Chec List Faunal Richness and the Checklist of Indian Mosquitoes

S (Diptera: Culicidae)

of Faunal richness and the checklist of Indian mosquitoes 1 2 2 1 2 1

ists D. R. Bhattacharyya , A. R. Rajavel , R. Natarajan , P. K. Mohapatra , P. Jambulingam , J. Mahanta L and A. Prakash 1, 3*

1 Regional Medical Research Centre, NE (Indian Council of Medical Research), Post Box No. 105, Dibrugarh – 786 001, Assam, India.

3 National Institute for Research in Environmental Health (Indian Council of Medical Research), Kamla Nehru Hospital Building, Bhopal – 462 001, 2 MadhyaVector Control Pradesh, Research India. Centre (Indian Council of Medical Research), Indiranagar, Puducherry – 605 006, Tamilnadu, India. * Corresponding Author. E-mail: [email protected]

Abstract: A review of published studies revealed that the Indian mosquito fauna comprises 393 species in

number49 genera of and species 41 subgenera. (176 species The in subfamily 33 genera Anophelinae and two groups contains of incertae 61 species sedis in; onei.e., “genusAedes ”followed sensu auctorum by Culicinae and with“Ochlerotatus 332 species” sensu in auctorum 11 tribes). andWith 48 the genera. recent taxonomic The tribe changes Aedini (subfamily in tribe Aedini, Culicinae) the Indian contains mosquito the highestgenera

Aedini are discussed. A total of 31 species are currently recognized in India for transmitting various mosquito- bornehave gone agents up offrom human 22 todiseases. 49. Changes A checklist to the for Indian the IndianAedini mosquitofauna subsequent species is to presented the reclassification and the need of tribefor a comprehensive study is emphasized.

10.15560/10.6.1342

DOI:

Introduction numbers, since the 1980s some faunistic surveys carried Mosquitoes, belonging to family Culicidae and order out in various parts of India detected many species which Diptera, are a large group of insects present throughout are new additions to the mosquito fauna of India. It seems the temperate and tropical regions and even beyond the that some of these records did not reach the mosquito catalog. As such, an up-to-date check list of mosquitoes of Region, which includes India, is regarded as one of the India is still not available, and the actual number of species richestArctic Circlebiogeographic of the world regions (Harbach for mosquitoes 2007). Theof the Oriental world, present in India, has to be approximated. Moreover, with along with the Neotropics (Gaston and Hudson 1994). the advancement of molecular biology, many researchers At present, a total of 3,540 recognized mosquito species, from non-entomology backgrounds are now working with divided between two subfamilies and 112 genera, are mosquitoes, and there is confusion about the informal taxonomic series, groups and subgroups to which mosquito fold increase of the present numbers can be foreseen with species belong. Confusion seems to have increased after morerecorded discoveries in the world and naming (Harbach of 2014).sibling Aspecies three tomainly five- in Anopheles, as a result of application of DNA-based (Reinert et al. 2004, 2006, 2008, 2009) as journals are the changes made to the classification of tribe Aedini mosquito biodiversity after Brazil, Indonesia, Malaysia and tribe Aedini brought major changes raising the number methods (Harbachet al2007).. 2007). India After is theranked monumental fifth in terms works of ofusing genera two insystems this tribe of classification. from 11 to 81 Reclassification along with changes of the of Christophers (1933) and Barraud (1934), not many to the spelling of names of some species in accordance Thailandcomprehensive (Foley biosystematics studies of Indian Culicidae with provisions of the International Code of Zoological have been undertaken. Since that time, many changes have Nomenclature. Hence, a checklist seems essential for taken place in the mosquito taxonomic literature. However, the entomologists and public health personnel working in the absence of any up-to-date monograph on Indian Culicidae, entomologists in India are still compelled to use familiarity with the new names. Considering this, an up- these two books. In this context, as mentioned by Reuben to-datein India check to reflect list of changesthe mosquito to nomenclature species of India to based promote on et al. (1993), it needs to be emphasized that publications of the Southeast Asian Mosquito Project are very useful the mosquito fauna of India and need for future work are alsothe current discussed. classification is provided. The present status of (1972) rightly proposed for a systematic study of Indian Culicidaefor the identification because of increasing of Indian mosquito-borne mosquito species. diseases, Berlin Materials and Methods changing ecology and advances in mosquito systematics. Information was collected from two online resource databases, namely systematic catalog of Culicidae and et al. 2014). Although fewer in mosquito taxonomic inventory and published materials The online systematic catalog of Culicidae listed 356 species in India (Gaffigan 1342 Bhattacharyya et al. | Mosquito biodiversity of India mainly since the 1980s (Bhattacharyya et al. 2000, 2002, prevalent and An. nigerrimus as rather uncommon species 2003, 2004a, 2004b, 2005, 2007, 2008, 2009; Khan et al. (Khan et al et al. 1987). Immature surveys 1998; Rajavel and Natarajan 2006, 2011; Rajavel et al. carried out in upper Brahmaputra valley detected An. 1998, 2004, 2005a, 2005b, 2005c, 2005d, 2011; Rajput crawfordi, An.. 1998; paraliae Tewari, An. peditaeniatus and An. sinensis and Singh 1987a, 1987b, 1987c, 1989, 1990; Reuben et of Hyrcanus Group from the north-east India, but did not al et al collect any specimen of An. nigerrimus (Khan et al. 1998). An. paraliae in these surveys information. 1993; Tewari on country . 1987;occurrence Tewari records and Hiriyan of mosquito 1991, were collected as immatures from a forest fringe area of 1992, 1995). The systematic catalog of Culicidae provides Assam.A few specimens Earlier considered identified as as a subspecies of An. lesteri, An. paraliae was later elevated to species status by Harrison et species. The mosquito Anopheles taxonomic, Culex inventory, Verrallina contains and al. (1990), but more recently it has been synonymyzed with up-to-dateUranotaenia information are available on worldin the Culicidae. website of The mosquito internal lesteri et al. (2013), hence it is listed as lesteri here. classificationtaxonomic inventory, of genera which is followed here for the However, distribution of this species is restricted to low Indian mosquito fauna belonging to these genera. Recent elevation by Taaicoastal areas of Malaysia, Brunei, Vietnam and abbreviations of generic and subgeneric names provided An. pursati. Hence, record of this generic and species names is provided in the check list speciesThailand in (Harrison India needs and further Scanlon investigation, 1975). This speciesincorporating is also (Appendixby Reinert 1).(2009a) are followed here. The authorship of easily misidentified with of r-DNA was used to differentiate member species of An. Results and Discussion hyrcanusDNA-based group analysis (Ma similaret al. 1998, to China, 2000a, where, 2000b). ITS2 Recently, marker Records indicate that the Indian mosquito fauna includes 393 species divided among 49 genera and 41 subgenera. An. crawfordi, An. peditaeniatus Subfamily Anophelinae contains 61 species in one genus ITS2and An. sequencing sinensis in of the some north-east specimens India of (Regional Hyrcanus Medical Group followed by subfamily Culicinae with 332 species in 11 confirmedResearch Centre, the presence Dibrugarh, of Assam, India; unpublished data). Similarly Barbirostris Group which includes 11 contains the highest number of species (176 species in species globally also has uncertain distribution records 33tribes genera and 48and genera. two groups Tribe Aediniof incertae of subfamily sedis, i.e., Culicinae “Aedes” for most of the species, particularly in Indonesia and sensu auctorum, “Ochlerotatus” sensu auctorum

Appendix 1. In India, 31 species are currently recognized) (Table wereIndian recognized sub-region, in this mainly group from because India. of An. identificationhodgkini was for1). transmittingThe check list various of the mosquito-borne Indian Culicidae pathogens; is presented these as detecteddifficulties from (Harrison Assam during and Scanlon larval collections 1975). Four (Khan species et al. 1998). Subfamily Anophelinae: Subfamily Anophelinae has Subgenus Cellia containing 35 species encompasses all are listed in Table 2. important vectors of human malaria in India. Recently, two Anopheles, with 61 formally named species divided between new species have been described from this subgenus from subgenerathree genera Anopheles globally. Indianand Cellia species. However, are confined the number to genus of India. An. pseudosundaicus belonging to Pyretophorus species will increase with the naming or recognition of synonymous names of sibling species in several groups et al. 2009). Another species belonging to Maculatus or complexes. In India, subgenus Anopheles contains 26 GroupSeries wasof the described Neocellia from Series coastal named areas as An.of Kerelakrishnai (Tyagi was species. In this subgenus, the assemblage of species in described (Sathe and Jagtap 2012). However, Harbach two most important groups (Hyrcanus and Barbirostris (2014) noted the lack of credible evidence to support the recognition of this proposed species. Hence, krishnai a highly complex group, which includes 30 closely related is regarded as nomen nudum and is not included in our Groups) is still uncertain in India. The Hyrcanus Group is list of Indian Anopheles. Anopheles dthali and Anopheles regions, with some species playing important role in superpictus have been known only from extreme north- species distributed widely in the Oriental and Palaearctic also not been recorded subsequently in India and therefore transmission of malaria and filariasis (Ma and Xu 2005). notwest included (Baluchistan) in the presentwhich is checklist. not part of India. They have sevenThis Group species was of also this incriminatedGroup were recordedas a vector from of Japanese India in Several anopheline taxa recognized earlier to be comparisonencephalitis to in 25 India species and recorded neighboring in China countries. (Ma et Only al. medically important are now found to be complexes or groups of morphologically indistinguishable species (Rattanarithikul et al. 2006). Some important complexes impossible1998, 2000a, unless 2000b). accompanied Morphological by associated identification immature of and groups recorded in India are, An. annularis (species skinsthese species(Harrison in adultand Scanlonstage seems 1975). to beMany very anophelinedifficult or A and B), An. culicifacies (species A, B, C, D and E), An. surveys carried out in India recorded An. nigerrimus of dirus An. fluviatilis An. Hyrcanus Group to be the most predominant, seemingly subpictus (species A, B, C, and D) and the An. sundaicus (cytotype (species D) complexes X), (Walton (species et al S, T, U and V), (Malhotra et al. 1987; Nagpal and Sharma 1987). However, With the advances in mosquito taxonomy several species surveysbecause carried their identification out in north-east was based India on and adult Western stage belonging to such groups or complexes. 1999; are now WHO formally 2007). named. An. maculatus, recognized as one such species done using adult as well as associated larval and pupal group, includes nine formally named species (Harbach characters,Ghats of South recorded India, An. where peditaeniatus species identification to be the most was 2014; Somboon et al. 2011). A recent study revealed the

1343 Bhattacharyya et al. | Mosquito biodiversity of India

Table 1 Member species of Culicidae in India. NUMBER OF SUBFAMILY TRIBE GENUS SUBGENUS SPECIES Anophelinae Anopheles Anopheles 26 Cellia 35 Culicinae Aedeomyiini Aedeomyia Aedeomyia 1 Aedini “Aedes” sensu auctorum 4 Aedimorphus 15 Armigeres Armigeres 9 Leicesteria 11 Ayurakitia 1 Bruceharrisonius 4 Christophersiomyia 4 Cancraedes 2 Collessius Collessius 3 Alloeomyia 1 Danielsia 2 Dendroskusea 5 Downsiomyia 6 Edwardsaedes 1 Finlaya 2 Fredwardsius 1 Gilesius 1 Himalaius 2 Heizmannia Heizmannia 11 Mattinglyia 2 Hopkinsius Yamada 1 Hulecoeteomyia 5 Jihlienius 1 Kenknightia 2 Lorrainea 2 Mucidus Mucidus 3 Neomelaniconion 1 Ochlerotatus 1 “Ochlerotatus” sensu auctorum “Finlaya” sensu auctorum 5 Paraedes 4 Petermattinglyius Petermattinglyius 2 Phagomyia 11 Rhinoskusea 3 Scutomyia 1 Stegomyia Actinothrix 2 Heteraspidion 2 Huangmyia 2 Stegomyia 1 Xyele 1 Without subgenus 11 Tewarius 3 Udaya 2 Verrallina Harbachius 4 Neomacleaya 16 Verrallina 3 Culicini Culex Culex 23 Culiciomyia 9 Eumelanomyia 10 Lophoceraomyia 28 Maillotia 1 Oculeomyia 6 Lutzia Metalutzia 4 Culisetini Culiseta Allotheobaldia 1 Culiseta 2 Ficalbia 1 Mimomyia Etorleptiomyia 1 Ficalbiini Mimomyia 4 Ingramia 1 Hodgesiini Hodgesia 1 Mansoniini Coquilletidia Coquilletidia 3 Mansonia Mansonioides 4 Orthopodomyia 5 Sebethini Malaya 2 Orthopodomyiini Topomyia Suaymyia 1 Topomyia 3 Tripteroides Rachionotomyia 5 Tripteroides 3 Toxorhynchites Toxorhynchites 9 Uranotaenia Pseudoficalbia 15 Toxorhynchitini Uranotaenia 13 Uranotaeniini 2 11 49 + 2 sensu auctorum 41 + 1 sensu auctorum 393 Total of each taxnomomic level

1344 Bhattacharyya et al. | Mosquito biodiversity of India presence of six member species of the Maculatus Group (Bhattacharyya et al. 2000). Recently a new species Ar. in the north-east India, with detection of An. rampae mahantai collected as immatures from endemic pitcher constituting a new country record (Singh et al. 2012). Such plant Nepenthes khasiana of Meghalaya state in the north- studies may be extended to other areas where Maculatus east India was described (Bhattacharyya et al. 2009). Ar. pallithorax, described from Yunnan, China was detected reported earlier from Yunnan province of China, was also from Namsai, Arunachal Pradesh (Rajavel et al. 2011). Group is present. Similarly, species X of the Dirus Complex, Genus Ayurakitia (Prakash et al. 2010). two species detected from mountainous areas of western detectedSubfamily in Haflong Culicinae: area Subfamily of Assam Culicinaein the north-east is comprised India : This genus is represented by only of 332 species divided into 11 tribes and 48 genera in India. collectedThailand form (Reinert Pandanus 1972). axils Recently, in Meghalaya, occurrence north-east of this Aedeomyia is Indiagenus andwas rearedreported to foradults, the firstwere time found from to India be Ayurakitia as larvae peytoni (Rajavel and Natarajan 2011). mosquitoTribe Aedeomyiini:catalog under A the single Indian genus mosquito fauna. Genus Bruceharrisonius: An earlier subgenus of genus included in this tribe. This genus is not listed in the Ochlerotatus (Reinert 2003) Bruceharrisonius was later one of the distribution localities for Aedeomyia catastica. elevated to generic rank (Reinert et al However, Tyson (1970) reported Andaman Islands as Assam and Western Ghats (Khan et al. 1998; Reuben et al. Aureostriatus Subgroup (Aureostriatus. 2006). Group) This and genus one 1993)This species and in was the also mangrove reported forests from ofmainland Sunderbans, India Westfrom speciesincludes of four Auronitens species Subgroup in India. (Alboannulatus Three species earlierGroup) inof Bengal (Rajavel et al. 2005a). subgenus Finlaya genus Aedes (Knight and Marks 1952) are now placed in genus Bruceharrisonius. Aedini contained 11 genera (Knight and Stone 1977 and Genus Christophersiomyia: A former subgenus of genus Tribe Aedini: In the earlier classification the tribe Aedes, now elevated to generic rank (Reinert et al. 2004), has raised the number of genera to 81 (Reinert et al. Christophersiomyia 2004,its supplements). 2006, 2008, Recent2009). Reinert reclassification (1999, 2000a) of this earlier tribe occur in India. Cr. gombakensis, resurrected Verrallina and Ayurakitia to generic rank. from Western Ghats, includes is a new five country species, record of which,(Reuben four et Revision of tribe Aedini resulted in the elevation of many al. 1993). reported for the first time former subgenera of Aedes to generic status (Reinert et Genus Cancraedes: Cancraedes was elevated to genus al. 2004). Downsiomyia was elevated to genus level from from former subgenus of Aedes (Reinert et al synonymy with Finlaya of Finlaya and allied taxa and Ochlerotatus and allied taxa Region, only two species occur in India. . 2009). Of (Reinert et al. 2006, 2008),. Further followed examination by comprehensive and revision the Genustotal ten Collessius species: Aincluded newly recognizedin this genus genus from Collessius Oriental phylogenetic study of tribe Aedini (Reinert et al. 2009) (Reinert et al. 2006) includes species formerly in resulted in recognition of more genera and rearrangement Pseudotaeniatus Subgroup of Mediovittatus Group, of taxa. subgenus Finlaya of genus Aedes (Knight and Marks 1952). In India this genus is represented by four species divided tribe Aedini includes 176 species in 33 genera and two into two subgenera. groupsAs perof incertae the current sedis species classification, (“Aedes” Indiansensu auctorum- fauna of Genus Danielsia: Danielsia was elevated to generic 4 and “Ochlerotatus” sensu auctorum- 5). Indian species rank from synonymy with Finlaya and included three taxa belonging to subgenus Finlaya genus Aedes are now placed (Reinert 2009b). In India this genus is comprised of two in 12 genera (Bruceharrisonius, Collessius, Downsiomyia, species formerly in Albotaeniatus Subgroup of subgenus Danielsia, Finlaya, Gilesius, Hopkinsius, Hulecoeteomyia, Finlaya, genus Aedes. Himalaius, Jihlienius, Ochlerotatus and Phagomyia). Genus Dendroskusea Similarly species of the subgenus Diceromyia of India are Dendroskusea now placed in three genera Dendroskusea, Petermattinglyius earlier placed in subgenus: All Diceromyia five species of included genus Aedes in genus. and Tewarius. Illustrated keys of these new genera present Genus Downsiomyia are found: Downsiomyia in India. These as a species genus werewas et al. (2010). resurrected from synonymy with Finlaya (Reinert et al taxonomicin Thailand wereinventory. provided Kaur by Rattanarithikul(2003) provided updated represented by six species in India which were earlier distributionFurther details maps are of availableIndian Aedini in the fauna. website of mosquito placed. 2006; in the Reinert Niveus and Subgroup Harbach of 2006).subgenus This Finlaya. genus is Genus Aedimorphus: Earlier recognized as a subgenus species Do. nivea (Ae. niveus) has been incriminated as a of genus Aedes, Aedimorphus is now elevated to generic vector of diurnally subperiodic Wuchereria bancrofti in The the rank (Reinert et al et al of the 67 species recognized in this genus, 15 species were (1995) redescribed this species from Andaman and recorded from India.. 2009). Mostly Oriental in distribution, Nicobar Islands.islands (Tewari . 1995). Tewari and Hiriyan Genus Armigeres Genus Edwardsaedes: Comprising of three species, the region and also in the Palaearctic and Australasian regions, genus Edwardsaedes is represented by only one species in the genus Armigeres: Distributedis represented mainly by in20 the species Oriental in India. India, between two subgenera Armigeres and Leicesteria. Genus Finlaya Finlaya was Rajput and Singh (1987c) reported the detection of one of the largest subgenus of genus Aedes comprising of Ar. dolichocephalus from Manipur state. Ar. joloensis, a : In the earlier classification rare mosquito species, was detected from upper Assam the genus Finlaya includes species only from Kochi Group 42 species in India. However, in the current classification, 1345 Bhattacharyya et al. | Mosquito biodiversity of India of subgenus Finlaya (Reinert et al. 2004, 2006; Reinert distribution of Pr. chrysoscuta, but pointed that Mattingly Finlaya are found in India. Fl. flavipennis is available otherwise, we retain Barraud’s record as Pr. mangroveand Harbach forest 2005). ecosystem Two species of Andamanof and Nicobar chrysoscuta(1958) questioned and include the records it in the from present India. checklist. Until evidence Islands (Rajavel and Natarajanwas reported 2006). for the first time from Genus Petermattinglyius: Petermattinglyius comprises Genus Fredwardsius: Fredwardsius vittatus is the only species included in this genus, which is present in India. placed in subgenus Diceromyia. Pe. franciscoi was reported Genus Gilesius of two species in India. These two species were earlier genus, only one species occurs in India. (Rajavel et al. 2005c). Genus Heizmannia: Of the two species included in this for theGenus first Phagomyia time from: mangrove Phagomyia forest was ecosystemelevated to of generic Orissa species in India, 11 species under subgenus Heizmannia status from synonymy with Finlaya and includes species and two species under: This subgenus genus is Mattinglyia represented. Rajput by 13 formerly in Gubernatoris Subgroup of subgenus Finlaya Hz. (Reinert et al. 2006). Phagomyia includes 11 species in aureochaeta in India from Manipur state. Similarly Hz. chengiand Singh (1987b) first reported the occurrence of Table 2. Vectors of mosquito-borne diseases in India. . 2005b). et al Malaria was reported for the first time from Jeypore hill Genus Himalaius Anopheles (Cellia) culicifacies s.l. aretracts found of Orissa in India. (Rajavel Anopheles (Cellia) baimaii Genus Hopkinsius: :Two Hopkinsius species contains representing two subgenera, this genus Anopheles (Cellia) fluviatilis s.l. Hopkinsius and Yamada Anopheles (Cellia) minimus s.l. subgenus Yamada occurs in India. Anopheles (Cellia) stephensi Genus Hulecoeteomyia. : OnlyReinert one et al species. (2006) belonging resurrected to Anopheles (Cellia) sundaicus s.l. Hulecoeteomyia to generic status from synonymy with Anopheles (Cellia) annularis s.l. Finlaya Anopheles (Cellia) jeyporiensis earlier placed in Chrysolineatus Subgroup of subgenus Anopheles (Cellia) philippinensis Finlaya. Five species included in this genus from India were Anopheles (Cellia) nivipes Anopheles (Cellia) varuna Genus Jihlienius Anopheles (Cellia) maculatus s.l. genus, only one is found in India. 2010; Rao 1984) : Of the three species recognized in this et al. Genus Kenknightia: Kenknightia comprised of 12 Japanese encephalitis (WHO 2007; Bhattacharyya species of which only two species represent this genus in Culex (Culex) vishnui India. Culex (Culex) pseudovishnui Genus Lorrainea: Presence of Lo. amesii and Lo. Culex (Culex) tritaeniorhynchus fumidus belonging to genus Lorrainea Culex (Culex) fuscocephala from India based on collections made in mangrove forests Culex (Culex) quinquefasciatus of Sundarbans, in West Bengal (Rajavel were et first al . reported2005a), Culex (Culex) gelidus et al. 2005c) and Andaman Culex (Culex) whitmorei Nicobar Islands (Rajavel et al. 2004). Previously, species of Culex (Oculeomyia) bitaeniorhynchus Bhitarkanika in Orissa (Rajavel Culex (Oculeomyia) infula Solomon Islands. Culex (Oculeomyia) epidesmus Anopheles (Anopheles) barbirostris s.l. thisGenus genus Mucidus were reported: Reinert from(2000b) Thailand, earlier Philippinestreated Mucidus and Anopheles (Anopheles) peditaeniatus as a subgenus of newly elevated genus . Ochlerotatus Anopheles (Cellia) subpictus s.l. Mucidus was elevated Mansonia (Mansonioides) annulifera to generic rank (Reinert et al Mansonia (Mansonioides) indiana MucidusHowever, in the later classification Mansonia (Mansonioides) uniformis of Mu. laniger was reported from. two 2004). forest Three areas species of Assam, of (Kanojia 2007) north-east are India known (Bhattacharyya to exist in India. et al Confirmed. 2005). occurrence Dengue Genus Neomelaniconion: Mainly distributed in Stegomyia (Stegomyia) aegypti Afrotropical region, only one species Neomelaniconion Stegomyia albopicta lineatopenne (Kaul et al. 1998; Das et al. 2004) including India. Chikungunya Genus Ochlerotatus represent this genus in the Oriental region Stegomyia (Stegomyia) aegypti species in India. Stegomyia albopicta (Mourya et al. 2001) Genus Paraedes: Subgenus: This genusof the genus includes Aedes only in onethe West Nile Paraedes has been raised to genus Culex (Culex) vishnui level (Reinert et al. 2004) and is represented by four Culex (Culex) quinquefasciatus earlier classification, (Paramasivan et al. 2003) of Pr. ostentatio was reported from a tropical forest Filariasis zonespecies of inDibrugarh India. In district, this genus Assam the (Bhattacharyya confirmed presence et al. Culex (Culex) quinquefasciatus 2002). Barraud (1934) earlier reported this species from Mansonia (Mansonioides) annulifera Malabar Coast of Kerela on the basis of female specimens. Mansonia (Mansonioides) uniformis Reinert (1981) on resurrection of Pr. chrysoscuta from Downsiomyia nivea Pr. ostentatio considered Barraud’s Indian record under et al. 1995) (Agarwal and Sashindran 2006; Tewari 1346 Bhattacharyya et al. | Mosquito biodiversity of India

India. Ph. feegradei was new addition to the mosquito fauna of India from the mangrove forest ecosystem of species of undetermined subgenus (Aedes kolhapuriensis, et al. 2005c). Aedesare now sangitee kept , asAedes provided panchgangee by the authors.and Aedes Four sangiti new) Genus Rhinoskusea: Rhinoskusea comprises of three were described from Kolhapur district of Maharashtra speciesOrissa (Rajavel in India. In this genus one new species Rh. (Girhe and Sathe 2001; Sathe and Girhe 2001). However, portonovoensis was described from mangrove forest of descriptions of these species were inadequate and mainly based on adult female characters. Comparisons Rh. wardi from mangrove forests of Andaman and Nicobar with closely related species were not available in the Islandsthe east iscoast a new (Tewari country and Hiriyanrecord (Rajavel1991). The and detection Natarajan of descriptions. Hence, collections of all life stages may be 2006). Reinert (2000b) earlier treated Rhinoskusea as required for comparing with other related species to a subgenus of genus Ochlerotatus. However, in the later establish their validity and placing them in relevant genera Rhinoskusea was elevated to generic status and subgenera. Genus Aedes is now restricted mainly in (Reinert et al. 2004). the Palaearctic and Nearctic Regions. We retained these classificationGenus Scutomyia : Scutomyia includes only one species species in the checklist as they were already in the list in India. of mosquito catalog and mosquito taxonomic inventory. Genus Stegomyia Ochlerotatus” Stegomyia includes 19 species in India. In this genus St. and subgenus “Finlaya” (Oc. auronitens, Oc. oreophilus, Oc. krombeini, earlier placed: The in medicallyScutellaris important Group of genus Similarly,sintoni, Oc. fivesuffusus species and retainedOc. versicolor in genus) are yet “ to be placed Aedes and subgenus Stegomyia (Huang 1979), was et al. 1987) and later from north-east India (Bhattacharyya et al. 2008). generain relevant and generaseven subgenera in the current in India. classification. Huangdetected (1975) first fromreported south this India species (Tewari to be very common in GenusTribe Culicini:Culex: Genus Culicini Culex includes is represented 81 species by 77 species in two Sri Lanka, which probably remained undetected as it was in six subgenera in India with Lophoceraomyia being mistaken for another common species, St. albopicta. Hence the predominant subgenus with 28 species, followed careful observations during entomological surveys are by subgenus Culex necessary to differentiate St. krombeini from St. albopicta, many important vectors of Japanese encephalitis (JE) with 23 species. This genusCulex , contains Vishnui of St. flavopicta in Assam, western Himalayas and Coorg Subgroup contains three most important vectors (Cx. (Barraudespecially 1934) in rural was and not forest considered areas byof HuangIndia. The(1972) records who tritaeniorhynchus,virus and microfilariae Cx. vishnuiin India. andIn subgenus Cx. pseudovishnui ) of restricted its distribution to Japan and Korea. Its inclusion JE in India. Presence of another three members in this by Kaur (2003) in Indian species is based on Barraud subgroup i.e. Cx. alienus, Cx. perplexus and Cx. whitei, which (1934). While St. pseudoalbopictus, St. subalbopictus and St. novalbopictus have been recorded in collections made the basis of larval, pupal and male phallosome characters in several parts of the country, St. flavopicta has not (Sirivanakarncan only be identified 1976), distinctivelycan create diagnostic from other problem members in onJE been obtained and hence it is not included in the present vector surveillance in areas of their occurrence. Barraud checklist. At present of the eight new subgenera recognized (1934) earlier collected larvae of Cx. whitei Cx. alienus which is regarded subgenera. Remaining 11 species under this genus are still as one of the uncommon members by Sirivanakarn from Haflong (1976) withoutfor this genus,subgeneric only eightplacement. Indian species are placed in five withinarea of Assam.this subgroup Few larvae were of collected from Assam. Cx. Genus Tewarius perplexus earlier known only from Andaman Islands was genus, three are found in India. Reinert (2006) placed et Te. agastyai, Te. reubenae: Of the and four Te. species nummatus included of subgenus in this al. 2005c). Diceromyia to a new genus Tewarius. Te. agastyai and Te. alsoIn reported subgenus from Culiciomyia, mangrove forests of Orissa (Rajavel reubenae reported Cx. harrisoni from Senapati district of Manipur. Hiriyan 1992). Cx. spathifurca Rajput and Singh (1987a) first Genus wereUdaya described: Udaya includesfrom Western three Ghats species (Tewari from andthe mosquito faunistic study in a mangrove forest ecosystem was reportedet al. for1998). the Cx. first scanloni time during known a Genus Verrallina: Divided into three subgenera this genusOriental contains region, 23 of specieswhich two in India. are found Subgenus in India. Neomacleaya wasof Tamil detected Nadu from (Rajavel Nagarhole National Park, Karnataka contains most of the species (16) followed by Harbachius (Rajavelearlier from et al . Indonesia,2011). Malaysia, Thailand and Vietnam (four) and Verrallina (three). A new species Ve. assamensis Subgenus Eumelanomyia is now represented by was described from Assam (Bhattacharyya et al. 2004a) Cx. hinglungensis was and Ve. consonensis Andaman and Nicobar Islands (Rajavel and Natarajan and10 speciesSingh 1989). in India. One species 2006). Ve. ceylonica waswas recorded recently forcollected the first from time Kerela from reportedIn subgenus first time Lophoceraomyia from Manipur one state new of India species (Rajput Cx. state (Vector Control Research Centre, Puducherry, India singhbhumensis unpublished report). Rajavel 2009). Several other species were also recorded Incertae sedis species: Nine species belonging to two from various parts was describedof India such from as Orissa Cx. lasiopalpis(Natarajan and groups “Aedes” sensu auctorum and “Ochlerotatus” sensu Cx. pholeter from south India (Reuben et al. 1993), Cx. auctorum, regarded to be of uncertain taxonomic position, quadripalpis from the northeast India (Bhattacharyya et al. 2003) and Cx. pilifemoralis and Cx. wilfredi from Jeypore formal status, the genus and subgenus of these species et al. 2005b), are new addition to are now under this head. Though, not considered as their hills, Orissa (Rajavel 1347 Bhattacharyya et al. | Mosquito biodiversity of India the mosquito fauna of India. Similarly Cx. aculeatus, Cx. Mosquito taxonomy provides essential inputs for vector paraculeatus and Cx. gracicornis from Assam, Cx. cubitatus control. In Vietnam, non-vector species An. varuna was from Andaman and Nicober Islands and Cx. inculus and Cx. An. minimus and targeted as vector demissus et (Bortel et al 2001). An. fluviatilis, a recognized primary al. 2011). Cx. wardi previously known only from Sri Lanka vectormisidentified of malaria as in the north-east India (Mohapatra was recently from collected Orissa are from new Kerela country state records (Vector (Rajavel Control et al. 1998) is now found to be a seasonal variant of An. Research Centre, Puducherry, unpublished report). minimus (Singh et al. 2010). In respect of malaria vectors, Maillotia is represented by only one Manguin et al. (2008) suggested that vector control in a species in India. region is dependent on the numbers and complexity of The subgenus Oculeomyia from synonymy the primary and secondary vectors and hence, requires as a new subgenus of genus Culex, which includes species study of the whole anopheline fauna in order to integrate earlierTanaka placed (2004) in resurrected the Bitaeniorhynchus Subgroup of the diverse information about vector system to formulate subgenus Culex appropriate and effective control strategy. Besides, sylvatic Cx. luzonensis resting cycles of some unknown arboviruses capable of spreading on vegetation . Thiswere subgenus collected includes from 6Alwar, species Rajasthan in India. (RajavelUnder this et alsubgenus. 2011). two females of of Chikungunya virus available in the web resource Genus Lutzia: It contains four species all in subgenus databaseto humans of may Chikungunya thrive in our Virus forest Net ecosystem. suggested The this history virus Metalutzia. A new species Lt. agranensis was described by to be originally circulated in the sylvatic cycle between Singh and Prakash (2008). forest dwelling mosquitoes and non-human primates, has now spread to different parts of the world including India.

India.Tribe Culisetini: This tribe includes a single genus. fauna study. Three species under two subgenera were recorded from TheseLimited facts studies clearly carried signify theout importancein some parts of mosquitoof India Ficalbia and Mimomyia Ficalbia and six since the 1980s detected several new species and many speciesTribe of Ficalbiini:Mimomyia Theare found tribe in includes India. two genera i.e. new country records. With the increasing mosquito borne . One species of diseases and changing ecology there is a need for the genus and represented by only one species in India. reassessment of Indian Culicidae fauna. Comprehensive Tribe Hodgesiini: Tribe Hodgesiini includes a single surveys with the incorporation of DNA-based methods Coquillettidia includes three species and genus such as DNA barcoding of mosquitoes (Pradeep Kumer et MansoniaTribe Mansoniini: is represented Two by genera four species are included in India. in this tribe. al. 2007) need to be undertaken on a large scale. The genus Acknowledgements: Researchers working in the Malariology Tribe Orthopodomyiini: Only one genus is included in department of Regional Medical Research Centre (RMRC), Dibrugarh are gratefully acknowledged for their assistance in the preparation of this onlythis tribe three which (Malaya is represented, Topomyia and by five Tripteroides species in) areIndia. found checklist. in India.Tribe GenusSabethini: Malaya Sabethini is represented includes by14 twogenera species of which and Tripteroides Literature Cited indicates that genus Tripteroides requires further studies problems, challenges and new initiatives. Medical Journal of Armed in India, especially by eight in bamboo species forested in India. areas Our of experience the north- Agarwal,Forces V.K. India and 62: V.K. 359–362 Sashindran. (doi: 2006. http://dx.doi.org/10.1016/S0377- Lymphatic Filariasis in India: 1237(06)80109-7). Barraud, P.J. 1934. The Fauna of British India, Including Ceylon and Burma. genus Topomyia was represented by only one species in Diptera. Vol V. Family Culicidae. Tribes Megarhinini and Culicini. India,east India. till Bhattacharyya Oriental in distribution et al. (2007) (Thurman added three 1959), more the www.mosquitocatalog. ). London: Taylor and Francis. 463 pp. (http:// high rainfall forested areas may increase the number of mosquitoesorg/files/pdfs/007000-28.pdf of India. Mosquito Systematics 4: 41–44 (http://www. species in from this Arunachalgenus. Pradesh. Further collections in Berlin, O.G.W. 1972. A proposal for a systematic study). of the culicine

Armigeresmosquitocatalog.org/files/pdfs/MS04N02P041.pdf joloensis (Diptera: Culicidae), a rare mosquito in upper Toxorhynchites includes nine species Bhattacharyya, D.R., A. Prakash, Entomon S.C. Tewari 25: 63–65. and J. Mahanta. 2000. in India.Tribe Tx. Toxorhynchitini: tyagii is a new Onlyspecies one described genus is from included Nilgiri in Bhattacharyya, D.R., A. Prakash, P.K. Mohapatra and J. Mahanta. 2002. A hills,this tribe. Western The genusGhats, southern India (Krishnamoorthy et noteAssam: on firstthe occurrencereport from of India. Aedes (Paraedes) ostentatio (Leicester) in Dibrugarh district, Assam, India (Diptera: Culicidae). Oriental Insects al. 2013). 36: 47–49. Uranotaenia is the only genus Bhattacharyya, D.R., A. Prakash, P.K. Mohapatra and J. Mahanta. in this tribe represented by 28 species (15 in subgenus Culex (Lophoceraomyia) quadripalpis, Culex Tribe Uranotaeniini: (Lophoceraomyia) mammilifer and Uranotaenia (Pseudoficalbia) Pseudoficalbia and 13 in subgenus Uranotaenia) in India. 2003.novobscura Occurrence in Assam. of Journal of the American Mosquito Control Ur. micans Association state (Rajput and Singh 1990). Subsequently, Ur. ohamai MQ0018.pdf). was first reported from India from Manipur 19: 13–14 (http://www.mosquitocatalog.org/files/pdfs/ was recorded from Western Ghats, South India (Reuben 2004a. Verrallina (Neomacleaya) assamensis (Diptera: Culicidae), et al. 1993). Ur. dibrugarhensis is a new species described Bhattacharyya,a new species D.R., from A. Prakash, Assam, S.C. India. Tewari, Journal P.K. Mohapatraof the American and J. Mahanta.Mosquito from Dibrugarh district of Assam (Bhattacharyya et al. Control Association 20: 115–120 (http://www.biodiversitylibrary. org/content/part/JAMCA/JAMCA_V20_N2_P115-120.pdf). 2004b). Ur. rutherfordi reported only from Sri Lanka Bhattacharyya, D.R., A. Prakash, P.K. Mohapatra and J. Mahanta. was recently collected from Kerela state (Vector Control 2004b. Uranotaenia dibrugarhensis, a new species in subgenus Research Centre, Puducherry, unpublished report). Pseudoficalbia (Diptera: Culicidae) from Assam, India. Journal of the American Mosquito Control Association 20: 1–5 (http://www. At least 31 species are recognized in India for biodiversitylibrary.org/content/part/JAMCA/JAMCA_V20_N1_P001- 005.pdf). transmitting various pathogens to humans (Table 2). 1348 Bhattacharyya et al. | Mosquito biodiversity of India

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Aedes and R. Rajendran. 2009. Description of a new species Anopheles (Diceromyia) from south India. Mosquito Systematics 24: 154–174 pseudosundaicus (Diptera: Culicidae) from Kerela, India. Zootaxa 2219: Tewari,( S.C. and J. Hiriyan. 1992. Description of two new species of ). 49–60 (http://www.mapress.com/zootaxa/2009/f/z02219p060f.pdf). Aedes (Finlaya) niveus (Diptera:http://www.mosquitocatalog.org/files/pdfs/MS24N02P154.pdf Culicidae) from Andaman and Nicobar, India. Mosquito VII. Genus Aedomyia Contributions of the Tewari,Systematics S.C. and 27: J. Hiriyan. 167–176 1995. ( Description of Tyson,American W.H. 1970. Entomological Contributions Institute to the ( Annmosquito Arbor fauna) 6: 1–27 of Southeast (http://www. Asia. pdfs/MS27N03P167.pdf). Theobald in Southeast). Asia. http://www.mosquitocatalog.org/files/Aedes (Stegomyia) krombeini Huang (Diptera: Culicidae) in India. Current mosquitocatalog.org/files/pdfs/WR44.pdf Anopheles Tewari,Science S.C., 56: J. 736–737. Hiriyan and R. Reuben. 1987. Occurrence of Walton,dirus C., J.M. Handley, C. Kuvangkadilok, F.H. Collins, R.E. Harbach, V. reaction.Baimai and Medical R.K. Butlin. and Veterinary 1999. Identification Entomology of five13: species24–32. of Wuchereria bancrofti infection in Nicobar Islands, India. Transactions complex from Thailand, usingAnopheles allele species specific complexes polymerase in Southchain Tewari,of the S.C., Royal J. Hiriyan Society and of TropicalR. Reuben. Medicine 1995. Epidemiologyand Hygiene 89: of 163–166.subperiodic and South East Asia World(http://apps.searo.who.int/pds_docs/B2406.pdf Health Organization, 2007. ). University of Maryland Agricultural Experimental . SEARO Technical Publication No. 57. 102 pp Thurman,Station E.B. Bulletin 1959. A-100: A contribution 182 pp. ( http://www.mosquitocatalog.org/ to a revision of the Culicidae of January 2014 northern Thailand. ) Received: December 2014 Paramasivan,files/pdfs/132400-2.pdf K.J. Krishnamoorthy, S. Dhananjeyan, L. Victorjerald Accepted: October 2014 Marco Gottschalk Tyagi, B.K., J. Hiriyan, S.C. Tewari, K. Ayanar, P. Samuel, N. Arunachalam, R. Published online: Editorial responsibility:

Appendix 1. Systematic checklist of mosquitoes of India.

Subfamily Anophelinae Genus Anopheles Meigen, 1818 Subgenus Anopheles Meigen, 1818

Anopheles Series ANGUSTICORNAitkenii SECTION Group Anopheles (Anopheles) aitkenii James, 1903 Anopheles (Anopheles) bengalensis Puri, 1930 Anopheles (Anopheles) insulaeflorum (Swellengrebel & Swellengrebel de Graaf, 1920) Anopheles (Anopheles) pinjaurensis Barraud, 1932 Culiciformis Group Anopheles (Anopheles) culiciformis Cogill, 1903 Anopheles (Anopheles) sintoni Puri, 1929 Lindesayi Group Gigas Complex Anopheles (Anopheles) baileyi Edwards, 1929 Anopheles (Anopheles) gigas Giles, 1901 Lindesayi Complex Anopheles (Anopheles) lindesayi Giles, 1900 Anopheles (Anopheles) nilgiricus Christophers, 1924 Plumbeus Group Anopheles (Anopheles) barianensis James, 1911 Lophoscelomyia Series Asiaticus Group Anopheles (Anopheles) annandalei Prashad, 1918 Interruptus Subgroup Anopheles (Anopheles) interruptus Puri, 1929

Myzorhynchus Series LATICORNBarbirostris SECTION Group Barbirostris Subgroup Anopheles (Anopheles) barbirostris van der Wulp, 1884 Anopheles (Anopheles) hodgkini Reid, 1962 Vanus Subgroup Anopheles (Anopheles) ahomi Chowdhury, 1929 Anopheles (Anopheles) barbumbrosus Strickland & Chowdhury, 1927 Hyrcanus Group Anopheles (Anopheles) argyropus (Swellengrebel, 1914) Anopheles (Anopheles) sinensis Wiedemann, 1828 Lesteri Subgroup Anopheles (Anopheles) crawfordi Reid, 1953 Anopheles (Anopheles) lesteri Baisas & Hu, 1936 Anopheles (Anopheles) peditaeniatus (Leicester, 1908) Nigerrimus Subgroup Anopheles (Anopheles) nigerrimus Giles, 1900 Anopheles (Anopheles) nitidus Harrison, Scanlon & Reid, 1973

Letifer Subgroup UmbrosusAnopheles Group (Anopheles) roperi Reid, 1950

Anopheles (Anopheles) umbrosus Subgenus Cellia Umbrosus Subgroup Myzomyia Series (Theobald, 1903) Theobald,Anopheles 1902 (Cellia) majidi Young & Majid, 1928

Anopheles (Cellia) jeyporiensis James, 1902 FunestusAconitus Group Subgroup

1351 Bhattacharyya et al. | Mosquito biodiversity of India

Appendix 1. Continued. Anopheles (Cellia) aconitus Donitz, 1902 Anopheles (Cellia) varuna Iyengar, 1924 Culicifacies Subgroup Anopheles (Cellia) culicifacies Giles, 1901 (species A, B, C, D and E) Minimus Subgroup

Anopheles (Cellia) fluviatilis FluMinimusviatilis Complex Complex Anopheles (Cellia) minimus James, 1902 (species S, T, U and V) Neocellia Series Anopheles (Cellia) karwari (James, 1903) Theobald, 1901 Anopheles (Cellia) moghulensis Christophers, 1924 Anopheles (Cellia) pulcherrimus Anopheles (Cellia) stephensi Liston, 1901 Anopheles (Cellia) theobaldi Giles, Theobald, 1901 1902 Annularis Group Anopheles (Cellia) annularis van der Wulp, 1884 (species A and B) Anopheles (Cellia) pallidus Anopheles (Cellia) philippinensis Ludlow, 1902 Nivipes Complex Theobald, 1901 Anopheles (Cellia) nivipes Jamesii Group Anopheles (Cellia) jamesii (Theobald, 1903) Anopheles (Cellia) pseudojamesi Strickland & Chowdhury, 1927 Anopheles (Cellia) splendidus Theobald, Koidzumi, 1901 1920 Maculatus group Anopheles (Cellia) pseudowillmori Anopheles (Cellia) willmori (James, 1903) Maculatus Subgroup (Theobald, 1910) Anopheles (Cellia) dravidicus Christophers, 1924 Anopheles (Cellia) maculatus Sawadwongporni Subgroup Anopheles (Cellia) sawadwongporni Theobald, Rattanarithikul 1901 & Green, 1987 Anopheles (Cellia) rampae Harbach & Somboon, 2011 Neomyzomyia Series Kochi Group Anopheles (Cellia) kochi Donitz, 1901 Leucosphyrus Group Hackeri Subgroup Anopheles (Cellia) mirans Sallum & Peyton, 2005 Leucosphyrus Subgroup Dirus complex Anopheles (Cellia) baimaii Sallum & Peyton, 2005 Anopheles (Cellia) elegans (James, 1903)

Anopheles (Cellia) tessellatus Paramyzomyia Tessellatus SeriesGroup Cinereus Group Theobald, 1901 Anopheles (Cellia) turkhudi Liston, 1901 Pyretophorus Series Anopheles (Cellia) pseudosundaicus Dhanajeyan, Leo & Rajendran, 2009) Anopheles (Cellia) vagus Donitz, 1902 (Tyagi, Hiriyan, Tewari, Ayanar, Samuel, Arunachalam, Paramsivam, Krishnamoorthy, Subpictus complex Anopheles (Cellia) subpictus Grassi, 1899 (species A, B, C and D) Sundaicus Complex Anopheles (Cellia) sundaicus (Rodenwaldt, 1925) (cytotype D) Subfamily Culicinae Tribe Aedeomyiini Genus Aedeomyia Subgenus Aedeomyia Theobald,Aedeomyia 1901 (Aedeomyia) catastica Knab, 1909 Tribe Aedini Theobald, 1901 Genus “Aedes” sensu auctorum Aedes kolhapuriensis Sathe & Girhe, 2001 Aedes panchgangee Sathe & Girhe, 2001 Aedes sangitee Sathe & Girhe, 2001 Aedes sangiti Girhe & Sathe, 2001 Genus Aedimorphus Aedimorphus alboscutellatus AedimorphusTheobald, 1903 caecus Aedimorphus culicinus (Edwards, (Theobald, 1922) 1905) Aedimorphus jamesi (Edwards,(Theobald, 1914) 1901) Aedimorphus lowisii Aedimorphus nigrostriatus (Barraud, 1927) Aedimorphus pallidostriatus (Theobald, 1910) Aedimorphus pampangensis (Ludlow, 1905) Aedimorphus pipersalatus (Giles, (Theobald, 1902) 1907) Aedimorphus punctifemoris (Ludlow, 1921) Aedimorphus stenoetrus

(Theobald, 1907) 1352 Bhattacharyya et al. | Mosquito biodiversity of India

Appendix 1. Continued. Aedimorphus syntheticus (Barraud, 1928) Aedimorphus taeniorhynchoides (Christophers, 1911) Aedimorphus trimaculatus Aedimorphus vexans (Meigen, 1830) Genus Armigeres (Theobald, 1905) Subgenus Armigeres Theobald,Armigeres 1901 (Armigeres) aureolineatus (Leicester, 1908) ArmigeresTheobald, (Armigeres) 1901 durhami (Edwards, 1917) Armigeres (Armigeres) joloensis (Ludlow, 1904) Armigeres (Armigeres) kesseli Ramalingam, 1987 Armigeres (Armigeres) kuchingensis Edwards, 1915 Armigeres (Armigeres) mahantai Bhattacharyya, Prakash, Mohapatra & Sarma, 2009 Armigeres (Armigeres) pallithorax Dong, Zhou & Dong, 2004 Armigeres (Armigeres) subalbatus (Coquillett, 1898) Armigeres (Armigeres) theobaldi Barraud, 1934 Subgenus Leicesteria Armigeres (Leicesteria) annulipalpis ArmigeresTheobald, (Leicesteria 1904) annulitarsis (Leicester, 1908) Armigeres (Leicesteria) cingulatus (Leicester,(Theobald, 1908) 1910) Armigeres (Leicesteria) dentatus Barraud, 1927 Armigeres (Leicesteria) digitatus (Edwards, 1914) Armigeres (Leicesteria) dolicocephalus (Leicester, 1908) Armigeres (Leicesteria) flavus (Leicester, 1908) Armigeres (Leicesteria) inchoatus Barraud, 1927 Armigeres (Leicesteria) longipalpis (Leicester, 1904) Armigeres (Leicesteria) magnus Armigeres (Leicesteria) omissus (Edwards, 1914) Genus Ayurakitia (Theobald, 1908) Ayurakitia peytoni (Reinert, 1972) Genus BruceharrisoniusThurman, Reinert, 1954 2003 Bruceharrisonius aureostriatus (Doleschall, 1857) Bruceharrisonius christophersi (Edwards, 1922) Bruceharrisonius doonii (Wattal, Bhatia & Kalra, 1958) Bruceharrisonius greenii Genus Christophersiomyia Barraud, 1923 Christophersiomyia annulirostris(Theobald, 1903) Christophersiomyia gombakensis (Mattingly, 1959) Christophersiomyia ibis (Barraud, (Theobald, 1931) 1905) Christophersiomyia thomsoni Genus Cancraedes Edwards, 1929 Cancraedes cancricomes (Edwards, (Theobald, 1922) 1905) Cancraedes simplex Genus Collessius Reinert, Harbach & Kitching, 2006 Subgenus Collessius Reinert, Harbach (Theobald, & Kitching, 1903) 2006 Collessius (Collessius) elsiae (Barraud, 1923) Collessius (Collessius) macdougalli (Edwards, 1922) Collessius (Collessius) shortti (Barraud, 1923) Subgenus Alloeomyia Reinert, Harbach & Kitching, 2008 Collessius (Alloeomyia) pseudotaeniatus (Giles, 1901) Genus Danielsia Danielsia albotaeniata (Leicester, 1904) Theobald,Danielsia 1904 lepchana (Barraud, 1923) Genus Dendroskusea Edwards, 1929 Dendroskusea kanarensis (Edwards, 1934) Dendroskusea micropterus (Giles, 1901) Dendroskusea periskelata (Giles, 1902) Dendroskusea ramachandrai (Reuben, 1967) Dendroskusea reginae (Edwards, 1922) Genus Downsiomyia Vargas, 1950 Downsiomyia albolateralis Downsiomyia albonivea (Barraud, 1934) Downsiomyia mohani (Knight, (Theobald, 1969) 1908) Downsiomyia nivea (Ludlow, 1903) Downsiomyia niveoides (Barraud, 1934) Downsiomyia novonivea (Barraud, 1934) Genus Edwardsaedes Belkin, 1962 Edwardsaedes imprimens (Walker, 1861) Genus Finlaya Finlaya flavipennis Giles, 1904 Theobald,Finlaya 1903 poicilia Genus Fredwardsius Reinert, 2000 Fredwardsius vittatus Theobald, (Bigot, 1903 1861) Genus Gilesius Reinert, Harbach & Kitching, 2006 Gilesius pulchriventer (Giles, 1901) Genus Himalaius Reinert, Harbach & Kitching, 2006 Himalaius gilli (Barraud, 1924) Himalaius simlensis (Edwards, 1922) Genus Heizmannia Ludlow, 1905 Subgenus Heizmannia Ludlow, 1905 Heizmannia (Heizmannia) aureochaeta (Leicester, 1908)

1353 Bhattacharyya et al. | Mosquito biodiversity of India

Appendix 1. Continued. Heizmannia (Heizmannia) chandi Edwards, 1922 Heizmannia (Heizmannia) chengi Lien, 1968 Heizmannia (Heizmannia) complex Heizmannia (Heizmannia) covelli Barraud, 1929 Heizmannia (Heizmannia) funerea (Leicester,(Theobald, 1908) 1910) Heizmannia (Heizmannia) greenii Heizmannia (Heizmannia) himalayensis Edwards, 1922 Heizmannia (Heizmannia) indica (Theobald, 1905) Heizmannia (Heizmannia) reidi Mattingly, 1957 Heizmannia (Heizmannia) viridis (Theobald,Barraud, 1929 1905) Subgenus Mattinglyia Lien, 1968 Heizmannia (Mattinglyia) discrepans (Edwards, 1922) Heizmannia (Mattinglyia) tripunctata Genus Hopkinsius Reinert, Harbach & Kitching, 2008 Subgenus Yamada Reinert, Harbach & Kitching, 1908 (Theobald, 1908) Hopkinsius (Yamada) albocinctus (Barraud, 1924) Genus Hulecoeteomyia Hulecoeteomyia chrysolineata HulecoeteomyiaTheobald, 1904 formosensis (Yamada, 1921) Hulecoeteomyia harveyi (Barraud, (Theobald, 1923) 1907) Hulecoeteomyia pallirostris (Edwards, 1922) Hulecoeteomyia saxicola (Edwards, 1922) Genus Jihlienius Reinert, Harbach & Kitching, 2006 Jihlienius unicinctus Edwards, 1922 Genus Kenknightia Reinert, 1990 Kenknightia dissimilis (Leicester, 1908) Kenknightia karwari (Barraud, 1924) Genus Lorrainea Belkin, 1962 Lorrainea amesii (Ludlow, 1903) Lorrainea fumida (Edwards, 1928) Genus Mucidus Subgenus Mucidus Theobald,Mucidus 1901 (Mucidus) laniger (Wiedemann, 1820) Mucidus Theobald, (Mucidus 1901) quasiferinus (Mattingly, 1961) Mucidus (Mucidus) scatophagoides Genus Neomelaniconion Newstead, 1907 Neomelaniconion lineatopenne (Ludlow, Theobald, 1905) 1901 Genus Ochlerotatus Lynch Arribalzaga, 1891

Ochlerotatus pullatus (Coquillett, 1904) Genus Unassi “Ochlerotatusgned to Subgenus” sensu auctorum Subgenus ‘Finlaya’ sensu auctorum Ochlerotatus (Finlaya) auronitens Edwards, 1922 Ochlerotatus (Finlaya) oreophilus Edwards, 1916 Ochlerotatus (Finlaya) sintoni (Barraud, 1924) Ochlerotatus (Finlaya) suffusus (Edwards, 1922) Ochlerotatus (Finlaya) versicolor (Barraud, 1924) Genus Paraedes Edwards, 1934 Paraedes barraudi Edwards, 1934 Paraedes chrysoscuta Paraedes menoni Mattingly, 1958 Paraedes ostentatio Leicester, (Theobald, 1908 1910) Genus Petermattinglyius Reinert, Harbach & Kitching, 2009 Subgenus Petermattinglyius Reinert, Harbach & Kitching, 2009 Petermattinglyius (Petermattinglyius) franciscoi (Mattingly, 1959) Petermattinglyius (Petermattinglyius) iyengari (Edwards, 1923) Genus Phagomyia Phagomyia assamensis Theobald,Phagomyia 1905 cacharana (Barraud, 1923) Phagomyia cogilli (Edwards, (Theobald, 1922) 1908) Phagomyia deccana (Barraud, 1923) Phagomyia feegradei (Barraud, 1934) Phagomyia gubernatoris (Giles, 1901) Phagomyia inquinata Edwards, 1922 Phagomyia khazani (Edwards, 1922) Phagomyia lophoventralis Phagomyia prominens (Barraud, 1923) Phagomyia stevensoni (Barraud, (Theobald, 1923) 1910) Genus Rhinoskusea Edwards, 1929 Rhinoskusea longirostris (Leicester, 1908) Rhinoskusea portonovoensis Rhinoskusea wardi (Reinert, 1976) Genus Scutomyia (Tewari & Hiriyan, 1992) Scutomyia albolineata Genus Stegomyia Theobald, 1904 Subgenus Actinothrix Reinert, HarbachTheobald, & Kitching, 1904 2009 Theobald,Stegomyia 1901 (Actinothrix) edwardsi Barraud, 1923 Stegomyia (Actinothrix) seampi (Huang, 1974) Subgenus Heteraspidion Reinert, Harbach & Kitching, 2009 Stegomyia (Heteraspidion) annandalei Stegomyia (Heteraspidion) craggy Barraud, 1923 Theobald, 1910 1354 Bhattacharyya et al. | Mosquito biodiversity of India

Appendix 1. Continued. Subgenus Huangmyia Reinert, Harbach & Kitching, 2009 Stegomyia (Huangmyia) mediopunctata Stegomyia (Huangmyia) perplexa Leicester, 1908 Subgenus Stegomyia Theobald, 1905 Stegomyia (Stegomyia) aegypti (Linnaeus, 1762) Subgenus Xyele Reinert,Theobald, Harbach 1901 & Kitching, 2006 Stegomyia (Xyele) desmotes Giles, 1904 Species without subgeneric placement Stegomyia w-alba Stegomyia gardnerii Ludlow, 1905 Stegomyia albopictaTheobald, (Skuse, 1895)1905 Stegomyia novalbopicta (Barraud, 1931) Stegomyia patriciae (Mattingly, 1954) Stegomyia pseudalbopicta Borel, 1928 Stegomyia subalbopicta (Barraud, 1931) Stegomyia unilineata Stegomyia krombeini (Huang, 1975) Stegomyia malayensis(Theobald, (Colless, 1962) 1906) Stegomyia scutellaris (Walker, 1858) Genus Udaya Udaya argyrurus (Edwards, 1934) Thurman,Udaya 1954 subsimilis (Barraud, 1927) Genus Tewarius Reinert, 2006 Tewarius agastyai Tewarius nummatus (Edwards, 1923) Tewarius reubenae (Tewari & Hiriyan, 1992) Genus Verrallina Subgenus Harbachius Reinert, 1999 (Tewari & Hiriyan, 1992) Theobald,Verrallina 1903 (Harbachius) abdita (Barraud, 1931) Verrallina (Harbachius) consonensis (Reinert, 1973) Verrallina (Harbachius) uniformis Verrallina (Harbachius) yusafi (Barraud, 1931) Subgenus Neomacleaya (Theobald, 1910) Verrallina (Neomacleaya) agrestis (Barraud, 1931) VerrallinaTheobald, (Neomacleaya 1907) andamanensis (Edwards, 1922) Verrallina (Neomacleaya) assamensis Verrallina (Neomacleaya) atria (Barraud, 1928) Verrallina (Neomacleaya) cauta (Barraud, Bhattacharyya, 1928) Tewari, Prakash, Mohapatra & Mahanta, 2004 Verrallina (Neomacleaya) ceylonica (Edwards, 1917) Verrallina (Neomacleaya) clavata (Barraud, 1931) Verrallina (Neomacleaya) comata (Barraud, 1931) Verrallina (Neomacleaya) indica Verrallina (Neomacleaya) pseudodiurna Verrallina (Neomacleaya) pseudomediofasciata (Theobald, 1907) Verrallina (Neomacleaya) rami (Barraud, (Theobald, 1928) 1910) Verrallina (Neomacleaya) seculata (Menon, 1950) (Theobald, 1910) Verrallina (Neomacleaya) unca Verrallina (Neomacleaya) vallistris (Barraud, 1928) Verrallina (Neomacleaya) yerburyi (Theobald, (Edwards, 1901) 1917) Subgenus Verrallina Verrallina (Verrallina) butleri Verrallina Theobald, (Verrallina 1903) dux (Dyar & Shannon, 1925) Verrallina (Verrallina) lugubris (Theobald, (Barraud, 1901)1928) Tribe Culicini Genus Culex Linnaeus, 1758 Subgenus Culex Linnaeus, 1758 Pipiens Group Culex (Culex) nilgiricus Edwards, 1916 Pipiens Subgroup Culex (Culex) quinquefasciatus Say, 1823

Culex (Culex) theileri Theileri Subgroup Culex (Culex) hutchinsoniTheobald, Barraud, 1903 1924 TrifilatusCulex Subgroup(Culex) vagans Wiedemann, 1828

Culex (Culex) fuscocephala UnivittatusCulex (Culex Subgroup) perexiguus Sitiens Group Theobald, 1907 Barraudi Subgroup Theobald, 1903 Culex (Culex) barraudi Edwards, 1922 Culex (Culex) edwardsi Barraud, 1923 Gelidus Subgroup Culex (Culex) gelidus Mimeticus Subgroup Culex (Culex) jacksoni Theobald, Edwards, 19011934 Culex (Culex) mimeticus Noe, 1899 Culex (Culex) mimuloides Barraud, 1924 Culex (Culex) mimulus Edwards, 1915 Culex (Culex) murrelli Lien, 1968

1355 Bhattacharyya et al. | Mosquito biodiversity of India

Appendix 1. Continued. Sitiens Subgroup Culex (Culex) sitiens Wiedemann, 1828 Culex (Culex) whitmorei (Giles, 1904) Vishnui Subgroup Culex (Culex) alienus Colless, 1957 Culex (Culex) pseudovishnui Colless, 1957 Culex (Culex) perplexus Leicester, 1908 Culex (Culex) tritaeniorhynchus Giles, 1901 Culex (Culex) vishnui Culex (Culex) whitei Barraud, 1923 Subgenus Culiciomyia Theobald, 1901

Culex (CuliciomyiaTheobald, 1907) fragilis Ludlow, 1903 Fragilis GroupCulex (Culiciomyia) nigropunctatus Edwards, 1926 Culex (Culiciomyia) pallidothorax Culex (Culiciomyia) ramakrishnii Wattal & Kalra, 1965 Culex (Culiciomyia) scanloni Bram,Theobald, 1967 1905 Culex (Culiciomyia) spathifurca (Edwards, 1915) Culex (Culiciomyia) viridiventer Giles, 1901 Shebbearei Group Culex (Culiciomyia) bailyi Barraud, 1934 Culex (Culiciomyia) shebbearei Barraud, 1924 Subgenus Eumelanomyia Mochthogenes Group Theobald, 1909 Culex (Eumelanomyia) foliatus Brug, 1932 FHinglungensisoliatus Subgroup Subgroup Culex (Eumelanomyia) castrensis Edwards, 1922 Culex (Eumelanomyia) hinglungensis Chu, 1957 Iphis Subgroup Culex (Eumelanomyia) iphis Barraud, 1924 Khazani Subgroup Culex (Eumelanomyia) khazani Edwards, 1922 Malayi Subgroup Culex (Eumelanomyia) malayi (Leicester, 1908) Pluvialis Subgroup Culex (Eumelanomyia) pluvialis Barraud, 1924

Culex (Eumelanomyia) tenuipalpis Barraud, 1924 TenuipalpisCulex (Eumelanomyia Subgroup ) mohani Sirivanakarn, 1977 Protomelanoconion Group Culex (Eumelanomyia) brevipalpis (Giles, 1902) Subgenus Lophoceraomyia

Theobald, 1905 FraudatrixCinctellus Group complex FraudatrixCulex Subgroup (Lophoceraomyia) cinctellus Edwards, 1922 Quadripalpis complex Culex (Lophoceraomyia) aculeatus Colless, 1965 Culex (Lophoceraomyia) paraculeatus Sirivanakarn, 1977 Culex (Lophoceraomyia) quadripalpis (Edwards, 1914) Rubithoracis complex Culex (Lophoceraomyia) rubithoracis (Leicester, 1908) Seniori complex Culex (Lophoceraomyia) seniori Barraud, 1934 Variatus complex Culex (Lophoceraomyia) cubitatus Colless, 1965 Culex (Lophoceraomyia) gracicornis Sirivanakarn, 1977 Culex (Lophoceraomyia) variatus (Leicester, 1908) Culex (Lophoceraomyia) macdonaldi Colless, 1965 Inculus complex Culex (Lophoceraomyia) inculus Colless, 1965 Minutissimus Subgroup Culex (Lophoceraomyia) minutissimus Culex (Lophoceraomyia) infantulus Edwards, 1922 Mammilifer Group (Theobald, 1907) Mammilifer Subgroup

Culex (Lophoceraomyia) flavicornis Barraud, 1924 FlavicornisCulex (Lophoceraomyia complex ) lasiopalpis Sirivanakarn, 1977 Culex (Lophoceraomyia) raghavanii Rahman, Chowdhury & Kalra, 1968 Singhbhumensis complex Culex (Lophoceraomyia) singhbhumensis Natarajan & Rajavel, 2009 Mammilifer complex Culex (Lophoceraomyia) mammilifer (Leicester, 1908) Culex (Lophoceraomyia) demissus Colless, 1965 Culex (Lophoceraomyia) wardi Sirivanakarn, 1977

Culex (Lophoceraomyia) uniformis Traubi complex (Theobald, 1905)

1356 Bhattacharyya et al. | Mosquito biodiversity of India

Appendix 1. Continued. Minor complex Culex (Lophoceraomyia) bengalensis Barraud, 1934 Culex (Lophoceraomyia) bicornutus Culex (Lophoceraomyia) minor (Leicester, 1908) Peytoni complex (Theobald, 1910) Culex (Lophoceraomyia) peytoni Bram & Rattanarithikul, 1967 Pholeter complex Culex (Lophoceraomyia) pholeter Bram & Rattanarithikul, 1967 Wilfredi Group Culex (Lophoceraomyia) pilifemoralis Culex (Lophoceraomyia) wilfredi Colless, 1965 Subgenus Maillotia Wang & Feng, 1964 Hortensis Group CulexTheobald, (Maillotia 1907) hortensis Subgenus Oculeomyia Bitaeniorhynchus Complex Ficalbi, 1889 Culex (OculeomyiaTheobald, 1907) bitaeniorhynchus Giles, 1901 Culex (Oculeomyia) infula Culex (Oculeomyia) luzonensis Sirivanakarn, 1976 Sinensis Complex Theobald, 1901 Culex (Oculeomyia) cornutus Edwards, 1922 Culex (Oculeomyia) epidesmus Culex (Oculeomyia) sinensis Genus Lutzia (Theobald, 1910) Subgenus Metalutzia Theobald, 1903 Theobald,Lutzia 1903 (Metalutzia) agranensis Singh & Prakash, 2008 Lutzia Tanaka, (Metalutzia 2000) fuscana Wiedemann, 1820 Lutzia (Metalutzia) halifaxii Lutzia (Metalutzia) vorax Edwards, 1921 Tribe Culisetini Theobald, 1903 Genus Culiseta Subgenus Allotheobaldia Brolemann, 1919 Culiseta Felt,(Allotheobaldia 1904 ) longiareolata (Macquart, 1838) Subgenus Culiseta Culiseta (Culiseta) alaskaensis (Ludlow, 1906) Culiseta Felt, 1904(Culiseta) niveitaeniata Tribe Ficalbiini Genus Ficalbia (Theobald, 1907) Ficalbia minima Genus Mimomyia Theobald, 1903 Subgenus Etorleptiomyia (Theobald, 1901) Theobald,Mimomyia 1903 (Etorleptiomyia) luzonensis (Ludlow, 1905) Subgenus Mimomyia Theobald, 1904 Mimomyia (Mimomyia) aurea (Leicester, 1908) MimomyiaTheobald, (Mimomyia 1903 ) chamberlaini Ludlow, 1904 Mimomyia (Mimomyia) hybrida (Leicester, 1908) Mimomyia (Mimomyia) intermedia (Barraud, 1929) Subgenus Ingramia Edwards, 1912 Mimomyia (Ingramia) fusca (Leicester, 1908) Tribe Hodgesiini Genus Hodgesia Belkin, 1962 Hodgesia bailyi Barraud, 1929 Tribe Mansoniini Genus Coquillettidia Dyar, 1905 Subgenus Coquillettidia Dyar, 1905 Coquillettidia (Coquillettidia) crassipes (van der Wulp, 1881) Coquillettidia (Coquillettidia) novochracea (Barraud, 1927) Coquillettidia (Coquillettidia) ochracea Genus Mansonia Blanchard, 1901 Subenus Mansonioides (Theobald, 1903) Mansonia (Mansonioides) annulifera Mansonia Theobald, (Mansonioides 1907 ) dives (Schiner, 1868) Mansonia (Mansonioides) indiana Edwards, (Theobald, 1930 1901) Mansonia (Mansonioides) uniformis Tribe Orthopodomyiini Genus Orthopodomyia Belkin, Heinemann & Page, 1970 (Theobald, 1901) Orthopodomyia albipes Leicester, 1904 Orthopodomyia anopheloides (Giles, 1903) Orthopodomyia andamanensis Barraud, 1934 Orthopodomyia flavicosta Barraud, 1927 Orthopodomyia flavithorax Barraud, 1927 Tribe Sabethini Genus Malaya Leicester, 1908 Malaya genurostris Leicester, 1908 Malaya jacobsoni (Edwards, 1930) Genus Topomyia Leicester, 1908 Subgenus Suaymyia Topomyia (Suaymyia) cristata Subgenus Topomyia Thurman,Leicester, 19081959 Topomyia (Topomyia) bifurcata Thurman, Dong, Wang 1959 & Lu, 1995

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Appendix 1. Continued. Topomyia (Topomyia) aureoventer Topomyia (Topomyia) hirtusa Gong, 1989 Genus Tripteroides Giles, 1904 (Theobald, 1910) Subgenus Rachionotomyia Tripteroides (Rachionotomyia) affinis (Edwards, 1913) Tripteroides Theobald, (Rachionotomyia 1905 ) aranoides Tripteroides (Rachionotomyia) coonorensis Mattingly, 1981 Tripteroides (Rachionotomyia) edwardsi (Barraud, (Theobald, 1929) 1901) Tripteroides (Rachionotomyia) serratus (Barraud, 1929) Subgenus Tripteroides Giles, 1904 Tripteroides (Tripteroides) indicus (Barraud, 1929) Tripteroides (Tripteroides) similis (Leicester, 1908) Tripteroides (Tripteroides) tarsalis Tribe Toxorhynchitini Genus Toxorhynchites Delfinado & Hodges, 1968 Subgenus Toxorhynchites Toxorhynchites Theobald, 1901 (Toxorhynchites) albipes (Edwards, 1922) ToxorhynchitesTheobald, (Toxorhynchites 1901 ) edwardsi (Barraud, 1924) Toxorhynchites (Toxorhynchites) gravelyi (Edwards, 1921) Toxorhynchites (Toxorhynchites) kempi (Edwards, 1921) Toxorhynchites (Toxorhynchites) klossi (Edwards, 1921) Toxorhynchites (Toxorhynchites) metallicus Leicester, 1904 Toxorhynchites (Toxorhynchites) minimus Toxorhynchites (Toxorhynchites) splendens (Wiedemann, 1819) Toxorhynchites (Toxorhynchites) tyagii Krishnamoorthy (Theobald, 1905) et al., 2013 Tribe Uranotaeniini Genus Uranotaenia Lynch Arribalzaga, 1891 Subgenus Pseudoficalbia Section A Bicolor series Theobald, 1912 Uranotaenia (Pseudoficalbia) atra Uranotaenia (Pseudoficalbia) bicolor Leicester, 1908 Uranotaenia (Pseudoficalbia) lutescens Theobald, Leicester, 1905 1908 Uranotaenia (Pseudoficalbia) obscura Edwards, 1915 Bimaculata Series Uranotaenia (Pseudoficalbia) nivipleura Leicester, 1908 Uranotaenia (Pseudoficalbia) novobscura Barraud, 1934 Recondita Series Uranotaenia (Pseudoficalbia) dibrugarhensis Bhattacharyya, Prakash, Mohapatra & Mahanta, 2004 Uranotaenia (Pseudoficalbia) husaini Qutubuddin, 1947 Uranotaenia (Pseudoficalbia) luteola Edwards, 1934 Uranotaenia (Pseudoficalbia) maculipleura Leicester, 1908 Uranotaenia (Pseudoficalbia) mattinglyi Qutubuddin, 1951 Uranotaenia (Pseudoficalbia) recondita Edwards, 1922 Uranotaenia (Pseudoficalbia) ohamai Uranotaenia (Pseudoficalbia) stricklandi Barraud, 1926 Section B Tanaka, Mizusawa & Saugstad, 1975 Maxima Series Uranotaenia (Pseudoficalbia) maxima Leicester, 1908 Subgenus Uranotaenia Lynch Arribalzaga Uranotaenia (Uranotaenia) alboannulata Uranotaenia (Uranotaenia) annandalei Barraud, 1926 Uranotaenia (Uranotaenia) campestris Leicester, (Theobald, 1908 1905) Uranotaenia (Uranotaenia) christophersi Barraud, 1926 Uranotaenia (Uranotaenia) edwardsi Barraud, 1926 Uranotaenia (Uranotaenia) hebes Barraud, 1931 Uranotaenia (Uranotaenia) lateralis Ludlow, 1905 Uranotaenia (Uranotaenia) longirostris Leicester, 1908 Uranotaenia (Uranotaenia) micans Leicester, 1908 Uranotaenia (Uranotaenia) macfarlanei Edwards, 1914 Uranotaenia (Uranotaenia) orientalis Barraud, 1926 Uranotaenia (Uranotaenia) rutherfordi Edwards, 1922 Uranotaenia (Uranotaenia) testacea

Theobald, 1905

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