An updated checklist of mosquito species (Diptera: Culicidae) from Madagascar

An updated checklist of 235 mosquito species from Madagascar is presented. The number of species has increased considerably compared to previous checklists, particularly the last published in 2003 (178 species). This annotated checklist provides concise information on endemism, taxonomic position, developmental stages, larval habitats, distribution, behavior, and vector-borne diseases potentially transmitted. The 235 species belong to 14 genera: Aedeomyia (3 species), Aedes (35 species), Anopheles (26 species), Coquillettidia (3 species), Culex (at least 50 species), Eretmapodites (4 species), Ficalbia (2 species), Hodgesia (at least one species), Lutzia (one species), Mansonia (2 species), Mimomyia (22 species), Orthopodomyia (8 species), Toxorhynchites (6 species), and Uranotaenia (73 species). Due to non-deciphered species complexes, several species remain undescribed. The main remarkable characteristic of Malagasy mosquito fauna is the high biodiversity with 138 endemic species (59%). Presence and abundance of species, and their association, in a given location could be a bio-indicator of environmental particularities such as urban, rural, forested, deforested, and mountainous habitats. Finally, taking into account that Malagasy culicidian fauna includes 64 species (27%) with a known medical or veterinary interest in the world, knowledge of their biology and host preference summarized in this paper improves understanding of their involvement in pathogen transmission in Madagascar.

This list was compiled using our own observations, and Internet and bibliographic references. The Culicidae mosquito fauna includes 235 species within 14 genera. The present taxon identifications are based on formally recognized genera, and subgenera. Their abbreviations follow taxonomic nomenclature from A Catalog of the Mosquitoes of the World [132], its supplements [131,235,236] and Reinert [188,189], and Wilkerson et al. [241] for the names of tribe Aedini. The author is given at first mention of a species.
Each species can be cited as follows: genus (subgenus) species, author(s) and date of first description, new name according to Wilkerson et al. [241], author(s) and date of first mention in Madagascar, endemism, development stages, larval habitats, distribution, trophic behavior, and vector-borne diseases potentially transmitted. The relationship between the species distribution and the importance of Malagasy biodiversity has been discussed, raising questions about the mosquito's evolution and differentiation.

Presentation of Malagasy mosquitoes 2.1 Genus Anopheles Meigen, 1818
The genus Anopheles is subdivided into eight subgenera. The subgenera Anopheles and Cellia are present in Madagascar. The subgenus Anopheles is represented by more than 183 species in the world [119]. In Madagascar, three species are present and they belong to the Myzorhynchus Series. One is an endemic species. The subgenus Cellia is represented by 224 species in the world. In Madagascar, 23 species occur. Ten are endemic, and one species (Anopheles mascarensis de Meillon) is present in Madagascar (Theobald), and An. (Cel.) theileri Edwards are absent from Madagascar (this study, [5]) but were ranked by WRBU among the mosquito species found on the island [244]. Three names of doubtful validity (An. arnoulti, An. courdurieri, and An. fuscicolor soalalaensis), regarded as nomen dubium by Brunhes et al. [32], are not listed in this document.

Genus Aedes Meigen, 1818
Aedes is the largest genus in the subfamily Culicinae with 930 species [241]. Thirty-eight subgenera were reported by Knight and Stone in 1977 [132]. This genus was recently subdivided into 74 subgenera, which was restored to its status prior to the year 2000 [241]. According to Wilkerson et al. [241], 12 subgenera are present in Madagascar. These subgenera are represented by 35 species, 18 species are endemic to Madagascar, and two species are found in Madagascar and in the Comoros archipelago. Theobald, 1903 In Madagascar, this subgenus includes eight species, three are endemic. This list and Arim [5] did not include Aedes (Aedimorphus) ovazzai that may be erroneously reported to be present on the island by WRBU [244]. In Africa, species were involved in transmission of RVFV [93,147]. d Aedes (Aedimorphus) albodorsalis Fontenille & Brunhes, 1984 Fontenille & Brunhes, 1984 [86] Endemic. Only the adult female has been described [86]. Found in the eastern [86] and western bioclimatic domains [15]. Anthropophilic, diurnal, seems closely related to northeastern forest areas [86]. Not involved in disease transmission in Madagascar. d Aedes (Aedimorphus) durbanensis (Theobald, 1903) Ravaonjanahary, 1978 [182] Pupal stages undescribed. In Madagascar, larval habitats are ponds, grassy bottom-land, and ditches [182]. Occurs in southern and western Malagasy domains [85,170]. No medical and veterinary importance in Madagascar. However, was found naturally infected with RVFV in Kenya [168]. d Aedes (Aedimorphus) domesticus  Doucet, 1951 [67] Only Malagasy species belonging to the Group Domesticus [80]. Larval stages morphologically close to Aedes leptolabis Edwards which is absent from Madagascar [230]. In Madagascar, larval habitats are puddles near the sea [67]. Presence reported in Madagascar by Doucet [67] in Vangaindrano (eastern domain) but never confirmed. In Africa, involved in transmission of Bunyamwera virus (BUNV) in Cameroon [1]. d Aedes (Aedimorphus) mathioti Fontenille & Brunhes, 1984 Fontenille & Brunhes, 1984 [86] Endemic. Only adult female was described [86]. Rare, occurs in the lowland forest of the eastern Malagasy domain (Onive river valley and region of Soanierana-Ivongo). Diurnal and anthropophilic [86]. Not involved in disease transmission. d Aedes (Aedimorphus) natronius Edwards, 1932 Arim, 1959 [5] Egg and pupal stages undescribed. No literature has reported its presence in Madagascar. However, this species occurs in Toliara, southern domain, as shown on the labels of specimens stored at the Institut de Recherche pour le Développement (IRD) of Montpellier [5]. Biology unknown. Not involved in disease transmission in Madagascar. In Africa, arboreal and crepuscular species [113] and involved in transmission of Uganda S virus (UGSV) [59]. Theobald, 1903 Only one species belonging to the subgenus Catageiomyia is in Madagascar.

Subgenus Coetzeemyia Huang, Mathis, & Wilkerson, 2010
This subgenus was recently created by Huang et al. [126] and is monotypic. (Theobald, 1912) Edwards, 1920 [75] Egg and pupal stages undescribed. Subgenus was changed on several occasions. Presence reported in Madagascar by Edwards in 1920 [75]. Adult stage morphologically close to Aedes dufouri Hamon which is endemic to La Réunion and likely absent from Madagascar. In Madagascar, larval habits are rock holes [144] and its biology seems to be related to the presence of mangrove [182]. Occurs in the western and southern biogeographic domains [85]. Caught in abundance in human landing catches in the Morondava region. No medical and veterinary importance in Madagascar [85], but found naturally infected with Spondweni virus (SPOV) in a mixedspecies batch of mosquitoes in Mozambique [161]. Theobald, 1911 In Madagascar, this subgenus is represented by five endemic species.

Subgenus Mucidus
d Aedes (Mucidus) scatophagoides  Brunhes, 1968 [182] Eggs undescribed. Presence reported for the first time in Madagascar by Brunhes in 1968 [182]. Larval habitats are warm temporary pools exposed to sunlight [182]. Occurs in the southern domain, particularly in the Antanimena area, in the semi-arid Androy region [182]. No medical and veterinary importance in Madagascar.

Subgenus Neomelaniconion Newstead, 1907
This subgenus includes 28 species. Six species of Neomelaniconion are present in Madagascar. Five species of them are endemic and were described from specimens formerly called Ae (Neo.) palpalis that is absent from Madagascar [5,143], but ranked among the mosquito species found on the island by WRBU [244]. Vertical transmission of RVFV was described in species belonging to this subgenus [147].  [143] Endemic. Only the adult female was described [143]. Sequence variations of the ribosomal DNA ITS2 consistent with morphological observations, indicating that this species belongs to the Group Sylvaticum [130]. Occurs in forest areas of the western (forest station Ampijoroa), central (forest relic near Anjiro), and south (degraded forest near Mahabo) regions [143]. No medical and veterinary importance in Madagascar.  [130].
No specific differentiation at the molecular level obtained between Ae. belleci and Ae. nigropterum. In Madagascar, larval habitats are temporary ponds characterized by being slightly brownish in color and full of dead leaves and located in forest areas of medium altitude (1000 m asl), near Ranomafana, in the eastern biogeographic domain [143]. Adult biology is unknown because adults were obtained only from larval rearing. No medical and veterinary importance in Madagascar. d Aedes (Neomelaniconion) circumluteolum (Theobald, 1908) Hamon, 1959 [182] Pupal stages undescribed. Presence reported in Madagascar in the southern domain by Hamon in 1959 [182] and confirmed by molecular study which indicated that Malagasy and South African specimens share a common origin [130]. In Madagascar, larval habitats are temporary pools and puddles [28]. Occurs in coastal areas, in Nosy Be, in western, eastern, and central biogeographic domains [85]. Never captured in the semi-arid bioclimatic domains of south and south-west Madagascar [85]. Diurnal and anthropophilic species in forested areas [143]. Found naturally infected with WNV in Ampijoroa and involved in transmission or dissemination of this virus in Madagascar [85]. In Africa, prefers to feed on cattle and may feed occasionally on humans [114,182]. Involved in transmission of SIMV [45], WNV [127], SPOV [160], and PGAV [1]. Found naturally [133] and experimentally [225] infected with RVFV. Endemic. Only adult stages (male and female) were described to date [143]. Sequence variations in ribosomal DNA ITS2 consistent with morphological observations, indicating that this species belongs to the Group Sylvaticum [130]. No specific differentiation at molecular level obtained between Ae. fontenillei and Ae. sylvaticum. In Madagascar, larval habitats may be small depressions in forest areas [143]. Adults collected from humans or using a hand-net in forest undergrowth. Only occurs in the Périnet forest, in the eastern biogeographic domain [143]. No medical and veterinary importance in Madagascar. Endemic. Only the adult stages (male and female) were described to date [143]. Sequence variations in ribosomal DNA ITS2 consistent with morphological observations, indicating that this species belongs to the Group Circumluteolus [130]. No specific differentiation at molecular level obtained between this species and Ae. belleci. Only occurs in the Périnet forest, in the eastern biogeographic domain [143]. Adults were collected from humans or using a hand-net in forest undergrowth. No medical and veterinary importance in Madagascar.  [143] Endemic. Only the adult stages (male and female) were described [143]. Sequence variations in ribosomal DNA ITS2 consistent with morphological observations, indicating that this species belongs to the Group Sylvaticum [130]. No specific differentiation at molecular level obtained between this species and Ae. fontenillei. Probably present in the northern and eastern biogeographic domains [143]. Occurs in Sainte-Marie island, on the eastern edge of Madagascar (from Sambava region to Manakara region), and in medium mountainous areas (900 m asl). Collected using a hand-net in forest undergrowth. No medical and veterinary importance in Madagascar.

Subgenus Ochlerotatus Lynch Arribálzaga, 1891
This subgenus includes 187 species in the world [241]. Only Ae. (Och.) ambreensis is present in Madagascar. The report of Aedes dufouri as present in Madagascar in Arim dataset [5] is doubtful as information on collection areas is not available. De facto this information was treated as an error. Aedes dufouri was described from Réunion Island and occurs on Europa island (French island in the Mozambique Channel) [99].

Subgenus Zavortinkius
d Aedes (Zavortinkius) monetus Edwards, 1935Edwards, 1935 Eggs and pupal stages undescribed. First reported in Madagascar and was also collected on the islands of Comoros, Mayotte, and Moheli [24]. Larval habitats are tree holes filled with rainwater and plant organic matter [182]. Occurs in all Malagasy biogeographic domains (except the central domain) [85,182]. Not involved in disease transmission. van Someren, 1949 [229] Endemic. Eggs undescribed. Larval habitats are sectioned trunks of Ravenala sp. [22], tree holes [207], rarely leaf axils of Pandanus [182], bamboo ovitraps, and leaf axils of agave [85]. Essentially present in the warmer and humid eastern coast of Madagascar, also occurs in Nosy Komba, in the Sambirano area, western and northern [85] and the central [210] domains. Not involved in disease transmission. Dyar, 1905 [71] This genus includes 57 species in the world [117]. These species represent three subgenera: Coquillettidia, Rhynchotaenia, and Austromansonia. In Madagascar, only the subgenus Coquillettidia is present and it is represented by three species. Two of them are endemic to Madagascar. Larval and pupal stages of Coquillettidia species derive their oxygen by puncture of the aerenchyma of aquatic plants. The report of Coquillettidia (Coquillettidia) aurites (Theobald) as present in Madagascar in Arim dataset [5] is doubtful as information on collection areas is not available. De facto this information was treated as an error.

Genus Culex Linnaeus, 1758
The genus Culex includes 26 subgenera and 769 species in the world [119]. In total, 45 species from 6 subgenera were described in Madagascar. They include two Culex salisburiensis subspecies (Culex salisburiensis salisburiensis and Culex salisburiensis coursi). In Madagascar, like in many regions of the world, the systematics of Culex have to be revisited. The majority of these species belong to the subgenus Culex. Ten species are endemic to Madagascar. Culex salisburiensis coursi was described only from a single specimen.
Doucet, 1950 [65] Endemic. Only larval stages were described [65]. The type was collected in a rice field in the Ambositra region [65]. This larval specimen has never been found in Madagascar since that time. Morphologically close to Culex guiarti by the presence of nine posterolateral combs of segment VIII and well visible teeth of the mentum [65].
Fontenille & Jupp, 1989 [88] Eggs undescribed. Morphologically close to Cx. univittatus, also present in Madagascar. In Madagascar, larval habitats are still unknown. Presence reported only from the Tsiroanomandidy region [88]. Not involved in disease transmission. In Africa, may be attracted to domestic ruminants, poultry, and humans [10]. Involved in transmission of at least 16 viruses (Alphavirus, Flavivirus, Bunyaviruses, Orbivirus) [1] and avian Plasmodium parasite in Cameroon [171]. Seems to have a lower vectorial capacity than Cx. univittatus for the transmission of WNV [88].
Doucet, 1949 [64] Egg and pupal stages undescribed. Larval stages might be confused with those of Cx. carleti, which were described in 1971 [29]. Larval habitats are in the rice fields, water holes, swamps, puddles, lakes, and flooded grasslands [63,67]. In Madagascar, occurs in the eastern domain [63,67]. 1758 Edwards, 1920 [75] Eggs appear to be very similar in surface morphology to those of Culex quinquefasciatus Say [2]. In Madagascar, larval habitats are irrigation drains, canals, tires, and water tanks [207,227]. In the central domain, occurs essentially in urban and suburban areas [227], also abundant in the Anorana rainforest, in the Anjozorobe district, in Antananarivo province [210]. A few specimens were recently collected in the western domain [15,170]. Zoophilic species, prefers to feed on cattle and poultry. Not involved in disease transmission in Madagascar. In Africa, involved in transmission of RVFV [123], SINV [242], and avian Plasmodium parasite (presence of sporozoites) [199]. In North America, vector of WNV [8].
Edwards, 1941 [80] Eggs undescribed. Morphologically close to Culex decens and Cx. invidiosus, and distinguishable by male genitalia morphology. In Madagascar, larval habitats are unknown. Occurs near Lake Alaotra (in the eastern domain) [80], in the Tsiroanomandidy region (central domain) [85], in the Ihosy and Ampandrandava regions (at the boundary between the southern and western domains) [80]. Not involved in the transmission of diseases. d Culex (Culex) quinquefasciatus Say, 1823 Edwards, 1920 [80] Eggs appear to be very similar in surface morphology to those of Culex tritaeniorhynchus Giles [205] and Culex pipiens [2]. Belongs to the subgroup pipiens of the Group Pipiens [115] and has about 40 synonyms. In Madagascar, larval habitats are sewage, tires, plastic containers, and metal drums (Tantely, unpublished data). Occurs in all geographic domains with a preference for urban environments [85]. Anthropophilic species [85] and may be attracted to domestic ruminants (Luciano Tantely, unpublished observation). Involved in the transmission of Wuchereria bancrofti [20], WNV, BABV, and endemic PERV [85]. In the world, has a variable host preference from high degree of mammalophily and anthropophily to a high degree of ornithophily [206]. Involved in transmission of RVFV [198], CHIKV, WNV, and MgV [1]. Involved in transmission of avian Plasmodium parasite (presence of sporozoites) in India and the United States of America [199].
Edwards, 1920 [75] Endemic. Eggs, larval and pupal stages undescribed. Morphologically close to Cx. simpsoni and male genitalia of these two species are morphologically identical [80]. Presence only reported in Antananarivo city [85]. Further studies are needed to guarantee the status of this endemic species. Adult biology unknown. Not involved in disease transmission.
Clerc & Coulanges, 1979 [40] Eggs undescribed. In Madagascar, larval habitats are unknown. Presence only reported in the Andasibe-Mantadia forest (or Périnet forest) in the eastern domain [40,85]. Rare species. Adult biology unknown. BABV and RVFV were isolated from a mixed batch of mosquito species, including this species, collected in Périnet forest [85]. In Africa, ornithophilic species and competent vector of WNV under laboratory conditions [150]. Potential vector of RVFV [147].
Brunhes & Ravaonjanahary, 1973 [30] Eggs and pupal stages undescribed. In Madagascar, larval habitats are grassy holes [30]. Occurs in the eastern and central domains [30,85] and caught in human landing catches during daytime catches [85]. Not involved in disease transmission. In Africa, involved in transmission of WNV, MOSV, SINV [45], CHIKV, BABV, and WSLV. MgV was isolated from this species [1]. Theobald, 1907 [219] This subgenus includes 55 species in the world. Six species occur in Madagascar, two of them are endemic species to the island. The report of Cx. semibrunneus Edwards as present in Madagascar in the Arim dataset [5] is doubtful as information on collection areas is not available. De facto this information was treated as an error.
Brunhes, 1967 [5] Eggs, pupal stages, and adult female undescribed. No literature has reported the presence of Cx. subaequalis in Madagascar. Occurs on the island, as shown on the labels of specimens stored at the IRD of Montpellier [5]. The locality type was not well specified. Biology unknown. Not involved in disease transmission.

Subgenus Kitzmilleria Danilov, 1989
This subgenus includes only one species in the world: Cx.
Coulanges et al., 1977 [48] Eggs undescribed. Formerly classified in the subgenus Culex. In Madagascar, rare species and reported in the Andasibe-Mantadia forest (forested area near Périnet) in the eastern area [85]. Biology unknown. Not involved in disease transmission. In Africa, involved in transmission of NTAV [17]. Theobald, 1907 This subgenus, rehabilitated by Tanaka in 2004, is represented by 19 species in the world. Five species are present in Madagascar and one species is endemic to the island. This study includes Cx. aurantapex Edwards that was not ranked among the mosquito species found on the island in WRBU [244]. The morphological diversity and the worldwide distribution of the species of subgenus Oculeomyia suggest that it is an old group [203]. Similar characters are observable in the larval stages for the majority of these species [124].
Brunhes & Rambelo, 1968 [28] Endemic to Madagascar and to the Comoros archipelago (Mohéli) [28]. Eggs undescribed. Larval habitats are temporary pools and small grassy pools under forest cover [28,143]. Occurs on the eastern slopes of the central highlands and in the eastern domain [28,85]. Captured in human landing catches during daytime catches [85]. Not involved in disease transmission.
Grjebine, 1955 [5] Eggs undescribed. No literature has reported its presence in Madagascar. However, this species occurs in Nosy Be, in the Sambirano area, as shown on the labels of specimens stored at the IRD of Montpellier [5]. Biology unknown. Not involved in disease transmission. Edwards, 1922Doucet, 1950 Eggs undescribed. In Madagascar, larval habitats are tree holes [85], and phytotelmata [27], fresh water marshes (in Mahajanga city) [112]. Occurs in the western [103] and eastern domains [85]. Adult biology unknown. Not involved in disease transmission.
Grjebine, 1952 [5] Egg and pupal stages undescribed. No literature has reported its presence in Madagascar. However, this species occurs in Manakara, eastern domain, as shown on the labels of specimens stored at the IRD of Montpellier [5]. Biology unknown. Not involved in disease transmission. Theobald, 1907 This subgenus includes nine species in the world [119]. Two species of this subgenus are reported in Madagascar. The report of Cx. avianus de Meillon as present in Madagascar in the WRBU dataset [244] is doubtful as information on collection areas is not available. De facto this information was treated as an error.

Subgenus Maillotia
In Madagascar, presence reported of two Culex salisburiensis subspecies (Culex salisburiensis salisburiensis and Culex salisburiensis coursi). The subspecies Culex salisburiensis coursi, endemic [63], described only from a single specimen and known only at larval stages collected from rice fields in the eastern domain. Sympatric with Culex salisburiensis salisburiensis in the Lake Alaotra region on the eastern slope of the central highlands [63]. For Culex salisburiensis salisburiensis, pupal stage and eggs undescribed. Adult biology unknown. Not involved in the transmission of diseases.
d Eretmapodites subsimplicipes Edwards, 1914 Doucet, 1951 [66] Eggs undescribed. In Madagascar, presence reported only in 1951 by Doucet in Périnet forest in the eastern domain [66]. Adult and larval biology unknown. Rare species and not involved in disease transmission. In Comoros archipelago, anthropophilic and nocturnal species [24]. In Kenya, involved in transmission of Okola virus (OKOV) [1].

Genus Ficalbia Theobald, 1903
The genus Ficalbia belongs to the tribe Ficalbiini with the genus Mimomyia. The genus Ficalbia is represented by only eight species in the world [119]. Four species occur in the Afrotropical region. In Madagascar, this genus is represented by two species. Among them, Fi. circumtestacea was not reported as present on the island by WRBU [244]. Some specimens collected in other bioclimatic domains by Fontenille [85] could not be identified with confidence. This observation suggests that this genus is probably insufficiently studied in Madagascar.
d Ficalbia circumtestacea (Theobald, 1908) Grjebine, 1986 [110] Eggs undescribed. In Madagascar, reported at the larval stage, in the eastern domain, in the Andasibe-Mantadia and Manakara regions [110] and adult stage in Antsalova district of the western domain [15]. Adult biology unknown. Not involved in disease transmission.

Genus Hodgesia Theobald, 1903
The Hodgesia is represented by 11 species in the world [119]. Four species occur in Afrotropical region, mainly in central Africa. Adult females of African specimens are indistinguishable [158]. In Madagascar, presence was reported by Fontenille [85] who captured seven adults in human landing catches during daytime catches in April 1984 in the Mandena forest (Taolagnaro) [85]. The Malagasy specimens are still unidentified to date. This is probably the reason why Arim and WRBU did not rank this genus among the mosquito genera found in Madagascar [5,244]. Two females are currently stored in the laboratory of vector taxonomy of IRD Montpellier. Two other female specimens were caught by Didier Fontenille, in human landing catches, in May 1983, in the Antetezana forest, along the eastern coast between Toamasina and Foulpointe (Gilbert Le Goff, unpublished observation). Capture efforts for larval stages in swamp areas of the eastern coast could facilitate specimen collection and species identification. Hodgesia is poorly known, and rarely feeds on humans and is not known to be involved in medical or veterinary pathogen transmission.

Genus Lutzia Theobald, 1903
The genus Lutzia was formerly classified in the genus Culex and it was subdivided into three subgenera represented by eight species in the world [119]. Only one species belonging to the subgenus Metalutzia is present in the Afrotropical region and Madagascar. Eggs undescribed. Larval stages are predators of mosquitoassociated species and usually found in association with other species in many larval habitats. In Madagascar, larval habitats are canoes, marshes, canals [67], swamps [103], tires, puddles, flooded lowlands [207], and rice fields [191]. Occurs in the Sambirano area (Nosy Be, Nosy Komba) [90], in the western, eastern, and central domains [67,85]. Not involved in disease transmission. In Africa, involved in transmission of NTAV [17], WNV [200], and many other viruses in the Central African Republic (SINV, BABV, Bobia virus [BIAV], MOSV, and KAMV) [1].

Genus Mimomyia Theobald, 1903
The genera Mimomyia and Ficalbia belong to the Ficalbiini tribe. The genus Mimomyia includes 45 species subdivided into three subspecies: Etorleptiomyia (7 species), Ingramia (21 species), and Mimomyia (17 species) [119]. In Madagascar, 22 species were reported, and 17 of them are endemic. The phylogenetic relationship between the genus Mimomyia and other Culicidae genus remains uncertain, and the morphological data suggest affinity with the genera Ficalbia and Hodgesia [121]. The biology of genus Mimomyia remains poorly known. The species of this genus have no medical or veterinary importance in Madagascar, although some species were found naturally infected with arboviruses, particularly in Senegal [1].

Subgenus Etorleptiomyia Theobald, 1904
The subgenus Etorleptiomyia includes seven species, occurring mainly in the Ethiopian, eastern, and Australian regions. Two species were found in Madagascar, one being endemic to the island. The species of the subgenus Ertoleptiomyia breed in a wide variety of terrestrial water accumulations (marshes, ponds). d Mimomyia (Etorleptiomyia) martinei  Doucet, 1951 [66] Endemic. Only the adult female was described [66]. Its existence and its membership to one subgenus were repeatedly questioned [157]. Without being able to provide indisputable evidence, some authors suggest that the description of the female stage could correspond to that of Mi. (Ingramia) spinosa [110,157]. If this were the case, these two species would be synonymous and retain the name Mimomyia martinei [110]. Pending further information, this species must be regarded as valid and inventoried in the Malagasy subgenus Ertoleptiomyia [119]. This endemic species was collected only once and is known only from the type locality (Périnet area) [66], where two adults were captured from an outdoor resting area, in hollow bamboo. This species was not found since that time.

Subgenus Ingramia
d Mimomyia (Ingramia) bernardi (Doucet, 1950) Doucet, 1950 [65] Endemic. Eggs undescribed. Belongs to the complex of six species cited above. All developmental stages are morphologically close to those of Mi. aurata. Larval habitats are in axils of Ravenala and Pandanus fronds in forested area [65]. Occurs in a large part of the eastern domain [65]. Biology unknown. Not involved in the transmission of disease. d Mimomyia (Ingramia) beytouti  Doucet, 1951 [67] Endemic. Eggs undescribed. Belongs to the complex of six species cited above. Morphologically close to Mi. collessi in the larval stage. Shows significant differences in the pupal stage. Larval habitats are leaf axils of Ravenala [67,110]. Occurs in eastern domain [110]. Biology unknown. Not involved in disease transmission. d Mimomyia (Ingramia) brygooi Grjebine, 1986 Grjebine, 1986 [110] Endemic. Adult female and eggs undescribed. Because of the similarity between the larval stages within the subgenus Ingramia, larval capture data may refer to one of the following species: Mi. brygooi, Mi. levicastilloi, and Mi. longicornis, or to Mi. ramalai. The only adult stage known is the two males used in the original description, they were obtained from larval rearing. Larval habitats are leaf axils of Pandanus and Ravenala [85]. Occurs in the eastern domain [110]. Adult biology unknown. Not involved in disease transmission.
d Mimomyia (Ingramia) collessi Grjebine, 1986 Grjebine, 1986 [110] Endemic. Eggs undescribed. Belongs to the complex of six species cited above. Morphologically close to Mi. beytouti. Occurs mainly in the eastern domain [110]. Adult biology unknown. Not involved in disease transmission.  Madagascar and Mi. grjebinei in the Comoros archipelago. Morphological variations observed from specimens collected in the Vohipeno region, allowing us to assume the presence of a complex of species. Larval habitats are mainly the leaf axils of Typhonodorum sp. and exceptionally the axils of fronds of Ravenala. Occurs in the eastern domain and locally on the west coast of Madagascar (Nosy Be, Morondava, and Mahajanga regions) [103]. Adult biology unknown. Not involved in disease transmission.

Subgenus Mimomyia Theobald, 1903
The subgenus Mimomyia includes 21 species, widely distributed throughout the Ethiopian and Oriental regions and extends to northern Australia and the South Pacific. The four Malagasy species have a wide distribution on the African mainland. The report of Mi. lacustris Edwards and Mi. pallida Edwards as present in Madagascar in the Arim dataset [5] is doubtful as information on collection areas is not available. De facto this information was treated as an error. The species of the subgenus Mimomyia develop in a wide variety of terrestrial water accumulations (ponds, marshes, ponds, and riverbanks). d Mimomyia (Mimomyia) hispida (Theobald, 1910) Doucet, 1951 [66] Eggs undescribed. In Madagascar, larval habitats are ponds and marshes containing abundant aquatic vegetation [110], muddy swamp water and rice fields with low vegetation [67]. Occurs in the central and eastern domains [65,67,85,110]. Adult biology unknown. In Africa, caught in human biting catches in West Africa [155] and feeds mainly on amphibians and cattle in Kenya [13]. Involved in transmission of BABV, BAGV, and WNV [222]. d Mimomyia (Mimomyia) mimomyiaformis (Newstead, 1907)  Eggs undescribed. In Madagascar, larval habitats are stagnant or slow moving water, with aquatic vegetation (swamps, irrigation canals, and rivers) within forested areas [66]. Occurs in the eastern domain, western coastal plains, in the Mahajanga region [110]. Adult biology unknown. In Africa, captured in human biting catches [155]. Not involved in disease transmission.
d Mimomyia (Mimomyia) plumosa  Doucet, 1951 [67] Eggs undescribed. In Madagascar, larval habitats are forest ponds with vegetation [110]. Occurs in the eastern domain (Périnet and Vangaindrano) and the Sambirano area. Adult biology unknown. Not involved in disease transmission. Adult biology unknown. In Africa, involved in transmission of Bunyaviruses of the Bwamba Group, a non-pathogenic virus to humans [1].
Grjebine, 1956 Eggs undescribed. In Madagascar, larval habitats are terrestrial water accumulations, invariably associated with aquatic plants, indispensable for breathing larvae. Occurs throughout the eastern domain, from the eastern margin of the central highlands (Moramanga Périnet) to the coastal lagoons of the south-eastern domain (Manakara and Vangaindrano regions). Also reported locally in the western domain (Mahajanga region). In Africa, feeds mainly on amphibians and occasionally on humans [13]. Involved in transmission of WNV, BABV, and BAGV [1].

Genus Orthopodomyia Theobald, 1904
This genus is represented by 36 [85,182,207], and ovitraps [85]. Occurs in the eastern and central domains and seems to be frequent at lower altitudes (below 800-900 m): from the sea to the eastern margins of the central highlands [27,85]. Adult biology unknown. Brunhes & Hervy, 1995 [27] Endemic. Egg and adult stages undescribed. Larval habitats are tree holes and cut bamboo. Only occurs in Antongil Bay, in the eastern domain [27]. d Orthopodomyia vernoni van Someren, 1949 [229] van Someren, 1949 [229] Endemic. Eggs undescribed. Larval habitats are many phytotelmata but sometimes artificial containers (tin cans and metal cans) [27,85]. Occurs in the western and southern domains of Madagascar, and can reach areas bordering other bioclimatic domains, at lower altitude (below 1000 m).

Genus Toxorhynchites Theobald, 1901
This genus includes four subgenera represented by 89 species in the world [119]. Six species occur in Madagascar. This study did not include Tx. brevipalpis that is reported to be present on the island by WRBU [244]. The Malagasy species belong only to the subgenus Afrorhynchus which dominates on the African mainland, and within the Group Pauliani which is endemic to Madagascar [190]. The external morphology of this group is homogeneous. Only male genitalia morphology allows differentiation of these species. The adult stages of Toxorhynchites are phytophagous and are not involved in transmission of pathogens. The larval stages are predators feeding on larval stages of other mosquito species. Its host preference allows us to consider Toxorhynchites mosquitoes as a biological control agent of vector mosquitoes [190]. In Madagascar, Toxorhynchites larva develops generally in many phytotelmata: Typhonodorum, Ravenala, Pandanus, Nepenthes madagascarensis, Colocasia, bamboo, and fruit shells [111]. Most Malagasy Toxorhynchites species occur in the eastern biogeographic domain [67,190]. Larval stages of this genus were collected by Rodhain et al. [195] in the Mahajanga area, of the western domain, but remained unidentified. The adult biology of Toxorhynchites of Madagascar is unknown and the Malagasy species has no medical or veterinary importance. d Toxorhynchites (Afrorhynchus) brunhesi Ribeiro, 2004 [190] Ribeiro, 2004 [190] Endemic. Eggs undescribed. The holotype male was collected in the Moramanga district located in the eastern domain, on the eastern margin of the central highlands.

Genus Uranotaenia Lynch Arribálzaga, 1891
Genus Uranotaenia is the only genus of Culicidae belonging to the tribe Uranotaeniini. This genus is subdivided into two subgenera and includes 267 species in the world: the subgenus Pseudoficalbia (146 species) and the subgenus Uranotaenia (121 species) [119]. The genus Uranotaenia occurs on all continents, with the exception of the Pacific Ocean islands and Antarctica. This genus is particularly well represented in the Afrotropical and Oriental regions. In Madagascar, a comprehensive and complete systematic revision was carried out on the genus Uranotaenia [51]. With this revision, the genus Uranotaenia is the best represented and probably the best known genus, regarding the number of species in Madagascar. Among 73 Malagasy species belonging to this genus, 65 species are endemic, and four species occur only in Madagascar and in the Comoros archipelago. Although we have little information on the host preferences and behavior of adults, we know that these mosquitoes prefer to feed on cold-blooded animals (reptiles, amphibians), and nevertheless are probably involved in disease transmission.
2.14.1 Subgenus Pseudoficalbia Theobald, 1912 [221] All Malagasy species (n = 52) of this subgenus are endemic. This study did not include Ur. comorensis, Ur. fusca, Ur. mashonaensis, Ur. nepenthes, Ur. ornate, Ur. pandani, and Ur. shillitonis that were reported to be present on the island by WRBU [244]. The larval habitats are always associated with small breeding sites and phytotelmata.  [67], puddles on tree trunks, banana leaves, and cut bamboo [51]. Occurs in the eastern domain [51], from sea level up to 1000 m. Endemic. Eggs undescribed. Belongs to section Nigripes. Morphologically close to Ur. ornata Theobald, which is absent from Madagascar. Larval habitats are dead leaves of Ravenala [67], leaf axils of Typhonodorum and Pandanus, gutters, mushrooms horns, rock holes, fallen tree trunks, puddles, and streams [51]. Occurs in the central and eastern domains. Abundant from eastern seaboard up to 1200 m asl [51].  [103] and Typhonodorum sp., banana leaves, bamboo, and mushroom caps [51]. Carnivorous larvae, which feed on mosquito larvae of the same or other species. Rare species, but widely distributed throughout Montagne d'Ambre in the northern domain, Mandraka in Antananarivo province, Ikongo of Fianarantsoa province in the central domain, and Andasibe forest in the eastern domain [51]. Endemic. Eggs undescribed. Belongs to section Nigripes. Larval habitats are cut bamboo and tree holes [51] and axils of Typhonodorum in Mayotte [145]. This species is present in Madagascar and Mayotte [145]. In Madagascar, occurs in the eastern domain and in the Montagne d'Ambre of the northern domain [51]. d Uranotaenia (Pseudoficalbia) lavieri Doucet, 1950 [65] Doucet, 1950 [65] Endemic. Eggs and adult stages undescribed. Belongs to section Annulata (representative of the nominal group). Larval habitats are cut bamboo [66] and tree holes [51]. Occurs in the Sambirano area (Nosy Be, Nosy Komba) [90], in rainforests of the eastern [66] and northern domains [51].  [14] Endemic. Only the larval stages were described [14]. Belongs to section Spinosa [51]. Larval habitats are leaf axils of Pandanus along the beachfront. Collected only once and known only from the type locality (Ambila-Lemaitso, Toamasina province of the eastern domain). Endemic. Only the larval stages were described [64]. Belongs to section Nigripes. Formerly synonymized with Ur. nepenthes and Ur. pandani which are endemic to the Seychelles archipelago [124,156]. Morphologically close to Ur. ornate which is absent from Madagascar [64]. Only type series of larval stages were caught to date. Larval habitats are dried leaf rachis of Neodypsis on the ground. Collected only once. Known only from the type locality (Ambohiby, Tsiroanomandidy region in the central domain) [64].  [14] Endemic. Only the larval stages were described [14]. Belongs to section Annulata. Larval habitats are cut trunks of Ravenala madagascariensis containing brown water and rotting vegetable matter. Its larval stages are usually found in association with Uranotaenia haddowi, Ur. manakaraensis, Ur. albimanus, and Ur. breviseta. Collected only once and known only from the type locality (Ifaho, Manakara region of the eastern domain).  [65], palm tree forming a gutter, coconuts, and the leaf axils of Pandanus [51]. Occurs in high altitude forests of the eastern and northern domains [51], between 700 and 1700 m asl.

Subgenus Uranotaenia Lynch Arribálzaga, 1891
This subgenus includes 21 species in Madagascar. Among them, Ur. alba was reported to be absent from the island by WRBU [244]. Fourteen species are endemic to the island, three are found in Madagascar  Endemic. Eggs and adult male remain to be described. Belongs to section and group Anopheloides. Larval habitats are tree holes with water containing organic matter. Collected only once and known only from the type locality (Masoala peninsula, in the eastern domain) [51]. d Uranotaenia (Uranotaenia) andavakae Doucet, 1950 [65] Doucet, 1950 [65] Endemic to Madagascar and the Comoros archipelago (Mayotte) [65]. Eggs undescribed. Belongs to section Anopheloides (Neireti Group). Larval habitats are cool and clear waters such as fountains, ditches with sphagnum, puddles, flooded meadows, rice fields, forest ponds, rock holes, and trickle of water along a stream [51,65]. Occurs in the eastern domain, in mountainous areas of the western, eastern, and central domains [51], and can reach about 1800 m asl. d Uranotaenia (Uranotaenia) anopheloides Brunhes & Razafindrasolo, 1975 [31] Brunhes & Razafindrasolo, 1975 [31] Endemic to Madagascar and to the Comoros archipelago (Mayotte) [31]. Eggs undescribed. Belongs to section Anopheloides. Larval stages are characterized by their bodies floating parallel to the surface of the water like those of Anopheles. Larval habitats are tree holes (mango tree, kapok tree, and Ravenala) [51]. Occurs in the western, central, and eastern domains, seems more abundant in the warmer regions of the western domain. Eggs are resistant to desiccation [51].  Theobald, 1904 Eggs undescribed. Belongs to section Alba (Balfouri Group). Morphologically close to Uranotaenia hebrardi. In Madagascar, larval habitats are canals containing iron hydroxide, rice fields, pools, marshes [63], grassy holes [103], muddy water under forest cover, swamps, and ponds [51]. Occurs in the Sambirano area (Nosy Be), in the western, eastern, and central domains. This species is found in Madagascar from sea level up to 1700 m asl [51]. Adult biology unknown. In Africa, feeds mainly on amphibians and cattle in Kenya [13]. Doucet, 1949 [64] Endemic. Only the larval stages were described [64]. Belongs to section and group Dumonti. Larval habitats are water holes and reed beds. Collected only once and known only from the type locality (Ambohiby mountain stream, at 1633 m asl, in the Tsiroanomandidy region of the central domain) [51,64]. Grjebine, 1953 [103] Endemic. Eggs undescribed. Originally described under the name of Uranotaenia chorleyi var. hamoni [103] and formally elevated to species rank by White [238]. Belongs to section Chorleyi (Hamoni Group). Larval habitats are forest ponds, rivers, holes containing clear and fresh water, rice fields surrounded by forest, and forest streams. Occurs in the central, eastern, and western domains and was mainly caught in mountainous areas, above 800 m asl [51]. d Uranotaenia (Uranotaenia) hebrardi da Cunha Ramos & Brunhes, 2004 [51] da Cunha Ramos & Brunhes, 2004 [51] Endemic. Belongs to section Alba. Morphologically close to Uranotaenia balfouri (Balfouri Group). Larval habitats are ponds, rice fields, and marshes. Occurs in the eastern, western, and central domains. Caught only as larvae and on five capture occasions in the Amborompotsy [191] and Manakara areas of Fianarantsoa province, the Marvoay area of Mahajanga province, and the Ambodimanga and Moramanga areas of Toamasina province [51]  Endemic. Eggs undescribed. This species belongs to section Chorleyi (Hamoni Group). Larval habitats are forest puddles, streams containing sphagnum, fresh and acidic water, cattle hoof prints, ditches, and grassy swamps. Occurs in the central, eastern, and southern domains [51] and is distributed between 450 m (Beraketa, Toliary province) and 1800 m asl (Ankaratra massif). d Uranotaenia (Uranotaenia) neireti Edwards, 1920 [75] Edwards, 1920 [75] Endemic. Eggs undescribed. Belongs to Anopheloides section. Larval habitats are hoof prints, lakes, and swamps areas. Present in the central and eastern domains and always between 900 and 2000 m asl [51].  [51] Endemic. Eggs, pupal and adult stages undescribed. Belongs to Chorleyi section (Hamoni Group). Larval habitats are slow-flowing water of small forest streams of primary rainforest. Collected only once and known only from the type locality (Lakato area, above 1000 m asl of the eastern domain) [51].

Discussion
The main remarkable characteristic of Malagasy mosquito fauna is the high biodiversity with 138 (of 235) endemic species (58.7%). This pattern is not only specific to mosquitoes since Madagascar has one of the highest rates of endemicity in the world, due to its insularity with long geographic isolation since the migration of the Indo-Malagasy subcontinent 155-120 million years ago. The isolation of the island dates from the Upper Cretaceous (83 million years). Paulian [173] highlighted this endemism rate in many vertebrate and invertebrate groups. The endemism rate is about 92-100% for terrestrial animals and 52-60% for flying animals (bats and birds) [100]. As an example, 75% (97/130) of Drosophila species [135], all of the 40 species of scorpions [149], and 91% (379/418) of ant species [84] are endemic to Madagascar. Among other hematophagous arthropods other than mosquitoes, the endemism rate is 92% (11/12) (Tables 1 and 2). This main difference is primarily due to the inclusion of an additional collection of Uranotaenia [14,51], Toxorhynchites [190], Aedeomyia [33], and Aedes belonging to the subgenus Neomelaniconion [143]. However, this number is moderately lower than that given in Internet references such as WRBU (253 species) [244] and Arim (245 species) [5], which include an enlarged area of several islands in the Mozambique Channel.
The report of the genus Culiseta (Culiseta longiareolata (Macquart)) as present in Madagascar in the Arim dataset [5] is doubtful as information on collection areas is not available. To date, there is no conclusive evidence of the presence of this tropical species in Madagascar and this information was treated as an error.
Taking into account that 3546 mosquito species are currently recognized in the world [119], 6.6% of them are present in Madagascar. In addition, 28% of the 804 known species in the Ethiopian zoogeographical area are present in Madagascar [144]. Moreover, despite decades of research, there is no doubt that new species remain to be discovered and that endemism could be higher than 60%. Indeed, some species have been reported only once (32 endemic species) and sometimes described from a single specimen with the possibility of misidentification (7 endemic species). In addition, species complexes exist, and several new species could occur without having been identified and described. On the other hand, due to deforestation, anthropogenization, and degradation of natural (and poorly explored) biotopes, several mosquito species have probably already disappeared.  Endemic mosquitoes restricted to Madagascar. § Endemic mosquitoes in Madagascar and the Comoros archipelago. # Species described and/or clearly identified. * The species Culex thalassius, mentioned with doubt in Madagascar by Knight and Stone [132] and listed in Madagascar by the site WRBU (www.mosquitocatalog.org/, August 2014) [244], has not been recorded in this table; similarly the species belonging to the Rima group surveyed but remaining unidentified by Rhodain et al. [195] have not been recorded. Cx. pipiens and Cx. quinquefasciatus were counted as two separate species following the conclusion of Harbach [118]. ** The four species of mosquitoes known only in mountains areas (altitude >1500 m.) are usually associated with the central bioclimatic domain.

Endemism rate of Malagasy mosquitoes
Mosquitoes already existed before separation of Madagascar and the Indian subcontinent from Africa, 156 million years ago, during the Upper Jurassic. The phylogeographic origins of sylvatic Malagasy mosquito species are not easy to decipher, and we do not yet know when and how founder species reached Madagascar. Differentiation occurred in nearly all the country's biogeographic domains. It was not observed in the southern domain because no strictly southern endemic species is known. However, there are Malagasy endemic mosquitoes in all biogeographic domains, including the south where more than 33% of species are endemic. The eastern domain, where the relict rainforest is still present, has the highest number of mosquito species (n = 190) and level of regional endemism (67%). Several species are microendemics, only present (or described) in a single biotope, often in forested areas (e.g., Ae. ambreensis, Ae. albodorsalis, Ae. mathioti, An. roubaudi, Mi. longicornis, Mi. mattinglyi, Mi. stellata, and Or. ankaratrensis, all species belonging to the genus Toxorhynchites, and eight species belonging to the genus Uranotaenia).
There is no endemic genus in Madagascar. Mosquito species endemism occurs in 9 of the 13 genera of mosquito, with variation of the endemism rate from 27% to 100%, according to the genus.
The genus Aedeomyia has 66% (2/3) endemism. This genus is distributed in the Australasian, Oriental, African, and Neotropical regions [117]. The known species would appear to be from a primitive population that developed during the fragmentation of Gondwana 100 million years ago [33].
The genus Anopheles has a cosmopolitan distribution [117]. All non-endemic Anopheles species are known from broad regions of Africa. The Anopheles fauna of Madagascar is typically Ethiopian and the occurrence of high endemism (46%; 12/26) supports evidence that its separation from the mainland must have taken place a long time ago [70].
The genus Coquillettidia is distributed in the Old World and Neotropical region [117]. In Madagascar, this genus has 66% (2/3) endemism.
For the genus Culex, the majority of non-endemic species are of African origin and only the subgenus Culex contains a few species occurring outside the African region (species belonging to the groups Univittatus and Pipiens). Culex ?The locality where this species was collected was not well specified. O: ornitophile, Z: zoophile, (ruminants), A: anthropophilic, GF: general feeder (Z, O, A). A: terrestrial water accumulation associated with agricultural activity, N: natural larval habitats associated with terrestrial habitat, P: phytotelmata (bamboo, tree hole, leaf axils), T: artificial containers (small plastic receptacles, tires, drums, cans, sewage) water accumulation (temporary or not). species, endemic to Madagascar, belong to five subgenera. Today, there are less than 24% endemic Culex species in Madagascar and the best known bioclimatic area for this genus is probably the central domain. In contrast, no endemic Culex species are known in the western, northern, and southern domains. There is no doubt that endemicity should be higher in this genus. Already recorded species belong to species complexes, and a revision of genus Culex would likely modify the species inventory in the future.
The genus Mimomyia has a high endemism rate (77%; 17/ 22). The subgenus Ingramia is well represented and all species belonging to this subgenus are endemic to Madagascar. The subgenus Etorleptiomyia is found from Africa to the Oriental and South Pacific regions. The subgenus Mimomyia is widely distributed on the African mainland and extends in the east to New Guinea, north-eastern Australia, and the South Pacific [110].
The genus Orthopodomyia is distributed throughout the Afrotropical, Nearctic, Neotropical, Palaearctic, and Oriental regions [117]. However, all Malagasy species of Orthopodomyia are endemic (8/8) and belong only to the Vernoni Group. Because of their morphological homogeneity, these species are probably derived from a single ancestral species [27].
The genus Toxorhynchites with four subgenera is distributed throughout the Afrotropical, Australasian, Neotropical, eastern Palaearctic, and Oriental regions [117]. All Malagasy species of Toxorhynchites are endemic (6/6) and belong only to the subgenus Afrorhynchus. This subgenus evolved here after the separation of Madagascar from the African mainland during the Tertiary period [190].
The genus Uranotaenia is represented by Pseudoficalbia and Uranotaenia subgenera. All species of subgenus Pseudoficalbia are endemic to Madagascar and affinity between Malagasy, African, and Indo-Malaysian species is observed [51]. These authors suggested that the subgenus Pseudoficalbia seems to have appeared in the Madagascar-Indian plate boundary during the Upper Cretaceous. Most species belonging to the subgenus Uranotaenia exhibit a marked affinity with African and Indian species and high endemic components are observed in different sections. Specific radiation seems to have appeared before Gondwana fragmentation [51].
There are no endemic species within the genera Eretmapodites, Ficalbia, Lutzia, and Mansonia. It is important to note that these genera are represented at most by four species. The genus Eretmapodites occurs only in the Afrotropical region, the two Ficalbia species have Afrotropical origin [110]; Lutzia is present in the Neotropical, Asian, Australasian, and African regions and Mansonia has worldwide distribution [117].

Bio-indicator species
According to Leclercq [139], a good bio-indicator should have a specific exigency allowing a link between its presence/absence and environmental particularities. Mosquitoes fit into this definition [62].

Urban species
Mosquito species which grow in peri-and para-domestic breeding sites are characteristic of urban areas [240]. Mosquito species growing in several artificial and polluted breeding sites exist in Malagasy urban habitats. It is not surprising to find a high abundance of Cx. quinquefasciatus and Ae. albopictus in Antananarivo, the capital and most urbanized city of Madagascar [85,176] where discarded bottles represent 64% of household waste [178] and the rice fields of the bottom-land are converted into polluted watercress fields [9].
The same situation is observed in Toamasina, the second largest city in Madagascar, where Cx. quinquefasciatus and Ae. albopictus have colonized several peri-and para-domestic breeding sites [85]. These species are already considered an urban species in the tropical and south-east Asian regions [240] and in the Mascareignes Archipelago [208].
Ancient entomological data have suggested the presence of Ae. aegypti as a bio-indicator of urban areas in Madagascar [85]. This is however probably not true because its distribution is actually limited to smaller anthropized and/or forested areas [176].
However, An. squamosus and An. coustani are the most abundant Anopheles species in the peripheral areas of the capital city Antananarivo [85], and in the suburban areas of the eastern and central domains where rice fields are abundant in and around cities [85,210].
A study performed in the Morondava region showed the abundance and concomitant presence of Ma. uniformis and Ma. africana in the cities of the western domain [85]. Indeed, all western plains, including within and on the outskirts of cities, are dotted with numerous swamps and ponds [36] which are typical habitats of these two species. These species constitute a major nuisance in some cities of south-east Asia and on the African mainland [240].

Forested and rural species
In total, 78 species are restricted to the rainforest biotope and can be considered as bio-indicator of this habitat. Mosquito species belonging to Anopheles of the Neomyzomyia series [108,112], Aedeomyia (Aedeomyia) madagascarica [33], Aedes (Aedimorphus) albodorsalis [86], Aedes (Diceromyia) sylvaticus [26], and endemic species of Mimomyia (Ingramia) [66,110] were collected only in rainforest habitats. Unlike other follow-up studies [85,227], Cx. pipiens was abundant only in the Anorana rainforest in the central domain [210]. Because of its abundance, Cx. pipiens could be considered as a bio-indicator of the rainforest habitat in the central domain. This finding raises important questions regarding the differentiation history or behavior modification of Cx. pipiens in relation to urban and suburban habitats [123,180,227]. Little information is available about its biology in Madagascar, even though a much lower proportion of this species was reported in the Andasibe rainforest of the eastern domain [85].
Several studies have reported that mosquito species of Ficalbia, Toxorhynchites, Mimomyia (Ingramia), and large numbers of Uranotaenia genera (especially subgenus Pseudoficalbia) occur only in the eastern region. These species lay their eggs in phytothelm breeding sites (Typhonodorum, Ravenala, Pandanus, and Nepenthes madagascarensis) which are characteristic of the eastern domain [35,122,136]. These observations confirm that these mosquito genera could be considered as good indicators of the eastern domain.
Eretmapodites quinquevittatus is considered as indicator species of degraded areas in all domains, with the exception of the central domain where this species is rare [85]. Four species Aedes brygooi, Ae. coulangesi, Ae. albiradius, and Ae. aegypti are confined to the driest forest of the western domain. In this domain, Ae. fryeri and Ae. cartroni species seem to be related to the presence of mangrove [182]. Anopheles merus was also found in dry regions in the western domain and far south-east of Madagascar and is probably associated with salt-water.
Few species are strongly represented in the warmer and dryer areas of the western domain, even though they occur in others domains: Ae. coulangesi, Ae. durbanensis, Ae. brygooi, Ae. albocephalus, Ae. tiptoni, Ae. cartroni, Cx. tritaeniorhynchus, and Ur. anopheloides are abundant in this area [31,85,182,194]. Similarly, Ae. scatophagoides is present only in the semi-arid Androy region, in the southern domain [182], while Ae. ambreensis occurs only in the northern domain [85]. These observations allow us to consider these species bio-indicators of their corresponding geographical domains.

Conclusion
In Madagascar, up to February 2016, 235 mosquito species belonging to 14 genera have been reported, with a high level of endemism. There is no doubt that this inventory will change with new species. The taxonomic status of species complexes described from only one specimen or from one stage should be deciphered. Further description of species belonging especially to the genus Hodgesia, and the subgenus Culex (Eumelanomyia), should be carried out to complete this inventory.