Francis Schaffner

17, rue du 152e R.I. – 68700 Steinbach, France

There are few major works on French mosquitoes. Those that exist are species catalogues (Séguy, 1925a; Hedeen, 1958, 1959; Callot & Rioux, 1965), sometimes including distribution and bibliography (Rageau et al., 1970; Moussiegt, 1986). A certain number of species listed in the literature cannot be considered as truly belonging to the French fauna. Furthermore, during the last few years, new species have been described, and taxonomic changes have occurred (synonyms, new sub-genera, etc.). The purpose of this paper is to update the list of species collected in France and Corsica.

Genus Anopheles Meigen 1818

Sub-genus Anopheles Meigen 1818

1 algeriensis Theobald 1903

claviger sensu lato:

2 claviger (Meigen 1804)

3* petragnanii Del Vecchio 1939

4 hyrcanus (Pallas 1771)

maculipennis sensu lato:

5 atroparvus Van Thiel 1927

6** labranchiae Falleroni 1926

7 maculipennis s.s. Meigen 1818

8 melanoon Hackett 1934

9 messeae Falleroni 1926

10** sacharovi Favre 1903

11 subalpinus Hackett & Lewis 1935

12** marteri Sénevet & Prunelle 1927

13 plumbeus Stephens 1828

Sub-genus Cellia Theobald 1902

Tribe Aedini

Genus Aedes Meigen 1818

Sub-genus Aedes Meigen 1818

15 cinereus Meigen 1818

16 esoensis rossicus Dolbeskin, Gorickaja & Mitrofanova 1930

17 geminus Peus 1970

Sub-genus Aedimorphus Theobald 1903

18 vexans vexans (Meigen 1830)

19 vittatus (Bigot 1861)

Sub-genus Finlaya Theobald 1903

20 geniculatus (Olivier 1791)

Sub-genus Ochlerotatus Lynch Arribalzaga 1891

Intrudens Group

21 diantaeus Howard, Dyar & Knab 1912

22 pullatus (Coquillett 1904)

Ochlerotatus Group

Annulipes Sub-group

23 annulipes (Meigen 1830)

24 cantans (Meigen 1818)

25 flavescens (Müller 1764)

26 surcoufi Theobald 1912

Communis Sub-group

27 cataphylla Dyar 1916

28 communis (De Geer 1776)

detritus (Haliday 1833)

29 sp. A Pasteur et al. 1977

30 sp. B Pasteur et al. 1977

31 nigrinus (Eckstein 1918)

32 punctor (Kirby 1837)

33 sticticus (Meigen 1838)

Dorsalis Sub-group

34 berlandi Séguy 1921

caspius caspius (Pallas 1771)

35 sp. A Cianchi et al. 1980

36 dorsalis (Meigen 1830)

37 mariae (Sergent & Sergent 1903)

38 pulcritarsis pulcritarsis (Rondani 1872)

Sub-genus Rusticoidus Shevchenko & Prudkina 1973

39 refiki Medschid 1928

40 rusticus (Rossi 1790)

Sub-genus Stegomyia

41*** aegypti (Linnaeus 1762)

Tribe Culicini

Genus Culex Linnaeus 1758

Sub-genus Barraudius Edwards 1921

42 modestus modestus Ficalbi 1889

Sub-genus Culex Linnaeus 1758

43** brumpti Galliard 1931

44 mimeticus Noe 1899

45 pipiens pipiens Linnaeus 1758 inc. form molestus Forskål 1775

46 theileri Theobald, 1903

47 torrentium Martini 1925

Sub-genus Maillotia Theobald 1907

48 hortensis hortensis Ficalbi 1889

Sub-genus Neoculex Dyar 1905

49 impudicus Ficalbi 1890

50 martinii Medschid 1930

51 territans Walker 1856

Tribe Culisetini

Genus Culiseta Felt, 1904

Sub-genus Allotheobaldia Brolemann 1919

52 longiareolata (Macquart 1838)

Sub-genus Culicella Felt 1904

53 fumipennis (Stephens 1825)

54 litorea (Shute 1928)

55 morsitans (Theobald 1901)

Sub-genus Culiseta Felt 1904

56 alaskaensis alaskaensis (Ludlow 1906)

57 annulata (Schrank 1776)

58 glaphyroptera (Schiner 1864)

59 subochrea (Edwards 1921)

Tribe Mansoniini

Genus Coquillettidia Dyar 1905

Sub-genus Coquillettidia Dyar 1905

60 buxtoni (Edwards 1923)

61 richiardii (Ficalbi 1889)

Tribe Orthopodomyiini

Genus Orthopodomyia Theobald 1904

62 pulcripalpis (Rondani 1872)

Tribe Uranotaeniini

Genus Uranotaenia Lynch Arribalzaga 1891

Sub-genus Pseudoficalbia Theobald 1912

63 unguiculata unguiculata Edwards 1913

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* this taxon should be petragnanii and not petragnani following the Latin rules of declension for specific names. The species was named by Del Vecchio in honour of Pr. Petragnani

** species present in Corsica only

*** sporadic species, often introduced via ports, not regarded as endemic in view of the rarity of observations (Rageau et al., 1970)

The organisation of this list takes into account the systematic classification used in the reference work: "A catalog of the mosquitoes of the world" and its additions (Knight & Stone, 1977; Knight, 1978; Ward, 1984; Gaffigan & Ward, 1985; Ward, 1992). Taxa are presented alphabetically within each taxonomic grouping.

Harbach (1994) has recently revised the internal classification of the genus Anopheles. Considering the small number of species, it was decided not to represent the sections, series, groups and sub-groups. On the other hand, certain relationships are deliberately highlighted by reference to "species complexes".

Considering that the sub-genus Ochlerotatus represents by far the greatest number of species, and that the morphological differences are sometimes important, it was decided to arrange the species according to the groups and sub-groups used by Mohrig (1969).

The sub-genus Rusticoidus was described by Shevchenko & Prudkina (1973) following a study of the male genitalia in the genus Aedes. They proposed Ae. refiki as haplotype, and included in the sub-genus the related species previously incorporated in the rusticus group. Even though the catalogue of mosquitoes (Knight & Stone, 1977 and its additions) classifies Ae. refiki as being the only European species in this sub-genus, we share the opinion of other authors (Shevchenko & Prudkina, 1973; Encinas Grandes, 1982) that Ae. rusticus is also a member of that sub-genus.

We have made several modifications to the last list of French Culicidae published by Callot & Rioux (1965). Their list recognised 62 taxa, including Ae. vexans which was accidentally omitted from the publication due to a typographical error.

An. subalpinus is a taxon that has recently been elevated to species status (Bullini et al., 1980; Cianchi et al., 1987; Ribeiro et al., 1988; Ward, 1992). Being morphologically very close to An. melanoon, the two species are often mistaken in the literature. An. subalpinus was first recorded in the Languedoc in 1957 (Rioux & Ruffié, 1957) and appears to be abundant in the south of France (Salieres et al., 1978).

Ae. esoensis rossicus is the most recently discovered species in France (Schaffner & Pfirsch, 1995). It has been recorded only in the north of Alsace, where it is relatively rare. However, large populations of this species occur in the flood plains of the Rhine in Germany (Becker & Ludwig, 1981).

Ae. geminus is a species related to Ae. cinereus, as described by Peus (1970). Since its discovery in France (Roman & Pichot, 1976), this species has been only rarely recorded, probably due to the difficulties posed by its identification (the need to distinguish adult males). It seems, however, that Ae. geminus is more abundant than Ae. cinereus (Schaffner, 1992; Schaffner & Pfirsch, 1995). Besides, it seems likely than a certain number of earlier records of Ae. cinereus are actually attributable to Ae.geminus. The distributions of both these taxa are still to be established.

Sibling species have been differentiated within two taxa of the genus Aedes (Ae. detritus and Ae. caspius caspius) using new genetic identification techniques, in particular isoenzymes studies. In addition to the demonstrated reproductive isolation, ecological, ethological and distributional differentiation has been added, thus confirming the validity of the two Ae. detritus species.

• Ae. detritus (both species "A" and "B"of Pasteur et al. (1977)), were recorded in Camargue (Pasteur et al., 1977). A geographical segregation has been found to exist along a north-south gradient, with "A" less common in the north (Atlantic coastline: Brutus et al., 1994). Autogeny, frequent in "A", is exceptional in "B"; species "A" develops only in hyper-saline larval sites; and the possibility of finding some morphological differentiation characters cannot be excluded (Agoulon, 1996).

• Ae. caspius caspius: only species "A" of Cianchi et al., 1980 has been identified in France, so far, on the Atlantic coastline (Agoulon, 1996).

Arnaud et al. (1976) resurrected the taxon Aedes surcoufi from synonymy with Aedes (Ochlerotatus) excrucians (Walker 1856) and considered it to be the sole representative of the "excrucians" complex in France. Accordingly, the name Ae. excrucians is replaced by Ae. surcoufi in the list of species. Certain authors (Dahl, C. & Nielsen, B.O. personal communication) consider, however, that the possibility of finding Ae. excrucians in the Alps should not be excluded. In the absence of any morphological analysis of alpine populations, we hold the opinion of Arnaud et al. (1976).

Aedes (Ochlerotatus) leucomelas (Meigen 1804) is removed from the list since it is not found in France. The only reports noted the occurrence of aggressive Ae. leucolemas females in the Jura (Roman, 1944; Roman & Morel, 1946), but emphasised the difficulties of distinguishing this species from Aedes cataphylla. A few years later the same specimens were classified as Ae. cataphylla (Roman, 1958). Hedeen (1959) and Rageau et al. (1970) subsequently suggested that the species probably occurred in France, considering its extensive Palaearctic range. However, as subsequent studies failed to locate the species in France, it seems reasonable to exclude it from the list. Nevertheless, Ae. leucomelas occurs in Germany, a few kilometres from the French border (Oberrheingebiet) (Becker & Ludwig, 1981) and therefore could potentially be encountered in the north-east of France.

Culex (Culex) laticinctus Edwards 1913 has been recorded in three different places: Hyères, in Var (Séguy, 1920a), Fontainebleau, in Seine-et-Marne (Séguy, 1921) and Lucciana, in Corsica (Galliard, 1927). All other references to this species in France actually relate to one or more of these records. We consider that the presence of a species can only be authenticated after larvae, adult males and adult females have been observed, together with the possibility of studying preserved specimens. None of the three French records matches these requirements. In addition to this is the fact that the species distribution is disjunct (Fontainebleau). We therefore do not consider this species to be present in France, at least until there are further records and more studies have been made.

Culex (Culex) univittatus Theobald 1901 was recorded in Corsica (Galliard, 1927) on the basis of larvae attributed to Culex perexiguus Theobald, 1903. Subsequently, the same author identified adult females from two other Corsican sites, but mentioned that there was no certainty of these specimens being Cx. univittatus (= perexiguus) as he had failed to obtain adults from the previously mentioned larvae (Galliard, 1928). Later, the discovery in 1931 of similar larvae from one of the 1927 sites, and the obtaining of adults (2 males and 2 females) allowed the author to describe the specimens as belonging to the new taxon Culex brumpti Galliard 1931. As no precise detail was given for the females from 1928, they remained classified as Cx. univittatus. However, even though various authors have retained them in their faunistic lists (Edwards, 1928; Séguy, 1932; Callot & Rioux, 1965; Moussiegt, 1986), others have removed the taxon (Lanzalavi, 1965; Rageau et al., 1970). In 1966, there was another mention of the species in an administrative document from Corsica (Ségui, 1966). The species was encountered three times, with a total of 4 larvae (no preserved specimens). Even though Cx. univittatus occurs in the Iberian Peninsula and in Italy (Dahl & White, 1978), we prefer not to include it in the French species list, due to the uncertainties in the literature.

Apart from the list by Callot & Rioux (1965) and species recently reported, the literature contains records of five other taxa in France. Four of these are invalid and one is considered doubtful.

Aedes (Finlaya) echinus (Edwards 1920) was reported for the first time (Séguy, 1920b) after a female specimen in bad condition was found in Meudon (Seine-et-Oise). Like Rageau et al. (1970) we consider that this record is doubtful as it is well outside the known range of the species (Mediterranean basin, especially North Africa). A second report (Ségui, 1966) mentions three discoveries of the species (124 larvae) in Corsica in 1966. Even though this report seems more reasonable from the biogeographical point of view, no specimens were preserved and the taxon is therefore excluded form the list until further surveys provide confirmation.

Aedes (Ochlerotatus) nigripes (Zetterstedt 1938) was mentioned by Séguy (1923) (4 females from Fontainebleau, June 1869, Dufour’s collection) and quoted by Hedeen (1959) on the same basis. On the other hand, Callot & Rioux (1965) excluded it from their list. Considering the extremely northerly circumpolar range of the species, it is most unlikely to belong to the French fauna.

Aedes (Ochlerotatus) caspius hargreavesi Edwards 1920 was described from Italy and is mentioned as present in France in the calalogues of Knight & Stone (1977) and Dahl & White (1978). Séguy (1923) considered that the species could possibly occur in the south of France. Lack of precise records and a general absence of data regarding this sub-species, the valadity of which requires confirmation, prevents us from retaining it in our lists.

Culex (Maillotia) deserticola Kirkpartick 1924 was recorded in Corsica by Ségui (1965) who stated that he had collected the species in the larval stage on five occasions during his surveys of that year. He also recorded larvae of Cx. hortensis. Differences between larvae of these two species are extremely small. For reasons of biogeographic disjunction (the species is known from North Africa and Asia Minor (Dahl & White, 1978)), and the absence of preserved specimen, we do not consider the species as part of the fauna of France.

Culex (Culex) prosecutor Séguy 1927 (=Culex pseudomimeticus Séguy 1925) was described by Séguy (1925b, 1927) from an incomplete larva found in the Camargue. Mattingly considers it to be a synonym of Cx. impudicus (in Sénevet & Andarelli, 1959). Although it is mentioned as present in France in various catalogues (Knight & Stone, 1977; Dahl & White, 1978; Minar, 1990), no new information has been published regarding this taxon. The description of the extremity of the abdomen matches that of larvae of the sub-genus Neoculex, which is represented in France (and in Europe) by Cx. impudicus, Cx. martinii and Cx. territans. No major taxonomic characters allow differentiation of the larvae of these three species. Both Cx. impudicus and Cx. martinii can be encountered in the area where Séguy found the larvae. Because of the lack of information about this specimen, we are unable to attribute it to any of these species. Therefore, we propose classifying Cx. prosecutor as nomina dubia.

We consider that the following species have a reasonable chance of being encountered in France (including Corsica):

Anopheles (Cellia) cinereus hispaniola (Theobald 1903): this species is common in Sardinia, and could therefore be discovered in Corsica (Aitken, 1954; Rageau et al., 1970).

Aedes (Ochlerotatus) leucomelas (Meigen 1804) and Culiseta (Culicella) ochroptera (Peus 1935): both species occur in Germany, close to the French border (Oberrheinebene and Schwarzwald respectively) (Becker & Ludwig, 1981). Populations could perhaps be discovered in the north-east of France.

Aedes (Stegomyia) albopictus (Skuse 1894) and Aedes (Ochlerotatus) atropalpus s.s. (Coquillett 1902): these species were introduced into Italy with used tyres imported from North America. The former occurs in Piedmont and Sardinia; it first appeared in 1990 and has ever since expanded its range (Knudsen et al., 1996). The latter was discovered in 1996 in Venice (Romi et al., 1997). Ae. albopictus poses health threats, due to its high capacity to transmit arboviruses. France is considered a "high introduction risk" country (Knudsen et al., 1996), and it is important to monitor closely the potential breeding niches (tyres, various containers, hollow trees) in the south of France and Corsica.

Aedes (Finlaya) echinus (Edwards 1920), Culex (Culex) laticinctus Edwards 1913 and Culex (Culex) univittatus Theobald 1901: as mentioned earlier, these Mediterranean species have not been positively recorded in France, but could nevertheless be present in the south of France and, in particular, Corsica.

The French Culicidae fauna comprises 63 species to date. Even though this fauna is relatively well known, this list is not definitive. Indeed, thorough investigations of poorly studied areas and border regions could result in the addition of many species currently recorded in neighbouring countries. Additionally, global climatic change will very probably result in a northward extension of the range of some species. Systematic monitoring would allow the detection of such new arrivals, whether due to range extensions or accidental introductions.

Acknowledgements — This paper was written within the scientific programme of the limitation de la nuisance due aux moustiques which is funded by the Conseil Général du Haut-Rhin. I also wish to thank my friend Jacques Brunhes (ORSTOM) for advice on presentation. I also wish to thank the editors for translating the paper into English.