosoma Lynch Arribálzaga = Jan. Subgenus Psorophora Robineau-Desvoidy = Pso.

Genus Udaya Thurman = Ud.

Genus Verrallina Theobald = Ve. Subgenus Harbachius Reinert= Har. Subgenus Neomacleaya Theobald = Nma. Subgenus Verrallina Theobald = Ver.

Genus Zavortinkius Reinert = Za.

Genus Zeugnomyia Leicester = Ze. Tribe Culicini

Genus Culex Linnaeus = Cx. Subgenus Acalleomyia Leicester = Aca. Subgenus Acallyntrum Stone & Penn = Acl. Subgenus Aedinus Lutz = Ads. Subgenus Afroculex Danilov = Afc. Subgenus Allimanta Casal & Garcia = Alm. Subgenus Anoedioporpa Dyar = And. Subgenus Barraudius Edwards = Bar. Subgenus Belkinomyia Adames & Galindo = Bel. Subgenus Carrollia Lutz = Car. Subgenus Culex Linnaeus = Cux. Subgenus Culiciomyia Theobald = Cui. Subgenus Eumelanomyia Theobald = Eum. Subgenus Kitzmilleria Danilov = Kit. Subgenus Lasiosiphon Kirkpatrick = Las. Subgenus Lophoceraomyia Theobald = Lop. Subgenus Maillotia Theobald = Mai. Subgenus Melanoconion Theobald = Mel. Subgenus Micraedes Coquillett = Mca. Subgenus Microculex Theobald = Mcx. Subgenus Neoculex Dyar = Ncx. Subgenus Nicaromyia Broche & Rodríguez Rodríguez = Nic. Subgenus Oculeomyia Theobald = Ocu. Subgenus Phenacomyia Harbach & Peyton = Phc. Subgenus Phytotelmatomyia Rossi & Harbach = Phy. Subgenus Sirivanakarnius Tanaka = Sir. Subgenus Tinolestes Coquillett = Tin.

Genus Deinocerites Theobald = De.

Genus Galindomyia Stone & Barreto = Ga.

Genus Lutzia Theobald = Lt. Subgenus Insulalutzia Tanaka = Ilt. Subgenus Metalutzia Tanaka = Mlt. Subgenus Lutzia Theobald = Lut.

Tribe Culisetini

Genus Culiseta Felt = Cs. Subgenus Allotheobaldia Broelemann = All. Subgenus Austrotheobaldia Dobrotworsky = Aut. Subgenus Climacura Howard, Dyar & Knab = Cli. Subgenus Culicella Felt = Cuc. Subgenus Culiseta Felt = Cus. Subgenus Neotheobaldia Dobrotworsky = Net. Subgenus Theomyia Edwards = Thm.

Tribe Ficalbiini

Genus Ficalbia Theobald = Fi.

Genus Mimomyia Theobald = Mi. Subgenus Etorleptiomyia Theobald = Eto. Subgenus Ingramia Edwards = Ing. Subgenus Mimomyia Theobald = Mim.

Tribe Hodgesiini

Genus Hodgesia Theobald = Ho.

Tribe Mansoniini

Genus Coquillettidia Dyar = Cq. Subgenus Austromansonia Belkin = Aus. Subgenus Coquillettidia Dyar = Coq. Subgenus Rhynchotaenia Brethés = Rhy.

Genus Mansonia Blanchard = Ma. Subgenus Mansonia Blanchard = Man. Subgenus Mansonioides Theobald = Mnd.

Tribe Orthopodomyiini

Genus Orthopodomyia Theobald = Or.

Tribe Sabethini

Genus Isostomyia Coquillett = Is.

Genus Johnbelkinia Zavortink = Jb.

Genus Kimia Vu Duc Huong & Harbach = Km.

Genus Limatus Theobald = Li.

Genus Malaya Leicester = Ml.

Genus Maorigoeldia Edwards = Mg. Genus Onirion Harbach & Peyton = On.

Genus Runchomyia Theobald = Ru. Subgenus Ctenogoeldia Edwards = Cte. Subgenus Runchomyia Theobald = Run.

Genus Sabethes Robineau-Desvoidy = Sa. Subgenus Davismyia Lane & Cerqueira = Dav. Subgenus Peytonulus Harbach = Pey. Subgenus Sabethes Robineau-Desvoidy = Sab. Subgenus Sabethinus Lutz = Sbn. Subgenus Sabethoides Theobald = Sbo.

Genus Shannoniana Lane & Cerqueira = Sh.

Genus Topomyia Leicester = To. Subgenus Miyagiella Harbach = Myg. Subgenus Suaymyia Thurman = Sua. Subgenus Topomyia Leicester = Top.

Genus Trichoprosopon Theobald = Tr.

Genus Tripteroides Giles = Tp. Subgenus Polylepidomyia Theobald = Pol. Subgenus Rachionotomyia Theobald = Rah. Subgenus Rachisoura Theobald = Rac. Subgenus Tricholeptomyia Dyar & Shannon = Tri. Subgenus Tripteroides Giles = Trp.

Genus Wyeomyia Theobald = Wy. Subgenus Antunesmyia Lane & Cerqueira = Ant. Subgenus Caenomyiella Harbach & Peyton = Cae. Subgenus Cruzmyia Lane & Cerqueira = Cru. Subgenus Decamyia Dyar = Dec. Subgenus Dendromyia Theobald = Den. Subgenus Dodecamyia Dyar = Dod. Subgenus Exallomyia Harbach & Peyton = Exm. Subgenus Hystatomyia Dyar = Hys. Subgenus Menolepis Lutz = Men. Subgenus Nunezia Dyar = Nuz. Subgenus Phoniomyia Theobald = Pho. Subgenus Prosopolepis Lutz = Prl. Subgenus Spilonympha Motta & Lourenço-de-Oliveira = Spi. Subgenus Wyeomyia Theobald = Wyo. Subgenus Zinzala Zavortink = Zin.

Tribe Toxorhynchitini

Genus Toxorhynchites Theobald = Tx. Subgenus Afrorhynchus Ribeiro = Afr. Subgenus Ankylorhynchus Lutz = Ank. Subgenus Lynchiella Lahille = Lyn. Subgenus Toxorhynchites Theobald = Tox.

Tribe Uranotaeniini

Genus Uranotaenia Lynch Arribálzaga = Ur. Subgenus Pseudoficalbia Theobald = Pfc. Subgenus Uranotaenia Lynch Arribálzaga = Ura.

S1 Fig. Valid generic level names and graphical timeline of nomenclature combination changes in Aedine taxa pre-2000 to date. Genera are indicated in blue, subgenera in green and synonomy (Syn.) in red. Author abbreviations include: D & S = Dyar and Shannon; H & R = Harbach & Rattanarithikul; R-D = Robineau-Desvoidy R, H & K = Reinert, Harbach & Kitching; S & P = Shevchenko & Prudkina. Generic abbreviations follow those in Reinert, Harbach & Kitching (2009) and herein as listed in S3 Appendix. *Nyctomyia is a replacement name for the originally pre-occupied genus name Nyx (Harbach et al., Zootaxa, 3683(2), 159-177 (2013).

Aedini valid generic Pre- 2000 2003 2004 2006 2007 2008 2009 2013 Herein level names 2000 Abraedes Ae. Oc. (Abr.) Abraedes Aedes (Abraedes) Zavortink 1970 (Abr.) Acartomyia Oc. Syn. Ae. (Och.) Acartomyia Aedes (Acartomyia) Theobald 1903 (Acy.) Aedes Ae. Aedes Meigen 1818 Aedimorphus Ae. (Adm.) Aedimorphus Aedes (Aedimorphus) Theobald 1903 Aglaonotus Petermattinglius Syn. Aedes

R, H & K 2009 (Aglaonotus) (Petermattinglius) Alloeomyia Syn. Aedes Collesius (Alloeomyia) R, H & K 2008 (Collessius) Alanstonea Ae. (Ala.) Alanstonea Aedes (Alanstonea) Mattingly 1960 Albuginosus Ae. (Alb.) Albuginosus Aedes (Albuginosus) Reinert 1987 Armigeres Armigeres Armigeres Theobald 1901 Ayurakitia Ae. Ayurakitia Aedes (Ayurakitia) Thurman 1954 (Ayu.) Aztecaedes Ae. Oc. (Azt.) Aztecaedes Aedes (Aztecaedes) Zavortink 1972 (Azt.) Belkinius Ae. (Blk.) Belkinius Aedes (Belkinius) Reinert 1982 Bifidistylus Bifidistylus Aedes (Bifidistylus) R, H & K 2009 Bohartius Stegomyia Syn. Aedes

R, H & K 2009 (Bohartius) (Stegomyia) Borichinda Borichinda Aedes (Borichinda) H & R 2007 Bothaella Ae. (Bot.) Bothaella Aedes (Bothaella) Reinert 1973 Bruceharrisonius Aedes Oc. (Brh.) Bruceharrisonius Reinert 2003 (Bruceharrisonius) Buvirilia Syn. Aedes Oc. (Buvirilia) R, H & K 2008 (Ochlerotatus) Cancraedes Ae. (Can.) Cancraedes Aedes (Cancraedes) Edwards 1929 Catageiomyia Aedes Syn. Ae. (Adm.) Catageiomyia Theobald 1903 (Catageiomyia) Catatassomyia Aedes Syn. Ae. (Stg.) Catatassomyia D & S 1925 (Catatassomyia) Chaetocruiomyia Ae. Syn. Aedes Oc. (Chaetocruiomyia) Theobald 1910 (Cha.) (Ochlerotatus) Christophersiomyia Aedes Ae. (Chr.) Christophersiomyia Barraud 1923 (Christophersiomyia) Chrysoconops Syn. Aedes Syn. Ae. (Och.) Oc. (Chrysoconops) Goeldi 1905 (Ochlerotatus) Collessius Collessius Aedes (Collessius) R, H & K 2006 Aedini valid generic Pre- 2000 2003 2004 2006 2007 2008 2009 2013 Herein level names 2000 Conopostegus Haemagogus Haemagogus (Conopostegus) Dyar 1925 (Conopostegus) Cornetius Cornetius Aedes (Cornetius) Huang 2005 Culicada Syn. Aedes Syn. Ae. (Och.) Oc. (Culicada) Felt 1904 (Ochlerotatus) Culicelsa Syn. Aedes Syn. Ae. (Och.) Oc. (Culicelsa) Felt 1904 (Ochlerotatus) Dahliana Dahliana Aedes (Dahliana) R, H & K 2006 Danielsia Syn. Ae. (Fin.) Danielsia Aedes (Danielsia) Theobald 1904 Dendroskusea Aedes Syn. Ae. (Dic.) Dendroskusea Edwards 1929 (Dendroskusea) Diceromyia Ae. (Dic.) Diceromyia Aedes (Diceromyia) Theobald 1911 Dobrotworskyius Aedes Dobrotworskyius R, H & K 2006 (Dobrotworskyius) Downsiomyia Aedes Syn. Ae. (Fin.) Downsiomyia Vargas 1950 (Downsiomyia) Edwardsaedes Aedes Ae. (Edw.) Edwardsaedes Belkin 1962 (Edwardsaedes) Elpeytonius Elpeytonius Aedes (Elpeytonius) R, H & K 2009 Empihals Syn. Aedes Oc. (Empihals) R, H & K 2008 (Ochlerotatus) Eretmapodites Eretmapodites Eretmapodites Theobald 1901 Finlaya Ae. Oc. (Fin.) Finlaya Aedes (Finlaya) Theobald 1903 (Fin.) Fredwardsius Ae. (Fre.) Fredwardsius Aedes (Fredwardsius) Reinert 2000 Georgecraigius Aedes Georgecraigius R, H & K 2006 (Georgecraigius) Geoskusea Ae. Oc. Oc. (Geo.) Geoskusea Geoskusea Aedes (Geoskusea) Edwards 1929 (Geo.) (Geo.) Gilesia Syn. Aedes Syn. Ae. (Och.) Oc. (Gilesia) Theobald 1903 (Ochlerotatus) Gilesius Gilesius Aedes (Gilesius) R, H & K 2006 Grabhamia Psorophora Psorophora (Grabhamia) Theobald 1903 (Grabhamia) Gymnometopa Ae. Aedes Oc. (Gym.) Gymnometopa Coquillett 1906 (Gym.) (Gymnometopa) Haemagogus Haemagogus Haemagogus Williston 1896 Halaedes Ae. Oc. (Hal.) Halaedes Aedes (Halaedes) Belkin 1962 (Hal.) Heizmannia Heizmannia Heizmannia Ludlow 1905 Heteraspidion Stegomyia Syn. Aedes

R, H & K 2009 (Heteraspidion) (Stegomyia) Aedini valid generic Pre- 2000 2003 2004 2006 2007 2008 2009 2013 Herein level names 2000 Himalaius Himalaius Aedes (Himalaius) R, H & K 2006 Hopkinsius Hopkinsius Aedes (Hopkinsius) R, H & K 2008 Horsfallius Syn. Aedes Georgecraigius (Horsfallius) R, H & K 2006 (Georgecraigius) Howardina Ae. Oc. (How.) Howardina Aedes (Howardina) Theobald 1903 (How.) Huaedes Ae. (Hua.) Huaedes Aedes (Huaedes) Huang 1968 Huangmyia Stegomyia Syn. Aedes

R, H & K 2009 (Huangmyia) (Stegomyia) Hulecoeteomyia Aedes Syn. Ae. (Fin.) Hulecoeteomyia Theobald 1904 (Hulecoeteomyia) Indusius Ae. (Ind.) Indusius Aedes (Indusius) Edwards 1934 Isoaedes Ae. (Isa.) Isoaedes Aedes (Isoaedes) Reinert 1979 Janthinosoma Psorophora Lynch-Arribalzaga Psorophora (Janthinsoma) (Janthinosoma) 1891 Jarnellius Jarnellius Aedes (Jarnellius) R, H & K 2006 Jihlienius Jihlienius Aedes (Jihlienius) R, H & K 2006 Juppius Syn. Aedes Oc. (Juppius) R, H & K 2009 (Ochlerotatus) Kenknightia Ae. Oc. (Ken.) Kenknightia Aedes (Kenknightia) Reinert 1990 (Ken.) Kompia Ae. Oc. (Kom.) Kompia Aedes (Kompia) Aitken 1941 (Kom.) Leicesteria Armigeres Armigeres (Leicesteria) Theobald 1904 (Leicesteria) Lepidokeneon Syn. Aedes Oc. (Lepidokeneon) R, H & K 2009 (Ochlerotatus) Leptosomatomyia Aedes Ae. (Lep.) Leptosomatomyia Theobald 1905 (Leptosomatomyia) Levua Stone & Ae. Oc. Oc. (Lev.) Levua Levua Aedes (Levua) Bohart 1944 (Lev.) (Lev.) Lewnielsenius Aedes Jarnellius (Lew.) Lewnielsenius R, H & K 2006 (Lewnielsenius) Lorrainea Ae. (Lor.) Lorrainea Aedes (Lorrainea) Belkin 1962 Luius Luius Aedes (Luius) R, H & K 2008 Macleaya Ae. Oc. (Mac.) Macleaya Aedes (Macleaya) Theobald 1903 (Mac.) Mattinglyia Heizmannia Heizmannia (Mattinglyia) Lien 1968 (Mattinglyia) Syn. Molpemyia Ae. Oc. (Mol.) Molpeymia Aedes (Molpemyia) Theobald 1910 (Fin.) Mucidus Ae. Oc. (Muc.) Mucidus Aedes (Mucidus) Theobald 1901 (Muc.) Aedini valid generic Pre- 2000 2003 2004 2006 2007 2008 2009 2013 Herein level names 2000 Mukwaya Stegomyia Syn. Aedes

R, H & K 2009 (Mukwaya) (Stegomyia) Neomacleaya Verrallina Verrallina (Neomacleaya) Theobald 1907 (Neomacleaya) Neomelaniconion Aedes Ae. (Neo.) Neomelanoconion Newstead 1907 (Neomelanoconion) Nothoskusea Ae. Oc. (Not.) Opifex (Not.) Opifex (Nothoskusea) Dumbleton 1962 (Not.) Nyctomyia Harbach Nyctomyia* Aedes (Nyctomyia) & Linton 2013 Ochlerotatus Lynch- Ae. Ochlerotatus Aedes (Ochlerotatus) Arribalzaga 1891 (Och.) Opifex Opifex Opifex Hutton 1902 Paraedes Ae. (Par.) Paraedes Aedes (Paraedes) Edwards 1934 Pardomyia Syn. Ae. (Muc.) Mucidus (Pardomyia) Syn. Aedes (Mucidus) Theobald 1907 Patmarksia Patmarksia Aedes (Patmarksia) R, H & K 2006 Petermattinglius Aedes Petermattinglius R, H & K 2009 (Petermattinglius) Phagomyia Syn. Ae. (Fin.) Phagomyia Aedes (Phagomyia) Theobald 1905 Pholeomyia Syn. Aedes Oc. (Pholeomyia) R, H & K 2008 (Ochlerotatus) Polyleptiomyia Aedes Syn. Ae. (Adm.) Polyleptiomyia Theobald 1905 (Polyleptiomyia) Protoculex Syn. Aedes Syn. Ae. (Och.) Oc. (Protoculex) Felt 1904 (Ochlerotatus) Protomacleaya Ae. Syn. Aedes Oc. (Protomacleaya) Theobald 1907 (Pro.) (Ochlerotatus) Pseudarmigeres Aedes Ae. (Pse.) Pseudarmigeres Stone & Knight 1956 (Pseudarmigeres) Pseudoskusea Ae. Syn. Aedes Oc. (Psk.) Pseudoskusea Oc.(Psk.) Theobald 1907 (Psk.) (Ochlerotatus) Psorophora Psorophora Psorophora R-D 1827 Rampamyia Rampamyia Aedes (Rampamyia) R, H & K 2006 Rhinoskusea Ae. Oc. (Rhi.) Rhinoskusea Aedes (Rhinoskusea) Edwards 1929 (Rhi.) Rusticoidus Ae. Syn. Aedes Oc. (Rus.) S & P 1973 (Rus.) (Ochlerotatus) Sallumia Oc. Sallumia Aedes (Sallumia) R, H & K 2008 (Sal.) Scutomyia Syn. Ae. (Stg.) Scutomyia Aedes (Scutomyia) Theobald 1904 Skusea Ae. (Sku.) Skusea Aedes (Skusea) Theobald 1903 Aedini valid generic Pre- 2000 2003 2004 2006 2007 2008 2009 2013 Herein level names 2000 Stegomyia Ae. (Stg.) Stegomyia Aedes (Stegomyia) Theobald 1901 Tanakius Tanakius Aedes (Tanakius) R, H & K 2004 Tewarius Tewarius Aedes (Tewarius) R, H & K 2006 Udaya Udaya Udaya Thurman 1954 Vansomerenis Aedes Vansomerensis R, H & K 2006 (Vansomerensis) Verrallina Verrallina Verrallina Theobald 1903 Woodius Syn. Aedes Oc. (Woodius) R, H & K 2009 (Ochlerotatus) Xyele Syn. Aedes Stegomyia (Xyele) R, H & K 2009 (Stegomyia) Yamada Syn. Aedes Hopkinsinsius (Yamada) R, H & K 2008 (Hopkinsius) Zavortinkius Ae. Oc. (Zav.) Zavortinkius Aedes (Zavortinkius) Reinert 1999 (Zav.) Zeugnomyia Zeugnomyia Zeugnomyia Leicester 1908 Zoomorphus Ochlerotatus Syn. Aedes

R, H & K 2009 (Zoromorphus) (Ochlerotatus)

Culiseta (Culiseta) inornata Culex (Culex) quinquefasciatus Orthopodomyia signifera Mansonia (Mansonia) titillans

32 97 Psorophora (Grabhamia) jamaicensis Psorophora (Grabhamia) columbiae 100 100 Psorophora (Psorophora) howardii 98 Psorophora (Psorophora) ciliata 99 Psorophora (Janthinosoma) ferox Psorophora (Janthinosoma) cyanescens

100 100 Mucidus (Pardomyia) quadripunctis 99 Mucidus (Pardomyia) aurantius aurantius 100 Mucidus (Mucidus) laniger Mucidus (Mucidus) alternans Fredwarsius vittatus Aedimorphus domesticus

3 Skusea pembaensis Indusius pulverulentus

100 Scutomyia arboricola 62 Scutomyia albolineata 13 Isoaedes cavaticus Borichinda cavernicola

97 Howardina walkeri Howardina sexlineata

100 Downsiomyia nivea Downsiomyia leonis

97 Dahliana geniculata Dahliana echinus Molpemyia pecuniosa 21 100 Rampamyia notoscripta Rampamyia albilabris Halaedes australis 87 59 Opifex (Opifex) fuscus Opifex (Nothoskusea) chathamicus ‘Ochlerotatus’ (‘Protomacleaya’) brelandi sensu auctorum 1 22 ‘Ochlerotatus’ (’Protomacleaya’) zoosophus sensu auctorum ‘Ochlerotatus’ (’Protomacleaya’) triseriatus sensu auctorum 72 ‘Ochlerotatus’ (’Finlaya’) biocellatus sensu auctorum 6 50 ‘Ochlerotatus’ (’Finlaya’) keefei ‘Ochlerotatus’ (’Finlaya’) candidoscutellum Luius fengi 29 100 Macleya (Macleaya) tremula Macleaya (Chaetocruiomyia) wattensis Dendroskusea periskelata 14 80 Dendroskusea reginae Dendroskusea micropterus Catageiomyia irritans 27 25 Catageiomyia tarsalis Catageiomyia argenteopunctata

100 Tewarius reubenae 8 Tewarius agastyai 100 Christophersiomyia thomsoni Christophersiomyia gombakensis ‘Ochlerotatus’ (’Finlaya’) oreophilus sensu auctorum 26 Jihliensius chungi 43 97 Phagomyia lophoventralis Phagomyia gubernatoris

100 Finlaya poicilia 57 Finlaya kochi 69 Danielsia harperi Danielsia albotaeniata

100 Diceromyia taylori 10 Diceromyia furcifer 100 Ayurakitia peytoni Ayurakitia griffithi

99 Albuginosus ngong 4 Albuginosus marshallii 97 Elpeytonius simulans Elpeytonius apicoannulatus Bifidistylus lamborni 19 8 Polyleptiomyia albocephala 20 94 Aedimorphus quasiunivittatus Aedimorphus gibbinsi ‘Ochlerotatus’ (‘Finlaya’) crossi sensu auctorum

28 93 Hopkinsius (Yamada) seoulensis 63 Hopkinsius (Yamada) albocinctus 37 Hopkinsius (Hopkinsius) ingrami Hopkinsius (Hopkinsius) embuensis

99 Edwardsaedes imprimens Edwardsaedes bekkui 2 Aedimorphus vexans vexans 54 24 Aedimorphus dentatus Aedimorphus cumminsii ‘Ochlerotatus’ (’Protomacleaya’) knabi sensu auctorum 0 ‘Ochlerotatus’ (’Protomacleaya’) burgeri sensu auctorum 2 ‘Ochlerotatus’ (’Protomacleaya’) kompi sensu auctorum 16 ‘Ochlerotatus’ (’Pro.’) homoeopus sensu auctorum 84 19 ‘Ochlerotatus’ (’Pro.’) terrens sensu auctorum ‘Ochlerotatus’ (’Pro.’) galindoi sensu auctorum Catatassomyia meronephada

100 Udaya lucaris Udaya argyrurus 10 100 Bothaella helenae Bothaella eldridgei Belkinius aurotaeniatus 8 99 Zeugnomyia lawtoni Zeugnomyia gracilis Himalaius gilli 68 100 Bruceharrisonius greenii Bruceharrisonius alektorovi 13 95 Vansomerensus pulchrithorax Vansomerensus luteostriata 40 99 Zavortinkius longipalpis 10 Zavortinkius fulgens 99 Kenknightia harbachi Kenknightia dissimilis

61 Leptosomatomyia aurimargo Huaedes wauensis Alanstonea brevitibia 21 59 100 Eretmapodites quinquevittatus Eretmapodites chrysogaster 70 99 Armigeres (Leicesteria) longipalpis 86 Armigeres (Leicesteria) flavus 91 Armigeres (Armigeres) subalbatus Armigeres (Armigeres) breinli Petermattinglyius whartoni 88 Petermattinglyius franciscoi 41 99 Petermattinglyius scanloni 5 Petermattinglyius iyengari 88 Pseudarmigeres michaelikati Pseudarmigeres argenteoventralis dunni 36 89 Heizmannia (Mattinglyia) catesi 99 Heizmannia (Mattinglyia) achaetae 100 Heizmannia (Heizmannia) scintillans Heizmannia (Heizmannia) complex

100 Lorrainea dasyorrhus Lorrainea amesii Cancraedes masculinus 8 2 100 55 Cancraedes penghuensis Cancraedes cancricomes 52 97 Geoskusea longiforceps Geoskusea baisasi 41 Sallumia hortator 16 Levua geoskusea

100 Rhinoskusea wardi Rhinoskusea longirostris Lewnielsenius muelleri 44 99 Jarnellius varipalpus Jarnellius sierrensis 1 16 Kompia purpureipes Aztecaedes ramirezi

10 38 Gymnometopa mediovittata Abraedes papago 18 99 Haemagogus (Haemagogus) splendens 99 Haemagogus (Haemagogus) equinus 99 Haemagogus (Conpostegus) leucotaeniatus Haemagogus (Conpostegus) leucocelaenus

99 Neomelaniconion palpale Neomelaniconion lineatopenne

3 95 Aedes esoensis Aedes cinereus

71 99 Paraedes ostentatio Paraedes barraudi

6 99 Verrallina (Harbachius) yusafi Verrallina (Harbachius) nobukonis 99 19 Verrallina (Verrallina) carmenti 12 Verrallina (Verrallina) butleri 93 Verrallina (Neomacleaya) pseudomediofasciata Verrallina (Neomacleaya) indica Aedimorphus punctifemoris Aedimorphus caecus 1 12 Aedimorphus trimaculatus 2 2 Aedimorphus orbitae Aedimorphus mediolineatus 84 24 Aedimorphus pallidostriatus 32 Aedimorphus ochraceus

93 Aedimorphus taeniorhynchoides Aedimorphus pipersalatus 9 96 Aedimorphus eritreae Aedimorphus dalzieli 21 Aedimorphus culicinus 1 47 Aedimorphus pampangensis Aedimorphus alboscutellatus

94 Dobrotworskyius tubbutiensis Dobrotworskyius alboannulatus

99 Patmarksia papuensis Patmarksia argyronotum 16 Georgecraigius (Horsfallius) fluviatilis 81 8 100 Georgecraigius (Georgecraigius) epactius Georgecraigius (Georgecraigius) atropalpus

98 Tanakius togoi 0 Tanakius savoryi 98 Hulecoeteomyia sherki 37 Hulecoeteomyia chrysolineata Gilesius pulchriventer

5 100 Collessius (Collessius) macfarlanei 66 Collessius (Collessius) elsiae 100 Collessius (Alloemyia) pseudotaeniatus Collessius (Alloemyia) banksi Cornetius cozi

43 Stegomyia (Subg. Unassigned) wadai Stegomyia (Subg. Unassigned) chemulpoensis

13 Stegomyia (Subg. Unassigned) poweri Stegomyia (Stegomyia) aegypti 27 20 Stegomyia (Subg. Unassigned) metallica Stegomyia (Subg. Unassigned) apicoargentea Stegomyia (Mukwaya) bromeliae 92 7 59 Stegomyia (Mukwaya) woodi Stegomyia (Mukwaya) simpsoni

32 Stegomyia (Subg. Unassigned) luteocephala 13 1 Stegomyia (Subg. Unassigned) africana 45 Stegomyia (Subg. Unassigned) dendrophila Stegomyia (Subg. Unassigned) deboeri Stegomyia (Subg. Unassigned) riversi

8 Stegomyia (Subg. Unassigned) scutellaris Stegomyia (Actinothrix) edwardsi Stegomyia (Zoromorphus) futunae 11 27 100 Stegomyia (Bohartius) saipanensis Stegomyia (Bohartius) pandani Stegomyia (Subg. Unassigned) albopicta 2 Stegomyia (Subg Unassigned) unilineata 50 Stegomyia (Subg. Unassigned) w-albus 54 Stegomyia (Subg. Unassigned) gardnerii 13 Stegomyia (Xyele) desmotes

65 100 St. (Huagmyia) perplexa 43 St. (Hua.) mediopunctata 81 St. (Heteraspidion) craggi St. (Het.) annandalei Ochlerotatus (Subg. Unassigned) theobaldi Ochlerotatus (Subg. Unassigned) calumnior Ochlerotatus (Subg. Unassigned) andersoni Ochlerotatus (Subg. Unassigned) albifasciatus

24 Ochlerotatus (Subg. Unassigned) sticticus Ochlerotatus (Subg. Unassigned) nivalis

33 Ochlerotatus (Subg. Unassigned) spencerii spencerii Ochlerotatus (Juppius) caballus

10 Ochlerotatus (Subg. Unassigned) ratcliffei Ochlerotatus (Subg. Unassigned) cantator

17 Ochlerotatus (Subg. Unassigned) pullatus Ochlerotatus (Subg. Unassigned) impiger

87 Ochlerotatus (Woodius) intrudens Ochlerotatus (Woodius) diantaeus

2 Ochlerotatus (Pholeomyia) flavifrons Ochlerotatus (Pholeomyia) calcariae

98 Ochlerotatus (Gilesia) mcdonaldi Ochlerotatus (Gilesia) aculeatus

17 Ochlerotatus (Empihals) vigilax Ochlerotatus (Buvirilia) edgari

25 Ochlerotatus (Culicelsa) taeniorhynchus Ochlerotatus (Culicelsa) mitchellae Ochlerotatus (Culicada) canadensis canadensis 0 Ochlerotatus (Subg. Unassigned) thibaulti 9 69 Ochlerotatus (Subg. Unassigned) aurifer Ochlerotatus (Lepidokeneon) spilotus 43 100 Ochlerotatus (Pseudoskusea) postspiraculosus Ochlerotatus (Pseudoskusea) bancroftianus Ochlerotatus (Subg. Unassigned) pionips 0 52 Ochlerotatus (Subg. Unassigned) punctor Ochlerotatus (Subg. Unassigned) hexodontus Ochlerotatus (Subg. Unassigned) communis 3 99 Ochlerotatus (Rusticoidus) rusticus Ochlerotatus (Rusticoidus) refiki Acartomyia zammitii 7 Ochlerotatus (Subg. Unassigned) caspius 16 17 Ochlerotatus (Subg. Unassigned) dorsalis Ochlerotatus (Subg. Unassigned) campestris

86 Ochlerotatus (Ochlerotatus) scapularis Ochlerotatus (Ochlerotatus) infirmatus 8 99 Ochlerotatus (Protoculex) serratus 7 Ochlerotatus (Protoculex) pertinax 100 Ochlerotatus (Chrysoconops) fulvus pallens Ochlerotatus (Chrysoconops) fulvus Ochlerotatus (Subg. Unassigned) riparius Ochlerotatus (Subg. Unassigned) flavescens

17 Ochlerotatus (Subg. Unassigned) stimulans 20 Ochlerotatus (Subg. Unassigned) squamiger

6 Ochlerotatus (Subg. Unassigned) grossbecki Ochlerotatus (Subg. Unassigned) cantans

17 Ochlerotatus (Subg. Unassigned) fitchii Ochlerotatus (Subg. Unassigned) excrucians Culiseta (Culiseta) inornata Culex (Culex) quinquefasciatus Orthopodomyia signifera Mansonia (Mansonia) titillans

45 96 Psorophora (Grabhamia) jamaicensis Psorophora (Grabhamia) columbiae 100 100 Psorophora (Psorophora) howardii 99 Psorophora (Psorophora) ciliata 99 Psorophora (Janthinosoma) ferox Psorophora (Janthinosoma) cyanescens

100 100 Mucidus (Pardomyia) quadripunctis 99 Mucidus (Pardomyia) aurantius aurantius 100 Mucidus (Mucidus) laniger Mucidus (Mucidus) alternans Ochlerotatus (Subg. Unassigned) theobaldi Ochlerotatus (Subg. Unassigned) pionips Ochlerotatus (Culicada) canadensis Ochlerotatus (Subg. Unassigned) calumnior Ochlerotatus (Subg. Unassigned) andersoni Fredwarsius vittatus 58 1 Skusea pembaensis Indusius pulverulentus

100 Scutomyia arboricola Scutomyia albolineata

72 Ochlerotatus (Subg. Unassigned) thibaulti Ochlerotatus (Subg. Unassigned) aurifer

25 Ochlerotatus (Subg. Unassigned) sticticus Ochlerotatus (Subg. Unassigned) nivalis

9 Ochlerotatus (Subg. Unassigned) ratcliffei Ochlerotatus (Subg. Unassigned) cantator

18 Ochlerotatus (Subg. Unassigned) pullatus Ochlerotatus (Subg. Unassigned) impiger

2 Ochlerotatus (Pholeomyia) flavifrons Ochlerotatus (Pholeomyia) calcariae

98 Ochlerotatus (Gilesia) mcdonaldi Ochlerotatus (Gilesia) aculeatus

13 Ochlerotatus (Empihals) vigilax Ochlerotatus (Buvirilia) edgari 74 18 Ochlerotatus (Culicelsa) taeniorhynchus Ochlerotatus (Culicelsa) mitchellae

100 Lorrainea dasyorrhus Lorrainea amesii

16 Isoaedes cavaticus Borichinda cavernicola

97 Howardina walkeri Howardina sexlineata

97 Dahliana geniculata Dahliana echinus Halaedes australis 83 44 Opifex (Opifex) fuscus Opifex (Nothoskusea) chathamicus ‘Ochlerotatus’ (‘Protomacleaya’) brelandi sensu auctorum 1 19 ‘Ochlerotatus’ (’Protomacleaya’) zoosophus sensu auctorum ‘Ochlerotatus’ (’Protomacleaya’) triseriatus sensu auctorum ‘Ochlerotatus’ (’Finlaya’) biocellatus sensu auctorum 4 47 ‘Ochlerotatus’ (’Finlaya’) keefei ‘Ochlerotatus’ (’Finlaya’) candidoscutellum Ochlerotatus (Lepidokeneon) spilotus 13 100 Ochlerotatus (Pseudoskusea) postspiraculosus Ochlerotatus (Pseudoskusea) bancroftianus Ochlerotatus (Subg. Unassigned) albifasciatus 2 25 Ochlerotatus (Subg. Unassigned) spencerii Ochlerotatus (Juppius) caballus Ochlerotatus (Subg. Unassigned) communis 3 100 Ochlerotatus (Rusticoidus) rusticus Ochlerotatus (Rusticoidus) refiki Luius fengi 30 99 Macleya (Macleaya) tremula Macleaya (Chaetocruiomyia) wattensis Lewnielsenius muelleri 33 Jarnellius varipalpus 28 100 Jarnellius sierrensis Dendroskusea periskelata 19 81 Dendroskusea reginae Dendroskusea micropterus Aedimorphus vexans 56 19 Aedimorphus dentatus Aedimorphus cumminsii Catageiomyia irritans 25 31 Catageiomyia tarsalis Catageiomyia argenteopunctata

100 Tewarius reubenae 5 Tewarius agastyai 100 Christophersiomyia thomsoni Christophersiomyia gombakensis ‘Ochlerotatus’ (’Protomacleaya’) knabi sensu auctorum 0 Molpemyia pecuniosa 16 100 Rampamyia notoscripta Rampamyia albilabris ‘Ochlerotatus’ (’Finlaya’) oreophilus sensu auctorum 26 Jihliensius chungi 44 97 Phagomyia lophoventralis Phagomyia gubernatoris

53 Ochlerotatus (Subg. Unassigned) punctor 0 Ochlerotatus (Subg. Unassigned) hexodontus 87 Ochlerotatus (Woodius) intrudens Ochlerotatus (Woodius) diantaeus Acartomyia zammitii 6 Ochlerotatus (Subg. Unassigned) caspius 15 16 Ochlerotatus (Subg. Unassigned) dorsalis Ochlerotatus (Subg. Unassigned) campestris

100 Diceromyia taylori 4 Diceromyia furcifer 100 Ayurakitia peytoni Ayurakitia griffithi

99 Albuginosus ngong 1 Albuginosus marshallii 97 Elpeytonius simulans Elpeytonius apicoannulatus Bifidistylus lamborni 4 Polyleptiomyia albocephala 15 94 Aedimorphus quasiunivittatus Aedimorphus gibbinsi ‘Ochlerotatus’ (’Protomacleaya’) burgeri sensu auctorum 2 ‘Ochlerotatus’ (’Protomacleaya’) kompi sensu auctorum 19 ‘Ochlerotatus’ (’Pro.’) homoeopus sensu auctorum 83 20 ‘Ochlerotatus’ (’Pro.’) terrens sensu auctorum ‘Ochlerotatus’ (’Pro.’) galindoi sensu auctorum ‘Ochlerotatus’ (‘Finlaya’) crossi sensu auctorum

28 93 Hopkinsius (Yamada) seoulensis 65 Hopkinsius (Yamada) albocinctus 36 Hopkinsius (Hopkinsius) ingrami Hopkinsius (Hopkinsius) embuensis

83 Ochlerotatus (Ochlerotatus) scapularis Ochlerotatus (Ochlerotatus) infirmatus 10 99 Ochlerotatus (Protoculex) serratus 5 Ochlerotatus (Protoculex) pertinax 100 Ochlerotatus (Chrysoconops) fulvus pallens Ochlerotatus (Chrysoconops) fulvus

100 Downsiomyia nivea 2 Downsiomyia leonis 2 100 Finlaya poicilia 58 Finlaya kochi 70 Danielsia harperi Danielsia albotaeniata Catatassomyia meronephada

99 Udaya lucaris Udaya argyrurus 6 100 Bothaella helenae Bothaella eldridgei Belkinius aurotaeniatus 6 100 Zeugnomyia lawtoni Zeugnomyia gracilis Ochlerotatus (Subg. Unassigned) flavescens Ochlerotatus (Subg. Unassigned) fitchii

18 Ochlerotatus (Subg. Unassigned) stimulans 17 Ochlerotatus (Subg. Unassigned) squamiger

1 Ochlerotatus (Subg. Unassigned) riparius Ochlerotatus (Subg. Unassigned) excrucians

1 Ochlerotatus (Subg. Unassigned) grossbecki Ochlerotatus (Subg. Unassigned) cantans

22 Kompia purpureipes Aztecaedes ramirezi

10 44 Gymnometopa mediovittata Abraedes papago 18 99 Haemagogus (Haemagogus) splendens 99 Haemagogus (Haemagogus) equinus 99 Haemagogus (Conpostegus) leucotaeniatus Haemagogus (Conpostegus) leucocelaenus Himalaius gilli 72 100 Bruceharrisonius greenii Bruceharrisonius alektorovi 13 95 Vansomerensus pulchrithorax Vansomerensus luteostriata 40 99 Zavortinkius longipalpis 7 Zavortinkius fulgens 99 Kenknightia harbachi Kenknightia dissimilis Cancraedes masculinus 100 21 Cancraedes penghuensis Cancraedes cancricomes 58 96 Geoskusea longiforceps Geoskusea baisasi 29 Sallumia hortator 8 Levua geoskusea

100 Rhinoskusea wardi Rhinoskusea longirostris

51 Leptosomatomyia aurimargo Huaedes wauensis Alanstonea brevitibia 15 46 100 Eretmapodites quinquevittatus Eretmapodites chrysogaster 66 99 Armigeres (Leicesteria) longipalpis 83 Armigeres (Leicesteria) flavus 89 Armigeres (Armigeres) subalbatus Armigeres (Armigeres) breinli Petermattinglyius whartoni 89 Petermattinglyius franciscoi 44 99 Petermattinglyius scanloni 3 Petermattinglyius iyengari 85 Pseudarmigeres michaelikati Pseudarmigeres argenteoventralis dunni 29 91 Heizmannia (Mattinglyia) catesi 99 Heizmannia (Mattinglyia) achaetae 100 Heizmannia (Heizmannia) scintillans Heizmannia (Heizmannia) complex

99 Edwardsaedes imprimens Edwardsaedes bekkui

4 99 Neomelaniconion palpale Neomelaniconion lineatopenne

6 95 Aedes esoensis Aedes cinereus

79 99 Paraedes ostentatio Paraedes barraudi

10 98 Verrallina (Harbachius) yusafi Verrallina (Harbachius) nobukonis 98 19 Verrallina (Verrallina) carmenti 15 Verrallina (Verrallina) butleri 93 Verrallina (Neomacleaya) pseudomediofasciata Verrallina (Neomacleaya) indica Aedimorphus domesticus Aedimorphus punctifemoris 2 Aedimorphus caecus

8 Aedimorphus trimaculatus 1 5 Aedimorphus orbitae Aedimorphus mediolineatus 83 26 Aedimorphus pallidostriatus 31 Aedimorphus ochraceus

90 Aedimorphus ochraceus Aedimorphus pipersalatus 7 96 Aedimorphus eritreae Aedimorphus dalzieli 19 Aedimorphus culicinus 1 45 Aedimorphus pampangensis Aedimorphus alboscutellatus

95 Dobrotworskyius tubbutiensis Dobrotworskyius alboannulatus

99 Patmarksia papuensis Patmarksia argyronotum 14 26 Georgecraigius (Horsfallius) fluviatilis 78 100 Georgecraigius (Georgecraigius) epactius Georgecraigius (Georgecraigius) atropalpus 1 Gilesius pulchriventer

99 Tanakius togoi 5 Tanakius savoryi 44 98 Hulecoeteomyia sherki Hulecoeteomyia chrysolineata

99 Collessius (Collessius) macfarlanei 67 Collessius (Collessius) elsiae 100 Collessius (Alloemyia) pseudotaeniatus Collessius (Alloemyia) banksi Cornetius cozi

41 Stegomyia (Subg. Unassigned) wadai Stegomyia (Subg. Unassigned) chemulpoensis Stegomyia (Subg. Unassigned) poweri 25 14 Stegomyia (Stegomyia) aegypti 20 Stegomyia (Subg. Unassigned) metallica 1 Stegomyia (Subg. Unassigned) apicoargentea Stegomyia (Mukwaya) bromeliae 90 9 58 Stegomyia (Mukwaya) woodi Stegomyia (Mukwaya) simpsoni

14 32 Stegomyia (Subg. Unassigned) luteocephala 1 Stegomyia (Subg. Unassigned) africana 42 Stegomyia (Subg. Unassigned) dendrophila Stegomyia (Subg. Unassigned) deboeri Stegomyia (Subg. Unassigned) riversi

9 Stegomyia (Subg. Unassigned) scutellaris Stegomyia (Actinothrix) edwardsi Stegomyia (Zoromorphus) futunae 5 20 100 Stegomyia (Bohartius) saipanensis Stegomyia (Bohartius) pandani Stegomyia (Subg. Unassigned) albopicta 5 Stegomyia (Subg. Unassigned) unilineata 50 Stegomyia (Subg. Unassigned) w-albus 54 Stegomyia (Subg. Unassigned) gardnerii 17 Stegomyia (Xyele) desmotes

65 100 St. (Huagmyia) perplexa 45 St. (Hua.) mediopunctata 82 St. (Heteraspidion) craggi St. (Het.) annandalei Aedini, 14 characters ordered, strict consensus of 4000 MPTs, L=6654 Strict consensus tree Culiseta (Culiseta) inornata Culex (Culex) quinquefasciatus Orthopodomyia signifera Mansonia (Mansonia) titillans Psorophora (Grabhamia) columbiae Ps. (Gra.) jamaicensis Ps. (Janthinosma) cyanescens Ps.(Jan.) ferox Ps. (Psorophora) ciliata Ps. (Pso.) howardii Polyleptiomyia albocephala Catageiomyia irritans Bifidistylus lamborni Aedimorphus punctifemoris Aztecaedes ramirezi Kompia purpureipes Molpemyia pecuniosa 'Oc.' ('Finlaya') biocellatus ‘Oc.’ (’Fin.’) oreophilus ‘Oc.’ (’Protomacleaya’) brelandi ‘Oc.’ (’Pro.’) burgeri ‘Oc.’ (’Pro.’) knabi ‘Oc.’ (’Pro.’) kompi ‘Oc.’ (’Pro.’) triseriatus ‘Oc.’ (’Pro.’) zoosophus Abraedes papago Gymnometopa mediovittata Elpeytonius apicoannulatus Elpeytonius simulans Catageiomyia argenteopunctatus Catageiomyia tarsalis Albuginosus marshallii Albuginosus ngong Dahliana echinus Dahliana geniculata Downsiomyia leonis Downsiomyia nivea Howardina sexlineata Howardina walkeri Kenknightia dissimilis Kenknightia harbachi ‘Oc.’ (‘Fin.’) candidoscutellum ‘Oc.’ (‘Fin.’) keefei Rampamyia albilabris Rampamyia notoscripta Vansomerensus luteostriata Vansomerensis pulchrithorax Zavortinkius fulgens Zavortinkius longipalpis Himalaius gilli Bruceharrisonius alektorovi Bruceharrisonius greenii Fredwarsius vittatus Indusius pulverulentus Skusea pembaensis Lewneilsenius muelleri Jarnellius sierrensis Jarnellius varipalpus Jihliensius chungi Phagomyia gubernatoris Phagomyia lophoventralis Luius fengi Macleaya (Chaetocuiomyia) wattensis Mc.(Macleaya) tremlula ‘Oc.’ (’Pro.’) galindoi Oc.Pro.terrens Dn.albotaeniata Dn.harperi Fl.kochi Fl.poicilia Oc.Aca.zammitii Ha.australis Op.Not.chathamicus Op.Opi.fuscus Hg.Con.leucocelaenus Hg.Con.leucotaeniatus Hg.Hag.equinus Hg.Hag.splendens Mu.Muc.alternans Mu.Muc.laniger Mu.Pdo.aurantius.aurantius Mu.Pdo.quadripunctis Oc.Fin.crossi Hk.Hpk.embuensis Hk.Hpk.ingrami Hk.Yam.albocinctus Hk.Yam.seoulensis Lo.amesii Lo.dasyorrhus Ae.Can.masculinus Ae.Can.cancricomes Ae.Can.penghuensis Oc.Lev.geoskusea Oc.Sal.hortator Oc.Geo.baisasi Oc.Geo.longiforceps Oc.Rhi.longirostris Oc.Rhi.wardi Cr.gombakensis Cr.thomsoni Te.agastyai Te.reubenae Hu.wauensis Lp.aurimargo As.brevitibia Er.chrysogaster Er.quinquevittatus Ar.Arm.breinli Ar.Arm.subalbatus Ar.Lei.flavus Ar.Lei.longipalpis Db.alboannulatus Db.tubbutiensis Pm.argyronotum Pm.papuensis Gc.Hor.fluviatilis Gc.Gec.atropalpus Gc.Gec.epactius Gi.pulchriventer Hl.chrysolineata Hl.sherki Ta.savoryi Ta.togoi Co.Alo.banksi Co.Alo.pseudotaeniatus Co.Col.elsiae Co.Col.macfarlanei Ae.Adm.gibbinsi Ae.Adm.quasiunivittatus Ae.Adm.vexans.vexans Ae.Adm.cumminsii Ae.Adm.dentatus Ed.bekkui Ed.imprimens Ne.lineatopenne Ne.palpale Ae.cinereus Ae.esoensis Pr.barraudi Pr.ostentatio Ve.Har.nobukonis Ve.Har.yusafi Ve.Ver.butleri Ve.Ver.carmenti Ve.Nma.indica Ve.Nma.pseudomediofasciata Ae.Adm.domesticus Ae.Adm.dalzieli Ae.Adm.eritreae Ae.Adm.culicinus Ae.Adm.alboscutellatus Ae.Adm.pampangensis Ae.Adm.pipersalatus Ae.Adm.taeniorhynchoides Ae.Adm.caecus Ae.Adm.orbitae Ae.Adm.trimaculatus Ae.Adm.mediolineatus Ae.Adm.ochraceus Ae.Adm.pallidostriatus Oc.Phl.calcariae Oc.Phl.flavifrons Oc.theobaldi Oc.Cul.mitchellae Oc.Cul.taeniorhynchus Oc.spilotus Oc.Psk.bancroftianus Oc.Psk.postspiraculosus Oc.canadensis.canadensis Oc.calumnior Oc.Och.infirmatus Oc.Och.scapularis Oc.Pcx.pertinax Oc.Pcx.serratus Oc.Gli.aculeatus Oc.Gli.mcdonaldi Oc.Emp.vigilax Oc.Buv.edgari Oc.Chs.fulvusfulvus Oc.Chs.fulvuspallens Oc.andersoni Oc.cantator Oc.ratcliffei Oc.aurifer Oc.thibaulti Oc.diantaeus Oc.intrudens Oc.impiger Oc.pullatus Oc.nivalis Oc.sticticus Oc.Excrucians Oc.fitchii Oc.grossbecki Oc.flavescens Oc.squamiger Oc.cantans Oc.riparius Oc.stimulans Oc.pionips Oc.hexodontus Oc.punctor Oc.communis Oc.Rus.refiki Oc.Rus.rusticus Oc.albifasciatus Oc.caballus Oc.spencerii.spencerii Oc.campestris Oc.caspius Oc.dorsalis Ay.griffithi Ay.peytoni Bc.cavernicola Ia.cavaticus Di.furcifer Di.taylori Sc.albolineata Sc.arboricola Di.Periskelata Di.micropterus Di.reginae Be.aurotaeniatus Di.meronephada Bo.eldridgei Bo.helenae Ud.argyrurus Ud.lucaris Ze.gracilis Ze.Lawtoni Di.whartoni Di.franciscoi Di.iyengari Di.scanloni Pa.argenteoventralis.dunni Pa.michaelikati Hz.Hez.complex Hz.Hez.scintillans Hz.Mat.achaetae Hz.Mat.catesi Ae.Cor.cozi St.africana St.apicoargentea St.luteocephala St.metallica St.aegypti St.poweri St.deboeri St.dendrophila St.bromeliae St.simpsoni St.woodi St.chemulpoensis St.wadai St.albopicta St.unilineata St.riversi St.scutellaris St.edwardsi St.futunae St.pandani St.saipanensis St.walbus St.gardnerii.gardnerii St.desmotes St.annandalei St.craggi St.mediopunctata St.perplexa Strict consensus tree Cs.Cus.inornata Cx.Cux.quinquefasciatus Or.signifera Ma.Man.titillans Ps.Gra.columbiae Ps.Gra.jamaicensis Ps.Jan.cyanescens Ps.Jan.ferox Ps.Pso.ciliata Ps.Pso.howardii Ae.Adm.albocephalus Ae.Adm.irritans Ae.Adm.lamborni Ae.Adm.punctifemoris Az.ramirezi Ko.purpureipes Mo.pecuniosa Oc.Fin.biocellatus Oc.Fin.oreophilus Oc.Pro.brelandi Oc.Pro.burgeri Oc.Pro.knabi Oc.Pro.kompi Oc.Pro.triseriatus Oc.Pro.zoosophus Ab.papago Gy.mediovittata Ae.Adm.apicoannulatus Ae.Adm.simulans Ae.Adm.argenteopunctatus Ae.Adm.tarsalis Al.marshallii Al.ngong Da.echinus Da.geniculata Do.leonis Do.nivea Hw.sexlineata Hw.walkeri Ke.dissimilis Ke.harbachi Oc.Fin.candidoscutellum Oc.Fin.keefei Ra.albilabris Ra.notoscripta Va.luteostriata Va.pulchrithorax Za.fulgens Za.longipalpis Hi.gilli Br.alektorovi Br.greenii Fr.vittatus In.pulverulentus Sk.pembaensis Ja.Lew.muelleri Ja.Jar.sierrensis Ja.Jar.varipalpus Ji.chungi Ph.gubernatoris Ph.lophoventralis Lu.fengi Mc.Cha.wattensis Mc.Mac.tremula ‘Oc.’ (’Pro.’) galindoi ‘Oc.’ (’Pro.’) homoeopus ‘Oc.’ (’Pro.’) terrens Danielsia albotaeniata Danielsia harperi Finlaya kochi Finlaya poicilia Acartomyia zammitii Halaedes australis Opifex (Nothoskusea) chathamicus Opifex (Opifex) fuscus Haemagogus (Conopostegus) leucocelaenus Haemagogus (Con.) leucotaeniatus Haemagogus (Haemagogus) equinus Haemagogus (Hag.) splendens Mucidus (Mucidus) alternans Mucidus (Muc.) laniger Mucidus (Pardomyia) aurantius Mucidus (Pdo.) quadripunctis ‘Oc.’ (’Finlaya’) crossi Hopkinsius (Hopkinsius) embuensis Hopkinsius (Hpk.) ingrami Hopkinsius (Yamada) albocinctus Hopkinsius (Yam.) seoulensis Lorrainea amesii Lorrainea dasyorrhus Cancraedes masculinus Cancraedes cancricomes Cancraedes penghuensis Levua geoskusea Sallumia hortator Geoskusea baisasi Geoskusea longiforceps Rhinoskusea longirostris Rhinoskusea wardi Christophersiomyia gombakensis Christophersiomyia thomsoni Tewarius agastyai Tewarius reubenae Huaedes wauensis Leptosomatomyia aurimargo Alanstonea brevitibia Eretmapodites chrysogaster Eretmapodites quinquevittatus Armigeres (Armigeres) breinli Armigeres (Arm.) subalbatus Armigeres (Leicesteria) flavus Armigeres (Lei.) longipalpis Dobrotworskyius alboannulatus Dobrotworskyius tubbutiensis Patmarksia argyronotum Patmarksia papuensis Georgecraigius (Horsfallius) fluviatilis Gc.(Georgecraigius) atropalpus Gc.(Georgecraigius) epactius Gilesius pulchriventer Hulecoeteomyia chrysolineata Hulecoeteomyia sherki Tanakius savoryi Tanakius togoi Collessius (Alloemyia) banksi Collessius (Alo.) pseudotaeniatus Collessius (Collessius) elsiae Collessius (Col.) macfarlanei Aedimorphus gibbinsi Aedimorphus quasiunivittatus Aedimorphus vexans vexans Aedimorphus cumminsii Aedimorphus dentatus Edwardsaedes bekkui Ed.imprimens Ne.lineatopenne Ne.palpale Ae.cinereus Ae.esoensis Pr.barraudi Pr.ostentatio Ve.Har.nobukonis Ve.Har.yusafi Ve.Ver.butleri Ve.Ver.carmenti Ve.Nma.indica Ve.Nma.pseudomediofasciata Ae.Adm.domesticus Ae.Adm.dalzieli Ae.Adm.eritreae Ae.Adm.culicinus Ae.Adm.alboscutellatus Ae.Adm.pampangensis Ae.Adm.pipersalatus Ae.Adm.taeniorhynchoides Ae.Adm.caecus Ae.Adm.orbitae Ae.Adm.trimaculatus Ae.Adm.mediolineatus Ae.Adm.ochraceus Ae.Adm.pallidostriatus Oc.Phl.calcariae Oc.Phl.flavifrons Oc.theobaldi Oc.Cul.mitchellae Oc.Cul.taeniorhynchus Oc.spilotus Oc.Psk.bancroftianus Oc.Psk.postspiraculosus Oc.canadensis.canadensis Oc.calumnior Oc.Och.infirmatus Oc.Och.scapularis Oc.Pcx.pertinax Oc.Pcx.serratus Oc.Gli.aculeatus Oc.Gli.mcdonaldi Oc.Emp.vigilax Oc.Buv.edgari Oc.Chs.fulvusfulvus Oc.Chs.fulvuspallens Oc.andersoni Oc.cantator Oc.ratcliffei Oc.aurifer Oc.thibaulti Oc.diantaeus Oc.intrudens Oc.impiger Oc.pullatus Oc.nivalis Oc.sticticus Oc.Excrucians Oc.fitchii Oc.grossbecki Oc.flavescens Oc.squamiger Oc.cantans Oc.riparius Oc.stimulans Oc.pionips Oc.hexodontus Oc.punctor Oc.communis Oc.Rus.refiki Oc.Rus.rusticus Oc.albifasciatus Oc.caballus Oc.spencerii.spencerii Oc.campestris Oc.caspius Oc.dorsalis Ay.griffithi Ay.peytoni Bc.cavernicola Ia.cavaticus Di.furcifer Di.taylori Sc.albolineata Sc.arboricola Di.Periskelata Di.micropterus Di.reginae Be.aurotaeniatus Di.meronephada Bo.eldridgei Bo.helenae Ud.argyrurus Ud.lucaris Ze.gracilis Ze.Lawtoni Di.whartoni Di.franciscoi Di.iyengari Di.scanloni Pa.argenteoventralis.dunni Pa.michaelikati Hz.Hez.complex Hz.Hez.scintillans Hz.Mat.achaetae Hz.Mat.catesi Ae.Cor.cozi St.africana St.apicoargentea St.luteocephala St.metallica St.aegypti St.poweri St.deboeri St.dendrophila St.bromeliae St.simpsoni St.woodi St.chemulpoensis St.wadai St.albopicta St.unilineata St.riversi St.scutellaris St.edwardsi St.futunae St.pandani St.saipanensis St.walbus St.gardnerii.gardnerii St.desmotes St.annandalei St.craggi St.mediopunctata St.perplexa Strict consensus tree Cs.Cus.inornata Cx.Cux.quinquefasciatus Or.signifera Ma.Man.titillans Ps.Gra.columbiae Ps.Gra.jamaicensis Ps.Jan.cyanescens Ps.Jan.ferox Ps.Pso.ciliata Ps.Pso.howardii Ae.Adm.albocephalus Ae.Adm.irritans Ae.Adm.lamborni Ae.Adm.punctifemoris Az.ramirezi Ko.purpureipes Mo.pecuniosa Oc.Fin.biocellatus Oc.Fin.oreophilus Oc.Pro.brelandi Oc.Pro.burgeri Oc.Pro.knabi Oc.Pro.kompi Oc.Pro.triseriatus Oc.Pro.zoosophus Ab.papago Gy.mediovittata Ae.Adm.apicoannulatus Ae.Adm.simulans Ae.Adm.argenteopunctatus Ae.Adm.tarsalis Al.marshallii Al.ngong Da.echinus Da.geniculata Do.leonis Do.nivea Hw.sexlineata Hw.walkeri Ke.dissimilis Ke.harbachi Oc.Fin.candidoscutellum Oc.Fin.keefei Ra.albilabris Ra.notoscripta Va.luteostriata Va.pulchrithorax Za.fulgens Za.longipalpis Hi.gilli Br.alektorovi Br.greenii Fr.vittatus In.pulverulentus Sk.pembaensis Ja.Lew.muelleri Ja.Jar.sierrensis Ja.Jar.varipalpus Ji.chungi Ph.gubernatoris Ph.lophoventralis Lu.fengi Mc.Cha.wattensis Mc.Mac.tremula Oc.Pro.galindoi Oc.Pro.homoeopus Oc.Pro.terrens Dn.albotaeniata Dn.harperi Fl.kochi Fl.poicilia Oc.Aca.zammitii Ha.australis Op.Not.chathamicus Op.Opi.fuscus Hg.Con.leucocelaenus Hg.Con.leucotaeniatus Hg.Hag.equinus Hg.Hag.splendens Mu.Muc.alternans Mu.Muc.laniger Mu.Pdo.aurantius.aurantius Mu.Pdo.quadripunctis Oc.Fin.crossi Hk.Hpk.embuensis Hk.Hpk.ingrami Hk.Yam.albocinctus Hk.Yam.seoulensis Lo.amesii Lo.dasyorrhus Ae.Can.masculinus Ae.Can.cancricomes Ae.Can.penghuensis Oc.Lev.geoskusea Oc.Sal.hortator Oc.Geo.baisasi Oc.Geo.longiforceps Oc.Rhi.longirostris Oc.Rhi.wardi Cr.gombakensis Cr.thomsoni Te.agastyai Te.reubenae Hu.wauensis Lp.aurimargo As.brevitibia Er.chrysogaster Er.quinquevittatus Ar.Arm.breinli Ar.Arm.subalbatus Ar.Lei.flavus Ar.Lei.longipalpis Db.alboannulatus Db.tubbutiensis Pm.argyronotum Pm.papuensis Gc.Hor.fluviatilis Gc.Gec.atropalpus Gc.Gec.epactius Gi.pulchriventer Hl.chrysolineata Hl.sherki Ta.savoryi Ta.togoi Co.Alo.banksi Co.Alo.pseudotaeniatus Co.Col.elsiae Co.Col.macfarlanei Ae.Adm.gibbinsi Ae.Adm.quasiunivittatus Ae.Adm.vexans.vexans Ae.Adm.cumminsii Aedimorphus dentatus Edwardsaedes bekkui Edwardsaedes imprimens Neomelaniconion lineatopenne Neomelaniconion palpale Aedes cinereus Aedes esoensis Paraedes barraudi Paraedes ostentatio Verrallina (Harbachius) nobukonis Verrallina (Har.) yusafi Verrallina (Verrrallina) butleri Verrallina (Ver.) carmenti Verrallina (Neomacleaya) indica Verrallina (Nma.) pseudomediofasciata Aedimorphus domesticus Aedimorphus dalzieli Aedimorphus eritreae Aedimorphus culicinus Aedimorphus alboscutellatus Aedimorphus pampangensis Aedimorphus pipersalatus Aedimorphus taeniorhynchoides Aedimorphus caecus Aedimorphus orbitae Aedimorphus trimaculatus Aedimorphus mediolineatus Aedimorphus ochraceus Aedimorphus pallidostriatus Ochlerotatus (Pholeomyia) calcariae Oc. (Phl.) flavifrons Oc. (Sub. Un.) theobaldi Oc. (Culicelsa) mitchellae Oc. (Cul.) taeniorhynchus Oc. (Leptokeneon) spilotus Oc. (Pseudoskusea) bancroftianus Oc. (Psk.) postspiraculosus Oc. (Culicada) canadensis Oc. (Sub. Un.) calumnior Oc. (Ochlerotatus) infirmatus Oc. (Och.) scapularis Oc. (Protoculex) pertinax Oc. (Pcx.) serratus Oc. (Gilesia) aculeatus Oc. (Gli.) mcdonaldi Oc. (Empihals) vigilax Oc. (Buvirilia) edgari Oc. (Chysoconops) fulvus Oc. (Chs.) fulvus pallens Oc. (Sub. Un.) andersoni Oc. (Sub. Un.) cantator Oc. (Sub. Un.) ratcliffei Oc. (Sub. Un.) aurifer Oc. (Sub. Un.) thibaulti Oc. (Woodius) diantaeus Oc. (Woo.) intrudens Oc. (Sub. Un.) impiger Oc. (Sub. Un.) pullatus Oc. (Sub. Un.) nivalis Oc. (Sub. Un.) sticticus Oc. (Sub. Un.) excrucians Oc. (Sub. Un.) fitchii Oc. (Sub. Un.) grossbecki Oc. (Sub. Un.) flavescens Oc. (Sub. Un.) squamiger Oc. (Sub. Un.) cantans Oc. (Sub. Un.) riparius Oc. (Sub. Un.) stimulans Oc. (Sub. Un.) pionips Oc. (Sub. Un.) hexodontus Oc. (Sub. Un.) punctor Oc.communis Oc.Rus.refiki Oc.Rus.rusticus Oc.albifasciatus Oc.caballus Oc.spencerii.spencerii Oc.campestris Oc.caspius Oc.dorsalis Ay.griffithi Ay.peytoni Bc.cavernicola Ia.cavaticus Di.furcifer Di.taylori Sc.albolineata Sc.arboricola Di.Periskelata Di.micropterus Di.reginae Be.aurotaeniatus Di.meronephada Bo.eldridgei Bo.helenae Ud.argyrurus Ud.lucaris Ze.gracilis Ze.Lawtoni Di.whartoni Di.franciscoi Di.iyengari Di.scanloni Pa.argenteoventralis.dunni Pa.michaelikati Hz.Hez.complex Hz.Hez.scintillans Hz.Mat.achaetae Hz.Mat.catesi Ae.Cor.cozi St.africana St.apicoargentea St.luteocephala St.metallica St.aegypti St.poweri St.deboeri St.dendrophila St.bromeliae St.simpsoni St.woodi St.chemulpoensis St.wadai St.albopicta St.unilineata St.riversi St.scutellaris St.edwardsi St.futunae St.pandani St.saipanensis St.walbus St.gardnerii.gardnerii St.desmotes St.annandalei St.craggi St.mediopunctata St.perplexa Strict consensus tree Cs.Cus.inornata Cx.Cux.quinquefasciatus Or.signifera Ma.Man.titillans Ps.Gra.columbiae Ps.Gra.jamaicensis Ps.Jan.cyanescens Ps.Jan.ferox Ps.Pso.ciliata Ps.Pso.howardii Ae.Adm.albocephalus Ae.Adm.irritans Ae.Adm.lamborni Ae.Adm.punctifemoris Az.ramirezi Ko.purpureipes Mo.pecuniosa Oc.Fin.biocellatus Oc.Fin.oreophilus Oc.Pro.brelandi Oc.Pro.burgeri Oc.Pro.knabi Oc.Pro.kompi Oc.Pro.triseriatus Oc.Pro.zoosophus Ab.papago Gy.mediovittata Ae.Adm.apicoannulatus Ae.Adm.simulans Ae.Adm.argenteopunctatus Ae.Adm.tarsalis Al.marshallii Al.ngong Da.echinus Da.geniculata Do.leonis Do.nivea Hw.sexlineata Hw.walkeri Ke.dissimilis Ke.harbachi Oc.Fin.candidoscutellum Oc.Fin.keefei Ra.albilabris Ra.notoscripta Va.luteostriata Va.pulchrithorax Za.fulgens Za.longipalpis Hi.gilli Br.alektorovi Br.greenii Fr.vittatus In.pulverulentus Sk.pembaensis Ja.Lew.muelleri Ja.Jar.sierrensis Ja.Jar.varipalpus Ji.chungi Ph.gubernatoris Ph.lophoventralis Lu.fengi Mc.Cha.wattensis Mc.Mac.tremula Oc.Pro.galindoi Oc.Pro.homoeopus Oc.Pro.terrens Dn.albotaeniata Dn.harperi Fl.kochi Fl.poicilia Oc.Aca.zammitii Ha.australis Op.Not.chathamicus Op.Opi.fuscus Hg.Con.leucocelaenus Hg.Con.leucotaeniatus Hg.Hag.equinus Hg.Hag.splendens Mu.Muc.alternans Mu.Muc.laniger Mu.Pdo.aurantius.aurantius Mu.Pdo.quadripunctis Oc.Fin.crossi Hk.Hpk.embuensis Hk.Hpk.ingrami Hk.Yam.albocinctus Hk.Yam.seoulensis Lo.amesii Lo.dasyorrhus Ae.Can.masculinus Ae.Can.cancricomes Ae.Can.penghuensis Oc.Lev.geoskusea Oc.Sal.hortator Oc.Geo.baisasi Oc.Geo.longiforceps Oc.Rhi.longirostris Oc.Rhi.wardi Cr.gombakensis Cr.thomsoni Te.agastyai Te.reubenae Hu.wauensis Lp.aurimargo As.brevitibia Er.chrysogaster Er.quinquevittatus Ar.Arm.breinli Ar.Arm.subalbatus Ar.Lei.flavus Ar.Lei.longipalpis Db.alboannulatus Db.tubbutiensis Pm.argyronotum Pm.papuensis Gc.Hor.fluviatilis Gc.Gec.atropalpus Gc.Gec.epactius Gi.pulchriventer Hl.chrysolineata Hl.sherki Ta.savoryi Ta.togoi Co.Alo.banksi Co.Alo.pseudotaeniatus Co.Col.elsiae Co.Col.macfarlanei Ae.Adm.gibbinsi Ae.Adm.quasiunivittatus Ae.Adm.vexans.vexans Ae.Adm.cumminsii Ae.Adm.dentatus Ed.bekkui Ed.imprimens Ne.lineatopenne Ne.palpale Ae.cinereus Ae.esoensis Pr.barraudi Pr.ostentatio Ve.Har.nobukonis Ve.Har.yusafi Ve.Ver.butleri Ve.Ver.carmenti Ve.Nma.indica Ve.Nma.pseudomediofasciata Ae.Adm.domesticus Ae.Adm.dalzieli Ae.Adm.eritreae Ae.Adm.culicinus Ae.Adm.alboscutellatus Ae.Adm.pampangensis Ae.Adm.pipersalatus Ae.Adm.taeniorhynchoides Ae.Adm.caecus Ae.Adm.orbitae Ae.Adm.trimaculatus Ae.Adm.mediolineatus Ae.Adm.ochraceus Ae.Adm.pallidostriatus Oc.Phl.calcariae Oc.Phl.flavifrons Oc.theobaldi Oc.Cul.mitchellae Oc.Cul.taeniorhynchus Oc.spilotus Oc.Psk.bancroftianus Oc.Psk.postspiraculosus Oc.canadensis.canadensis Oc.calumnior Oc.Och.infirmatus Oc.Och.scapularis Oc.Pcx.pertinax Oc.Pcx.serratus Oc.Gli.aculeatus Oc.Gli.mcdonaldi Oc.Emp.vigilax Oc.Buv.edgari Oc.Chs.fulvusfulvus Oc.Chs.fulvuspallens Oc.andersoni Oc.cantator Oc.ratcliffei Oc.aurifer Oc.thibaulti Oc.diantaeus Oc.intrudens Oc.impiger Oc.pullatus Oc.nivalis Oc.sticticus Oc.Excrucians Oc.fitchii Oc.grossbecki Oc.flavescens Oc.squamiger Oc.cantans Oc.riparius Oc.stimulans Oc.pionips Oc. (Sub. Un.) hexodontus Oc. (Sub. Un.) punctor Oc. (Sub. Un.) communis Oc. (Rusticoidus) refiki Oc. (Rus.) rusticus Oc. (Sub. Un.) albifasciatus Oc. (Juppius) caballus Oc. (Sub. Un.) spencerii Oc. (Sub. Un.) campestris Oc. (Sub. Un.) caspius Oc. (Sub. Un.) dorsalis Ayurakitia griffithi Ayurakitia peytoni Borichinda cavernicola Isoaedes cavaticus Diceromyia furcifer Diceromyia taylori Scutomyia albolineata Scutomyia arboricola Dendroskusea periskelata Dendroskusea micropterus Dendroskusea reginae Belkinius aurotaeniatus Catatassomyia meronephada Bothaella eldridgei Bothaella helenae Udaya argyrurus Udaya lucaris Zeugnomyia gracilis Zeugnomyia lawtoni Petermattinglyius whartoni Petermattinglyius franciscoi Petermattinglyius iyengari Petermattinglyius scanloni Pseudarmigeres a. dunni Pseudarmigeres michaelikati Heizmannia (Heizmannia) complex Hz. (Hez.) scintillans Hz. (Mattinglyia) achaetae Hz. (Mat.) catesi Cornetius cozi Stegomyia (Sub. Un.) africana Stegomyia (Sub. Un.) apicoargentea Stegomyia (Sub. Un.) luteocephala Stegomyia (Sub. Un.) metallica Stegomyia (Stegomyia) aegypti Stegomyia (Sub. Un.) poweri Stegomyia (Sub. Un.) deboeri Stegomyia (Sub. Un.) dendrophila Stegomyia (Mukwaya) bromeliae Stegomyia (Muk.) simpsoni Stegomyia (Muk.) woodi Stegomyia (Sub. Un.) chemulpoensis Stegomyia (Sub. Un.) wadai Stegomyia (Sub. Un.) albopicta Stegomyia (Sub. Un.) unilineata Stegomyia (Sub. Un.) riversi Stegomyia (Sub. Un.) scutellaris Stegomyia (Actinothrix) edwardsi Stegomyia (Zoromorphus) futunae Stegomyia (Bohartius) pandani Stegomyia (Boh.) saipanensis Stegomyia (Sub. Un.) w-albus Stegomyia (Sub. Un.) gardnerii Stegomyia (Xyele) desmotes Stegomyia (Heteraspidion) annandalei Stegomyia (Het.) craggi St. (Huangmyia) mediopunctata St. (Hua.) perplexa Aedini, all characters unordered, strict consensus of 832 MPTs, L=6609 Strict consensus tree

Culiseta (Culiseta) inornata Culex (Culex) quinquefasciatus Orthopodomyia signifera Mansonia (Mansonia) titillans Psorophora (Grabhamia) columbiae Ps. (Gra.) jamaicensis Ps. (Janthinosma) cyanescens Ps.(Jan.) ferox Ps. (Psorophora) ciliata Ps. (Pso.) howardii Fredswardsius vittatus Isoaedes cavaticus Indusius pulverulentus Skusea pembaensis Halaedes australis Opifex (Nothoskusea) chathamicus Opifex (Opifex) fuscus Bifidistylus lamborni Polyleptiomyia albocephala Aedimorphus gibbinsi Aedimorphus quasiunivittatus Aedimorphus vexans vexans Aedimorphus cumminsii Aedimorphus dentatus Edwardsaedes bekkui Edwardsaedes imprimens Neomelaniconion lineatopenne Neomelaniconion palpale Aedes cinereus Aedes esoensis Paraedes barraudi Paraedes ostentatio Verrallina (Harbachius) nobukonis Verrallina (Har.) yusafi Verrallina (Verrallina) carmenti Verrallina (Ver.) butleri Verrallina (Neomacleaya) indica Verrallina (Nma.) pseudomediofasciata Aedimorphus domesticus Aedimorphus dalzieli Aedimorphus eritreae Aedimorphus culicinus Aedimorphus alboscutellatus Aedimorphus pampangensis Aedimorphus pipersalatus Aedimorphus taeniorhynchoides Aedimorphus caecus Aedimorphus orbitae Aedimorphus trimaculatus Aedimorphus mediolineatus Aedimorphus ochraceus Aedimorphus pallidostriatus Catageiomyia irritans Catageiomyia argenteopunctatus Catageiomyia tarsalis Elpeytonius apicoannulatus Elpeytonius simulans Aedimorphus punctifemoris Albuginosus marshallii Albuginosus ngong Christophersiomyia gombakensis Christophersiomyia thomsoni Diceromyia furcifer Diceromyia taylori Tewarius agastyai Tewarius reubenae Hu.wauensis Lp.aurimargo As.brevitibia Er.chrysogaster Er.quinquevittatus Ar.Arm.breinli Ar.Arm.subalbatus Ar.Lei.flavus Ar.Lei.longipalpis Lo.amesii Lo.dasyorrhus Di.whartoni Di.franciscoi Di.iyengari Di.scanloni Pa.argenteoventralis.dunni Pa.michaelikati Hz.Hez.complex Hz.Hez.scintillans Hz.Mat.achaetae Hz.Mat.catesi Di.Periskelata Di.micropterus Di.reginae Bc.cavernicola Ay.griffithi Ay.peytoni Bo.eldridgei Bo.helenae Di.meronephada Ud.argyrurus Ud.lucaris Be.aurotaeniatus Ze.gracilis Ze.Lawtoni Sc.albolineata Sc.arboricola St.pandani St.saipanensis St.futunae St.edwardsi St.scutellaris St.riversi St.albopicta St.unilineata St.walbus St.gardnerii.gardnerii St.desmotes St.annandalei St.craggi St.mediopunctata St.perplexa Ae.Cor.cozi St.metallica St.apicoargentea St.africana St.luteocephala St.aegypti St.poweri St.deboeri St.dendrophila St.chemulpoensis St.wadai St.bromeliae St.simpsoni St.woodi Mu.Muc.alternans Mu.Muc.laniger Mu.Pdo.aurantius.aurantius Mu.Pdo.quadripunctis Oc.theobaldi Oc.Buv.edgari Oc.Emp.vigilax Oc.Cul.taeniorhynchus Oc.Cul.mitchellae Oc.spilotus Oc.calumnior Oc.Psk.bancroftianus Oc.Psk.postspiraculosus Oc.Geo.baisasi Oc.Geo.longiforceps Oc.Lev.geoskusea Oc.Rhi.longirostris Oc.Rhi.wardi Oc.Sal.hortator Ae.Can.cancricomes Ae.Can.masculinus Ae.Can.penghuensis Oc.Phl.flavifrons Oc.Phl.calcariae Oc.canadensis.canadensis Oc.cantator Oc.ratcliffei Oc.fitchii Oc.grossbecki Oc.flavescens Oc.squamiger Oc.cantans Oc.stimulans Oc.Excrucians Oc.riparius Oc.andersoni Oc.nivalis Oc.sticticus Oc.aurifer Oc.thibaulti Oc.Gli.aculeatus Oc.Gli.mcdonaldi Oc.Och.infirmatus Oc.Och.scapularis Oc.Chs.fulvusfulvus Oc.Chs.fulvuspallens Oc.Pcx.pertinax Oc.Pcx.serratus Oc.impiger Oc.pullatus Oc.diantaeus Oc.intrudens Oc.hexodontus Oc.punctor Oc.pionips Oc.communis Oc.Rus.refiki Oc.Rus.rusticus Oc.albifasciatus Oc.caballus Oc.spencerii.spencerii Oc.campestris Oc.dorsalis Oc.Aca.zammitii Oc.caspius Oc.Fin.crossi Hk.Hpk.embuensis Hk.Hpk.ingrami Hk.Yam.albocinctus Hk.Yam.seoulensis Oc.Fin.biocellatus Oc.Fin.candidoscutellum Oc.Fin.keefei Hi.gilli Br.alektorovi Br.greenii Va.luteostriata Va.pulchrithorax Ke.dissimilis Ke.harbachi Za.fulgens Za.longipalpis Ra.albilabris Ra.notoscripta Mo.pecuniosa Lu.fengi Mc.Cha.wattensis Mc.Mac.tremula Db.alboannulatus Db.tubbutiensis Pm.argyronotum Pm.papuensis Gc.Hor.fluviatilis Gc.Gec.atropalpus Gc.Gec.epactius Ta.savoryi Ta.togoi Hl.chrysolineata Hl.sherki Gi.pulchriventer Co.Alo.banksi Co.Alo.pseudotaeniatus Co.Col.elsiae Co.Col.macfarlanei Oc.Fin.oreophilus Ji.chungi Ph.gubernatoris Ph.lophoventralis Oc.Pro.kompi Oc.Pro.galindoi Oc.Pro.homoeopus Oc.Pro.terrens Oc.Pro.burgeri Oc.Pro.knabi Oc.Pro.brelandi Oc.Pro.zoosophus Oc.Pro.triseriatus Do.leonis Do.nivea Dn.albotaeniata Dn.harperi Fl.kochi Fl.poicilia Da.echinus Da.geniculata Hw.sexlineata Hw.walkeri Ko.purpureipes Az.ramirezi Ja.Lew.muelleri Ja.Jar.sierrensis Ja.Jar.varipalpus Ab.papago Gy.mediovittata Hg.Con.leucocelaenus Hg.Con.leucotaeniatus Hg.Hag.equinus Hg.Hag.splendens Strict consensus tree Cs.Cus.inornata Cx.Cux.quinquefasciatus Or.signifera Ma.Man.titillans Ps.Gra.columbiae Ps.Gra.jamaicensis Ps.Jan.cyanescens Ps.Jan.ferox Ps.Pso.ciliata Ps.Pso.howardii Fr.vittatus Ia.cavaticus In.pulverulentus Sk.pembaensis Ha.australis Op.Not.chathamicus Op.Opi.fuscus Ae.Adm.lamborni Ae.Adm.albocephalus Ae.Adm.gibbinsi Ae.Adm.quasiunivittatus Ae.Adm.vexans.vexans Ae.Adm.cumminsii Ae.Adm.dentatus Ed.bekkui Ed.imprimens Ne.lineatopenne Ne.palpale Ae.cinereus Ae.esoensis Pr.barraudi Pr.ostentatio Ve.Har.nobukonis Ve.Har.yusafi Ve.Ver.carmenti Ve.Ver.butleri Ve.Nma.indica Ve.Nma.pseudomediofasciata Ae.Adm.domesticus Ae.Adm.dalzieli Ae.Adm.eritreae Ae.Adm.culicinus Ae.Adm.alboscutellatus Ae.Adm.pampangensis Ae.Adm.pipersalatus Ae.Adm.taeniorhynchoides Ae.Adm.caecus Ae.Adm.orbitae Ae.Adm.trimaculatus Ae.Adm.mediolineatus Ae.Adm.ochraceus Ae.Adm.pallidostriatus Ae.Adm.irritans Ae.Adm.argenteopunctatus Ae.Adm.tarsalis Ae.Adm.apicoannulatus Ae.Adm.simulans Ae.Adm.punctifemoris Al.marshallii Al.ngong Cr.gombakensis Cr.thomsoni Di.furcifer Di.taylori Tewarius agastyai Tewarius reubenae Huaedes wauensis Leptosomatomyia aurimargo Alanstonea brevitibia Eretmapodites chrysogaster Eretmapodites quinquevittatus Armigeres (Armigeres) breinli Armigeres (Arm.) subalbatus Armigeres (Leicesteria) flavus Armigeres (Lei.) longipalpis Lorrainea amesii Lorrainea dasyorrhus Petermattinglyius whartoni Petermattinglyius franciscoi Petermattinglyius iyengari Petermattinglyius scanloni Pseudarmigeres a. dunni Pseudarmigeres michaelikati Heizmannia (Heizmannia) complex Hz. (Hez.) scintillans Hz. (Mattinglyia) achaetae Hz. (Mat.) catesi Dendroskusea periskelata Dendroskusea micropterus Dendroskusea reginae Borichinda cavernicola Ayurakitia griffithi Ayurakitia peytoni Bothaella eldridgei Bothaella helenae Catatassomyia meronephada Udaya argyrurus Udaya lucaris Belkinius aurotaeniatus Zeugnomyia gracilis Zeugnomyia lawtoni Scutomyia albolineata Scutomyia arboricola Stegomyia (Bohartius) pandani Stegomyia (Boh.) saipanensis Stegomyia (Zoromorphus) futunae Stegomyia (Actinothrix) edwardsi Stegomyia (Sub. Un.) scutellaris Stegomyia (Sub. Un.) riversi Stegomyia (Sub. Un.) albopicta Stegomyia (Sub. Un.) unilineata Stegomyia (Sub. Un.) w-albus Stegomyia (Sub. Un.) gardnerii Stegomyia (Xyele) desmotes Stegomyia (Heteraspidion) annandalei Stegomyia (Het.) craggi St. (Huangmyia) mediopunctata St. (Hua.) perplexa Cornetius cozi Stegomyia (Sub. Un.) metallica Stegomyia (Sub. Un.) apicoargentea Stegomyia (Sub. Un.) africana Stegomyia (Sub. Un.) luteocephala Stegomyia (Stegomyia) aegypti Stegomyia (Sub. Un.) poweri Stegomyia (Sub. Un.) deboeri Stegomyia (Sub. Un.) dendrophila Stegomyia (Sub. Un.) chemulpoensis Stegomyia (Sub. Un.) wadai Stegomyia (Mukwaya) bromeliae Stegomyia (Muk.) simpsoni Stegomyia (Muk.) woodi Mucidus (Mucidus) alternans Mucidus (Muc.) laniger Mu.Pdo.aurantius.aurantius Mu.Pdo.quadripunctis Oc.theobaldi Oc.Buv.edgari Oc.Emp.vigilax Oc.Cul.taeniorhynchus Oc.Cul.mitchellae Oc.spilotus Oc.calumnior Oc.Psk.bancroftianus Oc.Psk.postspiraculosus Oc.Geo.baisasi Oc.Geo.longiforceps Oc.Lev.geoskusea Oc.Rhi.longirostris Oc.Rhi.wardi Oc.Sal.hortator Ae.Can.cancricomes Ae.Can.masculinus Ae.Can.penghuensis Oc.Phl.flavifrons Oc.Phl.calcariae Oc.canadensis.canadensis Oc.cantator Oc.ratcliffei Oc.fitchii Oc.grossbecki Oc.flavescens Oc.squamiger Oc.cantans Oc.stimulans Oc.Excrucians Oc.riparius Oc.andersoni Oc.nivalis Oc.sticticus Oc.aurifer Oc.thibaulti Oc.Gli.aculeatus Oc.Gli.mcdonaldi Oc.Och.infirmatus Oc.Och.scapularis Oc.Chs.fulvusfulvus Oc.Chs.fulvuspallens Oc.Pcx.pertinax Oc.Pcx.serratus Oc.impiger Oc.pullatus Oc.diantaeus Oc.intrudens Oc.hexodontus Oc.punctor Oc.pionips Oc.communis Oc.Rus.refiki Oc.Rus.rusticus Oc.albifasciatus Oc.caballus Oc.spencerii.spencerii Oc.campestris Oc.dorsalis Oc.Aca.zammitii Oc.caspius Oc.Fin.crossi Hk.Hpk.embuensis Hk.Hpk.ingrami Hk.Yam.albocinctus Hk.Yam.seoulensis Oc.Fin.biocellatus Oc.Fin.candidoscutellum Oc.Fin.keefei Hi.gilli Br.alektorovi Br.greenii Va.luteostriata Va.pulchrithorax Ke.dissimilis Ke.harbachi Za.fulgens Za.longipalpis Ra.albilabris Ra.notoscripta Mo.pecuniosa Lu.fengi Mc.Cha.wattensis Mc.Mac.tremula Db.alboannulatus Db.tubbutiensis Pm.argyronotum Pm.papuensis Gc.Hor.fluviatilis Gc.Gec.atropalpus Gc.Gec.epactius Ta.savoryi Ta.togoi Hl.chrysolineata Hl.sherki Gi.pulchriventer Co.Alo.banksi Co.Alo.pseudotaeniatus Co.Col.elsiae Co.Col.macfarlanei Oc.Fin.oreophilus Ji.chungi Ph.gubernatoris Ph.lophoventralis Oc.Pro.kompi Oc.Pro.galindoi Oc.Pro.homoeopus Oc.Pro.terrens Oc.Pro.burgeri Oc.Pro.knabi Oc.Pro.brelandi Oc.Pro.zoosophus Oc.Pro.triseriatus Do.leonis Do.nivea Dn.albotaeniata Dn.harperi Fl.kochi Fl.poicilia Da.echinus Da.geniculata Hw.sexlineata Hw.walkeri Ko.purpureipes Az.ramirezi Ja.Lew.muelleri Ja.Jar.sierrensis Ja.Jar.varipalpus Ab.papago Gy.mediovittata Hg.Con.leucocelaenus Hg.Con.leucotaeniatus Hg.Hag.equinus Hg.Hag.splendens Strict consensus tree Cs.Cus.inornata Cx.Cux.quinquefasciatus Or.signifera Ma.Man.titillans Ps.Gra.columbiae Ps.Gra.jamaicensis Ps.Jan.cyanescens Ps.Jan.ferox Ps.Pso.ciliata Ps.Pso.howardii Fr.vittatus Ia.cavaticus In.pulverulentus Sk.pembaensis Ha.australis Op.Not.chathamicus Op.Opi.fuscus Ae.Adm.lamborni Ae.Adm.albocephalus Ae.Adm.gibbinsi Ae.Adm.quasiunivittatus Ae.Adm.vexans.vexans Ae.Adm.cumminsii Ae.Adm.dentatus Ed.bekkui Ed.imprimens Ne.lineatopenne Ne.palpale Ae.cinereus Ae.esoensis Pr.barraudi Pr.ostentatio Ve.Har.nobukonis Ve.Har.yusafi Ve.Ver.carmenti Ve.Ver.butleri Ve.Nma.indica Ve.Nma.pseudomediofasciata Ae.Adm.domesticus Ae.Adm.dalzieli Ae.Adm.eritreae Ae.Adm.culicinus Ae.Adm.alboscutellatus Ae.Adm.pampangensis Ae.Adm.pipersalatus Ae.Adm.taeniorhynchoides Ae.Adm.caecus Ae.Adm.orbitae Ae.Adm.trimaculatus Ae.Adm.mediolineatus Ae.Adm.ochraceus Ae.Adm.pallidostriatus Ae.Adm.irritans Ae.Adm.argenteopunctatus Ae.Adm.tarsalis Ae.Adm.apicoannulatus Ae.Adm.simulans Ae.Adm.punctifemoris Al.marshallii Al.ngong Cr.gombakensis Cr.thomsoni Di.furcifer Di.taylori Te.agastyai Te.reubenae Hu.wauensis Lp.aurimargo As.brevitibia Er.chrysogaster Er.quinquevittatus Ar.Arm.breinli Ar.Arm.subalbatus Ar.Lei.flavus Ar.Lei.longipalpis Lo.amesii Lo.dasyorrhus Di.whartoni Di.franciscoi Di.iyengari Di.scanloni Pa.argenteoventralis.dunni Pa.michaelikati Hz.Hez.complex Hz.Hez.scintillans Hz.Mat.achaetae Hz.Mat.catesi Di.Periskelata Di.micropterus Di.reginae Bc.cavernicola Ay.griffithi Ay.peytoni Bo.eldridgei Bo.helenae Di.meronephada Ud.argyrurus Ud.lucaris Be.aurotaeniatus Ze.gracilis Ze.Lawtoni Sc.albolineata Sc.arboricola St.pandani St.saipanensis St.futunae St.edwardsi St.scutellaris St.riversi St.albopicta St.unilineata St.walbus St.gardnerii.gardnerii St.desmotes St.annandalei St.craggi St.mediopunctata St.perplexa Ae.Cor.cozi St.metallica St.apicoargentea St.africana St.luteocephala St.aegypti St.poweri St.deboeri St.dendrophila St.chemulpoensis St.wadai St.bromeliae St.simpsoni St.woodi

Mucidus (Muc.) laniger Mucidus (Pardomyia) aurantius Mucidus (Pdo.) quadripunctis Oc. (Sub. Un.) theobaldi Oc. (Buvirilia) edgari Oc. (Empihals) vigilax Oc. (Cul.) taeniorhynchus Oc. (Culicelsa) mitchellae Oc. (Leptokeneon) spilotus Oc. (Sub. Un.) calumnior Oc. (Pseudoskusea) bancroftianus Oc. (Psk.) postspiraculosus Geoskusea baisasi Geoskusea longiforceps Levua geoskusea Rhinskusea longirostris Rhinoskusea wardi Sallumia hortator Cancraedes cancricomes Cancraedes masculinus Cancraedes penghuensis Ochlerotatus (Pholeomyia) flavifrons Oc. (Phl.) calcariae Oc. (Culicada) canadensis Oc. (Sub. Un.) cantator Oc. (Sub. Un.) ratcliffei Oc. (Sub. Un.) fitchii Oc. (Sub. Un.) grossbecki Oc. (Sub. Un.) flavescens Oc. (Sub. Un.) squamiger Oc. (Sub. Un.) cantans Oc. (Sub. Un.) stimulans Oc. (Sub. Un.) excrucians Oc. (Sub. Un.) riparius Oc. (Sub. Un.) andersoni Oc. (Sub. Un.) nivalis Oc. (Sub. Un.) sticticus Oc. (Sub. Un.) aurifer Oc. (Sub. Un.) thibaulti Oc. (Gilesia) aculeatus Oc. (Gli.) mcdonaldi Oc. (Ochlerotatus) infirmatus Oc. (Och.) scapularis Oc. (Chysoconops) fulvus Oc. (Chs.) fulvus pallens Oc. (Protoculex) pertinax Oc. (Pcx.) serratus Oc. (Sub. Un.) impiger Oc. (Sub. Un.) pullatus Oc. (Woodius) diantaeus Oc. (Woo.) intrudens Oc. (Sub. Un.) hexodontus Oc. (Sub. Un.) punctor Oc. (Sub. Un.) pionips Oc. (Sub. Un.) communis Oc. (Rusticoidus) refiki Oc. (Rus.) rusticus Oc. (Sub. Un.) albifasciatus Oc. (Juppius) caballus Oc. (Sub. Un.) spencerii Oc. (Sub. Un.) campestris Oc. (Sub. Un.) dorsalis Acartomyia zammitii Oc. (Sub. Un.) caspius ‘Oc.’ (’Finlaya’) crossi Hopkinsius (Hopkinsius) embuensis Hopkinsius (Hpk.) ingrami Hopkinsius (Yamada) albocinctus Hopkinsius (Yam.) seoulensis

Oc.Fin.candidoscutellum Oc.Fin.keefei Hi.gilli Br.alektorovi Br.greenii Va.luteostriata Va.pulchrithorax Ke.dissimilis Ke.harbachi Za.fulgens Za.longipalpis Ra.albilabris Ra.notoscripta Mo.pecuniosa Lu.fengi Mc.Cha.wattensis Mc.Mac.tremula Db.alboannulatus Db.tubbutiensis Pm.argyronotum Pm.papuensis Gc.Hor.fluviatilis Gc.Gec.atropalpus Gc.Gec.epactius Ta.savoryi Ta.togoi Hl.chrysolineata Hl.sherki Gi.pulchriventer Co.Alo.banksi Co.Alo.pseudotaeniatus Co.Col.elsiae Co.Col.macfarlanei Oc.Fin.oreophilus Ji.chungi Ph.gubernatoris Ph.lophoventralis Oc.Pro.kompi Oc.Pro.galindoi Oc.Pro.homoeopus Oc.Pro.terrens Oc.Pro.burgeri Oc.Pro.knabi Oc.Pro.brelandi Oc.Pro.zoosophus Oc.Pro.triseriatus Do.leonis Do.nivea Dn.albotaeniata Dn.harperi Fl.kochi Fl.poicilia Da.echinus Da.geniculata Hw.sexlineata Hw.walkeri Ko.purpureipes Az.ramirezi Ja.Lew.muelleri Ja.Jar.sierrensis Ja.Jar.varipalpus Ab.papago Gy.mediovittata Hg.Con.leucocelaenus Hg.Con.leucotaeniatus Hg.Hag.equinus Hg.Hag.splendens Strict consensus tree Cs.Cus.inornata Cx.Cux.quinquefasciatus Or.signifera Ma.Man.titillans Ps.Gra.columbiae Ps.Gra.jamaicensis Ps.Jan.cyanescens Ps.Jan.ferox Ps.Pso.ciliata Ps.Pso.howardii Fr.vittatus Ia.cavaticus In.pulverulentus Sk.pembaensis Ha.australis Op.Not.chathamicus Op.Opi.fuscus Ae.Adm.lamborni Ae.Adm.albocephalus Ae.Adm.gibbinsi Ae.Adm.quasiunivittatus Ae.Adm.vexans.vexans Ae.Adm.cumminsii Ae.Adm.dentatus Ed.bekkui Ed.imprimens Ne.lineatopenne Ne.palpale Ae.cinereus Ae.esoensis Pr.barraudi Pr.ostentatio Ve.Har.nobukonis Ve.Har.yusafi Ve.Ver.carmenti Ve.Ver.butleri Ve.Nma.indica Ve.Nma.pseudomediofasciata Ae.Adm.domesticus Ae.Adm.dalzieli Ae.Adm.eritreae Ae.Adm.culicinus Ae.Adm.alboscutellatus Ae.Adm.pampangensis Ae.Adm.pipersalatus Ae.Adm.taeniorhynchoides Ae.Adm.caecus Ae.Adm.orbitae Ae.Adm.trimaculatus Ae.Adm.mediolineatus Ae.Adm.ochraceus Ae.Adm.pallidostriatus Ae.Adm.irritans Ae.Adm.argenteopunctatus Ae.Adm.tarsalis Ae.Adm.apicoannulatus Ae.Adm.simulans Ae.Adm.punctifemoris Al.marshallii Al.ngong Cr.gombakensis Cr.thomsoni Di.furcifer Di.taylori Te.agastyai Te.reubenae Hu.wauensis Lp.aurimargo As.brevitibia Er.chrysogaster Er.quinquevittatus Ar.Arm.breinli Ar.Arm.subalbatus Ar.Lei.flavus Ar.Lei.longipalpis Lo.amesii Lo.dasyorrhus Di.whartoni Di.franciscoi Di.iyengari Di.scanloni Pa.argenteoventralis.dunni Pa.michaelikati Hz.Hez.complex Hz.Hez.scintillans Hz.Mat.achaetae Hz.Mat.catesi Di.Periskelata Di.micropterus Di.reginae Bc.cavernicola Ay.griffithi Ay.peytoni Bo.eldridgei Bo.helenae Di.meronephada Ud.argyrurus Ud.lucaris Be.aurotaeniatus Ze.gracilis Ze.Lawtoni Sc.albolineata Sc.arboricola St.pandani St.saipanensis St.futunae St.edwardsi St.scutellaris St.riversi St.albopicta St.unilineata St.walbus St.gardnerii.gardnerii St.desmotes St.annandalei St.craggi St.mediopunctata St.perplexa Ae.Cor.cozi St.metallica St.apicoargentea St.africana St.luteocephala St.aegypti St.poweri St.deboeri St.dendrophila St.chemulpoensis St.wadai St.bromeliae St.simpsoni St.woodi Mu.Muc.alternans Mu.Muc.laniger Mu.Pdo.aurantius.aurantius Mu.Pdo.quadripunctis Oc.theobaldi Oc.Buv.edgari Oc.Emp.vigilax Oc.Cul.taeniorhynchus Oc.Cul.mitchellae Oc.spilotus Oc.calumnior Oc.Psk.bancroftianus Oc.Psk.postspiraculosus Oc.Geo.baisasi Oc.Geo.longiforceps Oc.Lev.geoskusea Oc.Rhi.longirostris Oc.Rhi.wardi Oc.Sal.hortator Ae.Can.cancricomes Ae.Can.masculinus Ae.Can.penghuensis Oc.Phl.flavifrons Oc.Phl.calcariae Oc.canadensis.canadensis Oc.cantator Oc.ratcliffei Oc.fitchii Oc.grossbecki Oc.flavescens Oc.squamiger Oc.cantans Oc.stimulans Oc.Excrucians Oc.riparius Oc.andersoni Oc.nivalis Oc.sticticus Oc.aurifer Oc.thibaulti Oc.Gli.aculeatus Oc.Gli.mcdonaldi Oc.Och.infirmatus Oc.Och.scapularis Oc.Chs.fulvusfulvus Oc.Chs.fulvuspallens Oc.Pcx.pertinax Oc.Pcx.serratus Oc.impiger Oc.pullatus Oc.diantaeus Oc.intrudens Oc.hexodontus Oc.punctor Oc.pionips Oc.communis Oc.Rus.refiki Oc.Rus.rusticus Oc.albifasciatus Oc.caballus Oc.spencerii.spencerii Oc.campestris Oc.dorsalis Oc.Aca.zammitii Oc.caspius Oc.Fin.crossi Hk.Hpk.embuensis Hk.Hpk.ingrami Hk.Yam.albocinctus Hopkinsius (Yam.) seoulensis 'Oc.' ('Finlaya') biocellatus ‘Oc.’ (‘Fin.’) candidoscutellum ‘Oc.’ (‘Fin.’) keefei Himalaius gilli Bruceharrisonius alektorovi Bruceharrisonius greenii Vansomerensus luteostriata Vansomerensis pulchrithorax Kenknightia dissimilis Kenknightia harbachi Zavortinkius fulgens Zavortinkius longipalpis Rampamyia albilabris Rampamyia notoscripta Molpemyia pecuniosa Luius fengi Macleaya (Chaetocuiomyia) wattensis Macleaya (Macleaya) tremlula Dobrotworskyius alboannulatus Dobrotworskyius tubbutiensis Patmarksia argyronotum Patmarksia papuensis Georgecraigius (Horsfallius) fluviatilis Gc. (Georgecraigius) atropalpus Gc. (Georgecraigius) epactius Tanakius savoryi Tanakius togoi Hulecoeteomyia chrysolineata Hulecoeteomyia sherki Gilesius pulchriventer Collessius (Alloemyia) banksi Collessius (Alo.) pseudotaeniatus Collessius (Collessius) elsiae Collessius (Col.) macfarlanei ‘Oc.’ (’Fin.’) oreophilus Jihliensius chungi Phagomyia gubernatoris Phagomyia lophoventralis ‘Oc.’ (’Pro.’) kompi ‘Oc.’ (’Pro.’) galindoi ‘Oc.’ (’Pro.’) homoeopus ‘Oc.’ (’Pro.’) terrens ‘Oc.’ (’Pro.’) burgeri ‘Oc.’ (’Pro.’) knabi ‘Oc.’ (’Protomacleaya’) brelandi ‘Oc.’ (’Pro.’) zoosophus ‘Oc.’ (’Pro.’) triseriatus Downsiomyia leonis Downsiomyia nivea Danielsia albotaeniata Danielsia harperi Finlaya kochi Finlaya poicilia Dahliana echinus Dahliana geniculata Howardina sexlineata Howardina walkeri Kompia purpureipes Aztecaedes ramirezi Lewneilsenius muelleri Jarnellius sierrensis Jarnellius varipalpus Abraedes papago Gymnometopa mediovittata Haemagogus (Conopostegus) leucocelaenus Haemagogus (Con.) leucotaeniatus Haemagogus (Haemagogus) equinus Haemagogus (Hag.) splendens