SOUTHEAST ASIAN J TROP MED PUBLIC HEALTH

AEDINE SPECIES (DIPTERA: CULICIDAE: ) OCCURRING IN THAILAND WITH THEIR CURRENT GENERIC AND SUBGENERIC STATUS, AND NEW COUNTRY RECORDS

John F Reinert1, Ralph E Harbach2 and Rampa Rattanarithikul3

1Center for Medical, Agricultural and Veterinary (CMAVE), United States Department of Agriculture, Agricultural Research Service, Gainesville, and collaborator, Walter Reed Biosystematics Unit (WRBU), National Museum of Natural History, Smithsonian Institution, Washington, DC, USA; 2Department of Entomology, The Natural History Museum, Cromwell Road, London, United Kingdom; 3Museum of World and Natural Wonders, Chiang Mai, Thailand

Abstract. A list of the currently valid generic and species-level taxa of tribe Aedini in Thailand is provided. Its purpose is to enable entomologists, public health personnel and mosquito control workers to more quickly become familiar with recent formal taxonomic changes within the tribe. New country records of four species of are reported.

INTRODUCTION of eggs, fourth-instar larvae, pupae, females and genitalia, and males and genitalia, and 119 ae- Harrison et al (1990) provided the most re- dine species from all genera and subgenera of cent list of mosquito species (Culicidae) occur- the tribe recognized at the time. ring in Thailand. Because recent publications on The current classification that resulted from tribe Aedini have resulted in generic and/or the phylogenetic analyses, restricts subgeneric changes to mosquito species occur- Meigen to species in the former subgenus Aedes ring in Thailand, we provide here an updated list- which includes only taxa outside of Thailand. ing of these species. Genera Ayurakitia Thurman Alanstonea Mattingly, Bothaella Reinert, and Verrallina Theobald were resurrected to ge- Christophersiomyia Barraud, Diceromyia neric rank by Reinert (1999, 2000a). Genus Theobald, Edwardsaedes Belkin, was resurrected from synonymy in Reinert, Isoaedes Reinert, Lorrainea Belkin, the composite genus Aedes Meigen and the in- Neomelaniconion Newstead, Paraedes Edwards, cluded subgenera and species were transferred Scutomyia Theobald, and Stegomyia Theobald, to it (Reinert, 2000b). The new subgenus all formerly subgenera of Aedes, are now rec- Bruceharrisonius was described in Ochlerotatus ognized as genera. Former subgenera of Lynch Arribalzaga by Reinert (2003a). Heizmannia Ochlerotatus Lynch Arribalzaga, ie, Finlaya thelmae Mattingly was transferred from subge- Theobald, Kenknightia Reinert, Mucidus nus Mattinglyia Lien to subgenus Heizmannia Theobald, and Rhinoskusea Edwards, also have Ludlow (Reinert, 2003b). Further extensive revi- full generic status. Downsiomyia Vargas was el- sion and modification of Aedini was provided by evated to generic rank from synonymy with Reinert et al (2004) and involved the development Finlaya and includes the species formally in the of phylogenetic relationships resulted from cla- Niveus Group. Genus Tanakaius Reinert, distic analyses of 172 morphological characters Harbach and Kitching was described as new by Correspondence: John F Reinert, Center for Medical, Reinert et al (2004). Armigeres Theobald, Agricultural and Veterinary Entomology (CMAVE), Ayurakitia Thurman, Heizmannia Ludlow, Udaya United States Department of Agriculture, Agricultural Thurman, Verrallina Theobald, and Zeugnomyia Research Service, 1600/1700 SW 23rd Drive, Leicester are still recognized as genera and their Gainesville, FL 32608-1067, USA. Tel: 352 274 5997 subgeneric classifications remain unchanged. E-mail: [email protected] Adhering to the International Code of Zoological

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Nomenclature (International Commission on Genus Armigeres Theobald, 1901 Zoological Nomenclature, 1999), the termina- Subgenus Armigeres Theobald, 1901 tions of some species names were changed to Ar. (Arm.) aureolineatus (Leicester, 1908) agree in gender with newly recognized genera. Ar. (Arm.) bhayungi Thurman and Twenty-six genera and 162 species/sub- Thurman, 1958 species of Aedini are currently recognized in Ar. (Arm.) confusus Edwards, 1915 Thailand, but only 129 species are now classi- Ar. (Arm.) foliatus Brug, 1931 fied in one or other of the genera. In this coun- Ar. (Arm.) jugraensis (Leicester, 1908) try, species previously included in subgenera Ar. (Arm.) kesseli Ramalingam, 1987 Bruceharrisonius Reinert (2 species), Finlaya Ar. (Arm.) kuchingensis Edwards, 1915 Theobald (19 species), and Ochlerotatus (1 spe- Ar. (Arm.) malayi (Theobald, 1901) cies) of Ochlerotatus, and 9 species of subge- Ar. (Arm.) maximus Edwards, 1922 nus Aedimorphus Theobald and 2 species of Ar. (Arm.) moultoni Edwards, 1914 subgenus Cancraedes Edwards of Aedes can- Ar. (Arm.) subalbatus (Coquillett, 1898) not be placed in a currently recognized genus Ar. (Arm.) theobaldi Barraud, 1934 and are regarded as incertae sedis (a Latin term Subgenus Leicesteria Theobald, 1904 meaning ‘of uncertain taxonomic position’) within Ar. (Lei.) annulipalpis (Theobald, 1910) Aedini. These species are indicated in the list Ar. (Lei.) annulitarsis (Leicester, 1908) below as belonging to genus (subgenus) of au- Ar. (Lei.) balteatus Macdonald, 1960 thors, eg Aedes (Aedimorphus) sensu auctorum. Ar. (Lei.) cingulatus (Leicester, 1908) The incertae sedis generic-level designations are Ar. (Lei.) dentatus Barraud, 1927 not to be interpreted as having formal status. Ar. (Lei.) digitatus (Edwards, 1914) They are merely temporary repositories for the Ar. (Lei.) dolichocephalus (Leicester, species listed below them. As an example, 1908) Aedes (Aedimorphus) vexans Meigen is a valid Ar. (Lei.) flavus (Leicester, 1908) species of uncertain generic and subgeneric Ar. (Lei.) inchoatus Barraud, 1927 placement that was previously placed in subge- Ar. (Lei.) longipalpis (Leicester, 1904) nus Aedimorphus of Aedes. Once the phyloge- Ar. (Lei.) magnus (Theobald, 1908) netic relationships of incertae sedis species have Ar. (Lei.) omissus (Edwards, 1914) been analyzed, they will be retained in their cur- Ar. (Lei.) pectinatus (Edwards, 1914) rent genera, assigned to new genera, or placed Ar. (Lei.) pendulus (Edwards, 1914) in genera resurrected from synonymy with the Ar. (Lei.) traubi Macdonald, 1960 genera in which they were previously assigned. Ar. (Lei.) vimoli Thurman and Thurman, However, we emphasize that until such studies 1958 have been accomplished, incertae sedis species Genus Ayurakitia Thurman, 1954 should continue to be cited with their accus- Ay. griffithi Thurman, 1954 tomed generic and subgeneric affiliations. Stud- Ay. peytoni (Reinert, 1972) ies are currently underway to resolve the tax- Genus Bothaella Reinert, 1973 onomy of the incertae sedis taxa. Bo. eldridgei (Reinert, 1973) Bo. helenae (Reinert, 1973) Abbreviations used for genera (2 letters) and Genus Christophersiomyia Barraud, 1923 subgenera (3 letters) follow Reinert (2001) and Cr. annulirostris (Theobald, 1905) Reinert et al (2004). Cr. ibis (Barraud, 1931) Cr. thomsoni (Theobald, 1905) SPECIES WITH TAXONOMICALLY VALID Genus Diceromyia Theobald, 1911 GENERIC AND SUBGENERIC PLACEMENT Di. franciscoi (Mattingly, 1959) Di. iyengari (Edwards1923) Tribe Aedini Belkin, 1962 Di. pseudonummata (Reinert, 1973) Genus Alanstonea Mattingly, 1960 Di. scanloni (Reinert, 1970) As. treubi (De Meijere, 1910) Di. whartoni (Mattingly, 1965)

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Genus Downsiomyia Vargas, 1950 Lo. amesii (Ludlow, 1903) Do. albolateralis (Theobald, 1908) Lo. fumida (Edwards, 1928) Do. alboniveus (Barraud, 1934) Genus Mucidus Theobald, 1901 Do. ganapathi (Colless, 1958) Mu. laniger (Wiedemann, 1820) Do. harinasutai (Knight, 1978) Mu. quasiferinus (Mattingly, 1961) Do. inermis (Colless, 1958) Genus Neomelaniconion Newstead, 1907 Do. leonis (Colless, 1958) Ne. lineatopenne (Ludlow, 1905) Do. litorea (Colless, 1958) Genus Paraedes Edwards, 1934 Do. mikrokopion (Knight and Harrison, Pr. ostentatio (Leicester, 1908) 1988) Pr. thailandensis (Reinert, 1976) Do. niveoides (Barraud, 1934) Genus Rhinoskusea Edwards, 1929 Do. novonivea (Barraud, 1934) Rh. longirostris (Leicester, 1908) Do. pexa (Colless, 1958) Genus Scutomyia Theobald, 1904 Do. pseudonivea (Theobald, 1905) Sc. albolineata (Theobald, 1904) Do. subnivea (Edwards, 1922) Genus Stegomyia Theobald, 1901 Do. vana (Colless, 1958) St. aegypti (Linnaeus, 1762) Genus Edwardsaedes Belkin, 1962 St. albopicta (Skuse, 1895) Ed. imprimens (Walker, 1860) St. annandalei Theobald, 1910 Genus Finlaya Theobald, 1903 St. craggi Barraud, 1923 Fl. flavipennis Giles, 1904 St. desmotes Giles, 1904 Fl. poicilia Theobald, 1903 St. gardnerii imitator Leicester, 1908 Genus Fredwardsius Reinert, 2000 St. malayensis (Colless, 1962) Fr. vittatus (Bigot, 1861) St. malikuli (Huang, 1973) Genus Heizmannia Ludlow, 1905 St. novalbopicta (Barraud, 1931) Subgenus Heizmannia Ludlow, 1905 St. patriciae (Mattingly, 1954) Hz. (Hez.) aureochaeta (Leicester, 1908) St. perplexa Leicester, 1908 Hz. (Hez.) chengi Lien, 1968 St. pseudalbopicta Borel, 1928 Hz. (Hez.) communis (Leicester, 1908) St. seatoi (Huang, 1969) Hz. (Hez.) complex (Theobald, 1910) St. w-albus Theobald, 1905 Hz. (Hez.) covelli Barraud, 1929 Genus Tanakaius Reinert, Harbach and Kitching, Hz. (Hez.) demeilloni Mattingly,1970 2004 Hz. (Hez.) macdonaldi Mattingly, 1957 Ta. togoi (Theobald, 1907) Hz. (Hez.) mattinglyi Thurman, 1959 Genus Udaya Thurman, 1954 Hz. (Hez.) persimilis Mattingly, 1970 Ud. argyrurus (Edwards, 1934) Hz. (Hez.) propinqua Mattingly, 1970 Genus Verrallina Theobald, 1903 Hz. (Hez.) proxima Mattingly, 1970 Subgenus Harbachius Reinert, 1999 Hz. (Hez.) reidi Mattingly, 1957 Ve. (Har.) indecorabilis Leicester, 1908 Hz. (Hez.) scanloni Mattingly, 1970 Ve. (Har.) yusafi (Barraud, 1931) Hz. (Hez.) scintillans Ludlow, 1905 Subgenus Neomacleaya Theobald, 1907 Hz. (Hez.) taiwanensis Lien, 1968 Ve. (Nma.) adusta (Laffoon, 1946) Hz. (Hez.) thelmae Mattingly, 1970 Ve. (Nma.) andamanensis (Edwards, Subgenus Mattinglyia Lien, 1968 1922) Hz. (Mat.) achaetae (Leicester, 1908) Ve. (Nma.) atria (Barraud, 1928) Hz. (Mat.) catesi (Lien, 1968) Ve. (Nma.) clavata (Barraud, 1931) Genus Isoaedes Reinert, 1979 Ve. (Nma.) cretata (Delfinado, 1967) Ia. cavaticus (Reinert, 1979) Ve. (Nma.) cyrtolabis (Edwards, 1928) Genus Kenknightia Reinert, 1990 Ve. (Nma.) gibbosa (Delfinado, 1967) Ke. dissimilis (Leicester, 1908) Ve. (Nma.) hispida (Delfinado, 1967) Ke. harbachi (Reinert, 1990) Ve. (Nma.) incerta (Edwards, 1922) Genus Lorrainea Belkin, 1962 Ve. (Nma.) latipennis (Delfinado, 1967)

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Ve. (Nma.) notabilis (Delfinado, 1967) Oc. (Fin.) elsiae (Barraud, 1923) Ve. (Nma.) phnoma (Klein, 1973) Oc. (Fin.) feegradei (Barraud, 1934) Ve. (Nma.) protuberans (Delfinado, 1967) Oc. (Fin.) formosensis (Yamada, 1921) Ve. (Nma.) pseudodiurna (Theobald, Oc. (Fin.) harveyi (Barraud, 1923) 1910) Oc. (Fin.) jugraensis (Leicester, 1908) Ve. (Nma.) sohni (Reinert, 1974) Oc. (Fin.) khazani (Edwards, 1922) Ve. (Nma.) torosa (Delfinado, 1967) Oc. (Fin.) lophoventralis (Theobald, 1910) Ve. (Nma.) unca (Theobald, 1901) Oc. (Fin.) macfarlanei Edwards, 1914 Ve. (Nma.) vallistris (Barraud, 1928) Oc. (Fin.) prominens (Barraud, 1923) Subgenus Verrallina Theobald, 1903 Oc. (Fin.) pseudotaeniatus (Giles, 1901) Ve. (Ver.) butleri (Theobald, 1901) Oc. (Fin.) pulchriventer (Giles, 1901) Ve. (Ver.) dux (Dyar and Shannon, 1925) Oc. (Fin.) reinerti (Rattanarithikul and Ve. (Ver.) lugubris (Barraud, 1928) Harrison, 1988) Genus Zeugnomyia Leicester, 1908 Oc. (Fin.) saxicola Edwards, 1922 Ze. gracilis Leicester, 1908 Oc. (Fin.) shortii (Barraud, 1923) Oc. (Fin.) simlensis (Edwards, 1922) INCERTAE SEDIS SPECIES WITH Oc. (Fin.) unicinctus (Edwards, 1922) PROVISIONAL GENERIC AND Subgenus Ochlerotatus Lynch Arribalzaga, SUBGENERIC PLACEMENT 1891, sensu auctorum Oc. (Och.) vigilax (Skuse, 1889) Genus Aedes Meigen, 1818, sensu auctorum Subgenus Aedimorphus Theobald, 1903, DISCUSSION sensu auctorum Harrison et al (1990) compiled a list of mos- Ae. (Adm.) alboscutellatus (Theobald, quitoes occurring in Thailand which included 1905) several new country records and the elimination Ae. (Adm.) caecus (Theobald, 1901) of incorrect records. They noted that the inclu- Ae. (Adm.) culicinus Edwards, 1922 sion of Hz. (Hez.) taiwanensis needed confirma- Ae. (Adm.) mediolineatus (Theobald, tion. Mattingly (1970) indicated that he was un- 1901) sure of records of Hz. (Hez.) indica (Theobald) Ae. (Adm.) orbitae Edwards, 1922 from Thailand. Zeugnomyia gracilis was reported Ae. (Adm.) pallidostriatus (Theobald, for the first time from southern Thailand by 1907) Benjaphong and Rattanarithikul (1991). Ae. (Adm.) pampangensis (Ludlow, 1905) Armigeres (Lei.) cingulatus is a valid species (see Ae. (Adm.) pipersalatus (Giles, 1902) Thurman, 1959) and not a synonym of Ar. (Lei.) Ae. (Adm.) vexans vexans (Meigen, 1830) longipalpis as indicated in Stone and Knight Subgenus Cancraedes Edwards, 1929, (1977). Thurman (1959, pages 97 and 100) sensu auctorum stated that only records of Ar. cingulatus by Ae. (Can.) indonesiae Mattingly, 1958 Barraud (1927), Borel (1930), and Edwards (in Ae. (Can.) kohkutensis Mattingly, 1958 Barraud,1934) were synonymous with Ar. Genus Ochlerotatus Lynch Arribalzaga, 1891, longipalpus. sensu auctorum Subgenus Bruceharrisonius Reinert, 2003, New country records for Thailand reported sensu auctorum here include the following four species of Oc. (Brh.) christophersi (Edwards, 1922) Armigeres that were examined by one of us (RR): Oc. (Brh.) greenii (Theobald, 1903) Ar. confusus, Ar. foliatus, Ar. pendulus and Ar. Subgenus Finlaya Theobald, 1903, sensu traubi. auctorum Oc. (Fin.) albotaeniatus (Leicester, 1904) ACKNOWLEDGEMENTS Oc. (Fin.) assamensis (Theobald, 1908) Appreciation is expressed to Kenneth J Oc. (Fin.) chrysolineatus (Theobald, 1907) Linthicum (CMAVE) for providing facilities to JFR,

Vol 36 No. 2 March 2005 415 SOUTHEAST ASIAN J TROP MED PUBLIC HEALTH and to Bruce A Harrison (Public Health Pest Reinert JF. Restoration of Verrallina to generic rank in Management, North Carolina Department of tribe Aedini (Diptera: Culicidae) and descriptions Environment and Natural Resources, Winston- of the genus and three included subgenera. Salem, NC ), Graham B White (Department of Contrib Am Entomol Inst (Gainesville) 1999; 31: Entomology, University of Florida, Gainesville, 1-83. FL), and Leopoldo M Rueda (WRBU), for review- Reinert JF. Restoration of Ayurakitia to generic rank in ing the manuscript. tribe Aedini and a revised definition of the genus. J Am Mosq Control Assoc 2000a; 16: 57-65. REFERENCES Reinert JF. New classification for the composite genus Aedes (Diptera: Culicidae: Aedini), elevation of Barraud PJ. A revision of the culicine mosquitoes of subgenus Ochlerotatus to generic rank, reclas- India. Part XX. The Indian species of Armigeres sification of the other subgenera, and notes on (including Leicesteria), with descriptions of two certain subgenera and species. J Am Mosq Con- new species. Indian J Med Res 1927; 14: 533- trol Assoc 2000b; 16: 175-88. 48. Reinert JF. Revised list of abbreviations for genera and Barraud PJ. The Fauna of British India, Including subgenera of Culicidae (Diptera) and notes on Ceylon and Burma. Diptera, Vol V. Family Culi- generic and subgeneric changes. J Am Mosq cidae. Tribes Megarhinini and Culicini. London: Control Assoc 2001; 17: 51-5. Taylor and Francis, 1934. Reinert JF. Description of Bruceharrisonius, a new Benjaphong N, Rattanarithikul R. Zeugnomyia gracilis subgenus of Ochlerotatus, and a redescription of Leicester (Diptera: Culicidae), a new genus and its type species Oc. (Brh.) greenii. J Am Mosq species occurrence record from Thailand. Mosq Control Assoc 2003a; 19: 309-22. Syst 1991; 23:53. Borel E. Les moustiques de la Cochinchine et du Sud- Reinert JF. Transfer of Heizmannia thelmae from sub- Annam. Monograph III, Collection de la Societe genus Mattinglyia to subgenus Heizmannia of de Pathologie Exotique, Paris, France, 1930. genus Heizmannia and new characters for sepa- rating the two subgenera. J Am Mosq Control Harrison BA, Rattanarithikul R, Peyton EL, Mong- Assoc 2003b; 19: 433-4. kolpanya K. Taxonomic changes, revised occur- rence records and notes on the Culicidae of Thai- Reinert JF, Harbach RE, Kitching IJ. Phylogeny and land and neighboring countries. Mosq Syst 1990; classification of Aedini ( Diptera: Culicidae) based 22: 196-227. on morphological characters of all life stages. International Commission on Zoological Nomenclature. Zool J Linn Soc 2004; 142:289-368. International Code of Zoological Nomenclature. Stone A, Knight KL. A catalog of the mosquitoes of the 4th ed. London: International Trust for Zoological world (Diptera: Culicidae). College Park, MD: The Nomenclature, 1999. Thomas Say Foundation, 1977. Mattingly PF. Contributions to the mosquito fauna of Thurman EHB. A Contribution to a Revision of the Southeast Asia. - VI. The genus Heizmannia Culicidae of Northern Thailand. Bulletin A-100, Ludlow in Southeast Asia. Contrib Am Entomol University of Maryland Agricultural Experiment Inst (Ann Arbor) 1970; 5: 1-104. Station, College Park, MD, 1959.

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