Aedini (Reinert, Harbach & Kitching, 2004)

Reinert, John F., Harbach, Ralph E. & Kitching, Ian J., 2009, Phylogeny and classification of tribe Aedini (Diptera: Culicidae), Zoological Journal of the Linnean Society 157 (4), pp. 700-794 : 704-710

Aedini ingroup taxa

Abraedes Zavortink

Aedes Meigen

‘ Aedes ’ sensu auctorum ‘ Aedimorphus ’ Theobald

‘ Cancraedes ’ Edwards

‘ Cornetius ’ Huang Alanstonea Mattingly

Albuginosus Reinert

Armigeres Theobald Armigeres

Leicesteria Theobald

Ayurakitia Thurman

Aztecaedes Zavortink

Belkinius Reinert

Borichinda Harbach &

Rattanarithikul

Bothaella Reinert papago (Zavortink) * 1 cinereus Meigen * 2 esoensis Yamada 2 albocephalus (Theobald) 3 alboscutellatus (Theobald) 2,4,5 apicoannulatus (Edwards) 3 argenteopunctatus (Theobald) 3

caecus (Theobald) 4 culicinus Edwards 4 cumminsii (Theobald) 3 dalzieli (Theobald) 3 dentatus (Theobald) 3 domesticus (Theobald) * 3 eritreae Lewis 3 gibbinsi Edwards 3 irritans (Theobald) 3 lamborni Edwards 3 mediolineatus (Theobald) 4 ochraceus (Theobald) 3 orbitae Edwards 4 pallidostriatus (Theobald) 4 pampangensis (Ludlow) 4 pipersalatus (Giles) 4 punctifemoris (Ludlow) 4 quasiunivittatus (Theobald) 3 simulans (Newstead & Carter) 3 taeniorhynchoides (Christophers) 4 tarsalis (Newstead) 3 trimaculatus (Theobald) 4 vexans vexans (Meigen) 1,2,3,4,5 cancricomes Edwards * 4 masculinus Mattingly 4 penghuensis Lien 4

cozi Cornet * 3 brevitibia (Edwards) 4 marshallii (Theobald) * 3

ngong (van Someren) 3

breinli (Taylor) 5 subalbatus (Coquillett) 2,4 flavus (Leicester) 4 longipalpis (Leicester) * 4 griffithi Thurman * 4 peytoni (Reinert) 4 ramirezi (Vargas & Downs) * 1 aurotaeniatus (Edwards) * 4 cavernicola Rattanarithikul & Harbach * 4

eldridgei (Reinert) 4 helenae (Reinert) * 4 Genus Subgenus Species Distribution

Bruceharrisonius Reinert alektorovi (Stackelberg) 2 greenii (Theobald) * 4

Christophersiomyia Barraud gombakensis (Mattingly) 4 thomsoni (Theobald) * 4

Collessius Reinert , Alloeomyia Reinert , banksi (Edwards) 4 Harbach & Kitching Harbach & Kitching pseudotaeniatus (Giles) * 4 Collessius elsiae (Barraud) 4

macfarlanei (Edwards) * 4

Dahliana Reinert , echinus (Edwards) 2 Harbach & Kitching geniculata (Olivier) * 2 Danielsia Theobald albotaeniata Leicester * 4 harperi (Knight) 4

Diceromyia Theobald franciscoi (Mattingly) 4 furcifer (Edwards) * 3

iyengari (Edwards) 4

meronephada (Dyar & Shannon) 4

micropterus (Giles) 4

periskelata (Giles) 4

reginae (Edwards) 4

scanloni (Reinert) 4

taylori (Edwards) 3

whartoni (Mattingly) 4

Dobrotworskyius Reinert , alboannulatus (Macquart) 5 Harbach & Kitching tubbutiensis (Dobrotworsky) * 5 Downsiomyia Vargas leonis (Colless) 4 nivea (Ludlow) * 4

Edwardsaedes Belkin bekkui (Mogi) 2 imprimens (Walker) * 2,4,5

Eretmapodites Theobald chrysogaster Graham 3 quinquevittatus Theobald * 3

Finlaya Theobald kochi (Dönitz) * 5 poicilia Theobald 4,5

Fredwardsius Reinert vittatus (Bigot) * 2,3,4 Georgecraigius Reinert , Georgecraigius atropalpus (Coquillett) * 1,2* Harbach & Kitching epactius (Dyar & Knab) 1,6 Horsfallius Reinert , fluviatilis (Lutz) * 6

Harbach & Kitching

Gilesius Reinert, Harbach pulchriventer (Giles) * 2,4 & Kitching Gymnometopa Coquillett mediovittata (Coquillett) * 6 Haemag ogus Williston Conopostegus Dyar leucocelaenus (Dyar & Shannon) * 6 leucotaeniatus (Komp) 6

Haemagogus equinus Theobald 6

splendens Williston * 6

Halaedes Belkin australis (Erichson) * 5 Heizmannia Ludlow Heizmannia complex (Theobald) 4 scintillans Ludlow * 4

Mattinglyia Lien achaetae (Leicester) 4

catesi (Lien) * 4

Himalaius Reinert , gilli (Barraud) * 4 Harbach & Kitching Genus Subgenus Species Distribution Hopkinsius Reinert , Hopkinsius

Harbach & Kitching

Yamada Reinert, Harbach & Kitching Howardina Theobald

Huaedes Huang

Hulecoeteomyia Theobald

Indusius Edwards Isoaedes Reinert Jarnellius Reinert , Jarnellius Harbach & Kitching

Lewnielsenius Reinert,

Harbach & Kitching Jihlienius Reinert, Harbach & Kitching Kenknightia Reinert

Kompia Aitkin

Leptosomatomyia Theobald

Lorrainea Belkin

Luius Reinert, Harbach

& Kitching

Macleaya Theobald Chaetocruiomyia Theobald

Macleaya

Molpemyia Theobald

Mucidus Theobald Mucidus

Pardomyia Theobald

Neomelaniconion Newstead Ochlerotatus Lynch Acartomyia Reinert, Arribálzaga Harbach & Kitching Buvirilia Reinert, Harbach & Kitching Chrysoconops Goeldi

Culicelsa Felt

Empihals Reinert ,

Harbach & Kitching

Geoskusea Edwards

Gilesia Theobald

Levua Stone & Bohart Ochlerotatus embuensis (Edwards) 3 ingrami (Edwards) * 3 albocinctus (Barraud) 4 seoulensis (Yamada) * 2 sexlineata (Theobald) 6 walkeri (Theobald) * 6 wauensis (Huang) * 5 chrysolineata (Theobald) * 4 sherki (Knight) 4 pulverulentus (Edwards) * 4 cavaticus (Reinert) * 4 sierrensis (Ludlow) 1 varipalpus (Coquillett) * 1 muelleri (Dyar) * 1

chungi (Lien) * 4

dissimilis (Leicester) * 4 harbachi (Reinert) 4 purpureipes (Aitken) * 1 aurimargo (Edwards) * 5 amesii (Ludlow) 4 dasyorrhus (King & Hoogstraal) * 5 fengi (Edwards) * 4

wattensis (Taylor) 5

tremula Theobald* 5 pecuniosa (Edwards) 5 alternans (Westwood) * 5 laniger (Wiedemann) 4 aurantius aurantius (Theobald) * 4,5 quadripunctis (Ludlow) 4 lineatopenne (Ludlow) 4,5 palpale Newstead* 3 zammitii (Theobald) * 2

edgari (Stone & Rosen) * 5

fulvus fulvus (Wiedemann) * 6 fulvus pallens (Ross) 1,6 mitchellae (Dyar) 1 taeniorhynchus (Wiedemann) * 1,6 vigilax (Skuse) * 3*,4,5

baisasi (Knight & Hull) 4 longiforceps (Edwards) 5 aculeatus (Theobald) * 5 mcdonaldi (Belkin) 5 geoskusea (Amos) * 5 infirmatus (Dyar & Knab) 1 scapularis (Rondani) * 1,6 Genus Subgenus Species Distribution

Protoculex Felt pertinax (Grabham) 6 serratus (Theobald) * 6 Pholeomyia Reinert , calcariae (Marks) * 5 Harbach & Kitching flavifrons (Skuse) 5 Pseudoskusea Theobald bancroftianus (Edwards) 5 postspiraculosus (Dobrotworsky) 5 Rhinoskusea Edwards longirostris (Leicester) * 4,5 wardi (Reinert) 4 Rusticoidus Shevchenko refiki (Medschid) * 2 & Prudkina rusticus (Rossi) 2 Sallumia Reinert , hortator (Dyar & Knab) * 6 Harbach & Kitching Unassigned to subgenus albifasciatus (Macquart) 6 andersoni (Edwards) 5 aurifer (Coquillett) 1 caballus (Theobald) 2,3 calumnior (Belkin, Heinemann & Page) 6 campestris (Dyar & Knab) 1 canadensis canadensis (Theobald) 1 cantans (Meigen) 2 cantator (Coquillett) 1 caspius (Pallas) 2 communis (de Geer) 1,2 diantaeus (Howard, Dyar & Knab) 1,2 dorsalis (Meigen) 1,2,4 excrucians (Walker) 1,2 fitchii (Felt & Young) 1,2 flavescens (Müller) 1,2 grossbecki (Dyar & Knab) 1 hexodontus (Dyar) 1,2 impiger (Walker) 1,2 intrudens (Dyar) 1,2 nivalis (Edwards) 5 pionips (Dyar) 1,2 pullatus (Coquillett) 1,2 punctor (Kirby) 1,2 ratcliffei (Marks) 5 riparius (Dyar & Knab) 1,2 spencerii spencerii (Theobald) 1 spilotus (Marks) 5 squamiger (Coquillett) 1 sticticus (Meigen) 1,2 stimulans (Walker) 1 theobaldi (Taylor) 5 thibaulti (Dyar & Knab) 1,2

‘ Ochlerotatus ’ sensu ‘ Finlaya ’ sensu auctorum biocellatus (Taylor) 5

auctorum candidoscutellum (Marks) 5 crossi (Lien) 4 keefei (King & Hoogstraal) 5 oreophilus Edwards 2,4

Genus Subgenus Species Distribution

‘ Protomacleaya ’ sensu auctorum

Opifex Hutton Nothoskusea Dumbleton

Opifex

Paraedes Edwards

Patmarksia Reinert, Harbach & Kitching

Phagomyia Theobald

Pseudarmigeres Stone & Knight

Psorophora Robineau- Grabhamia Desvoidy Theobald

Janthinosoma Lynch Arribálzaga

Psorophora Rampamyia Reinert, Harbach & Kitching

Scutomyia Theobald

Skusea Theobald

Stegomyia Theobald brelandi (Zavortink) 1 burgeri (Zavortink) 1 galindoi (Schick) 6 homoeopus (Dyar) 6 knabi (Coquillett) 6 kompi (Vargas & Downs) 6 terrens (Walker) 6 triseriatus (Say) * 1 zoosophus (Dyar & Knab) 1 chathamicus (Dumbleton) * 5

fuscus Hutton * 5 barraudi Edwards * 4 ostentatio (Leicester) 4 argyronotum (Belkin) 5 papuensis (Taylor) * 5 gubernatoris (Giles) * 4 lophoventralis (Theobald) 4 argenteoventralis dunni (Evans) * 3 michaelikati (van Someren) 3 columbiae (Dyar & Knab) 1,6 jamaicensis Theobald * 6 cyanescens (Coquillett) 1,6 ferox (von Humboldt) 1,6 ciliata (Fabricius) * 1,6 howardii Coquillett 1,6 albilabris (Edwards) 5 notoscripta (Skuse) * 5 albolineata Theobald * 4,5 arboricola (Knight & Rozeboom) 4 pembaensis (Theobald) * 3 aegypti (Linnaeus) * 1*,2*,3,4*, 5*,6* africana Theobald 3 albopicta (Skuse) 1*,2,3*,4,5, 6* annandalei Theobald 4 apicoargentea Theobald 3 bromeliae (Edwards) 3 chemulpoensis (Yamada) 2 craggi Barraud 4 deboeri (Edwards) 3 dendrophila (Edwards) 3 desmotes Giles 4 edwardsi Barraud 4 futunae (Belkin) 5 gardnerii gardnerii Ludlow 4 luteocephala Newstead 3 mediopunctata Theobald 4 metallica (Edwards) 3 pandani (Stone) 5 perplexa Leicester 4 poweri Theobald 3 riversi (Bohart & Ingram) 4 saipanensis (Stone) 5 scutellaris (Walker) 5

with 400¥ magnification is needed to observe the very slender distal parts of many setae of larvae and pupae for the measurement of total length. Pinned adults were examined with a binocular stereomicroscope utilizing cold white light delivered by a fiber-optic illumination system. An adjustable examination stage with biaxial rotation capability (see fig. 125.3 in Russell et al., 1963) allowed observation and illumination of specimens at any angle.

In general, we examined three to six specimens (range 1–20) of each life stage and structure for each species. However, we had to code some characters from literature sources for those taxa where specimens of certain life stage were unavailable (i.e. characteristics of most eggs and some fourth-instar larvae and pupae). Examples of species not available in the larval stage are ‘ Ae. (Adm.)’ apicoannulatus, Hz. (Mat.) catesi , Oc. andersoni, Oc. (Phl.) flavifrons, Oc. (Chs.) fulvus fulvus and Oc. nivalis , and the pupal stage are ‘ Ae. (Adm.)’ apicoannulatus , ‘ Ae. (Adm.)’ argenteopunctatus , Db. alboannulatus and Oc. (Chs.) fulvus fulvus . Missing data are denoted by a ‘?’ in the data set (Appendix 1). States of continuous characters were determined either by clear gaps in the observed counts or measurements (e.g. character 36), or by reference to observed intraspecific variation (e.g. character 50). Characters that could not be scored due to absence of homologous structures (‘dependent characters’) are indicated by a dash ‘-’, e.g. conditions of setae 6-S and 9-S on the larval siphon of Mansonia , which lacks these setae. All multistate characters were initially treated as unordered. However, it may be argued that continuous measurement characters, such as lengths or ratios derived from lengths, should be treated as ordered sequences ( Thiele, 1993). Hence we also undertook analyses with the 14 such length ratio characters (13, 14, 17, 20, 62, 67, 72, 84, 107, 138, 161, 273, 280 and 318; see below) treated as ordered. Polymorphic characters are explicitly coded as exhibiting only those states observed. A few characters used in our previous studies ( Reinert et al., 2004, 2006, 2008) were re-evaluated and modified, and a number of new characters were added to complete the present data set.

Specimens from the following collections were examined during the study: National Museum of Natural History, Smithsonian Institution, Washington, District of Columbia, USA ; The Natural History Museum , London, UK ; Bohart Museum , University of California, Davis, California, USA ; Canadian National Collection of Insects , Arachnids and Nematodes, Ottawa, Ontario, Canada ; Department of Zoology and Entomology, University of Queensland, Brisbane, Australia ; Australian National Insect Collection , CSIRO Division of Entomology, Canberra, Australia ; Laboratoire / Cellule Entomologie, EID Mediterranée, Montpellier, France ; Connecticut Agricultural Experiment Station , New Haven, Connecticut, USA ; Florida State Collection of Arthropods , Division of Plant Industry, Gainesville, Florida, USA ; Florida Medical Entomology Laboratory , University of Florida, Vero Beach, Florida, USA ; National Institute of Malariology , Parasitology and Entomology, Hanoi, Vietnam ; Center for Research in Medical Entomology, Madurai, Tamil Nadu, India ; Vector Control Research Center , Indira Nagar, Pondicherry, India ; Laboratoire de Taxonomie des Vecteurs , Centre IRD de Montpellier, France ; and the collection of the first author.

Abbreviations used in the text for generic-level taxa follow Reinert (2001a) and Reinert et al. (2006, 2008). New generic-level taxa proposed herein are described in the section following the ‘Concluding Comments’. Abbreviations for all genera of Aedini recognized as a result of the present study are listed in the ‘Abbreviations of Aedine Genera’ section following the ‘New Generic-Level Taxa’. All currently valid species of Aedini , with their generic and subgeneric placement, are listed in Appendix 2. Due to the unavailability of specimens for study and inadequate published descriptions, some species could not be assigned to generic-level taxa with certainty and remain in the category of incertae sedis.