Clement Ramsdale¹ & Keith Snow²

1Varndean Lodge, London Road, Brighton, BN1 6YA and ² Department of Environmental Sciences, University of East London, Romford Road, London E15 4LZ UK Email: clem.ramsdale@claranet.co.uk; k.r.snow@uel.ac.uk

Maps and distribution data are presented for the eighteen species of Anopheles currently recognised in Europe.

Eighteen species of Anopheles are recognised in Europe: fourteen in the subgenus Anopheles, which is found throughout the continent, and four in the subgenus Cellia, which has a southerly distribution in Europe. In the past, Anopheles distribution in Europe has been the subject of many treatises at country level and has been the subject of reviews, most recently by Dahl & White (1978), White (1978), Stegnii (1991) and Jetten & Takken (1994). The present work aims to extend and update previous reviews by mapping distributions as precisely as is currently possible. To this end, maps have been produced showing the known distribution of all eighteen members of the genus. These are accompanied by details of the locations and the sources of the information, compiled from published records and personal communications. There are no records of any mosquito species from Iceland and there is only one record (An. maculipennis s.s.) from Luxembourg (Schaffner, 1999).

Records fall into five categories: 1. villages, towns and cities e.g. Frankfurt am Main, Barcelona; 2. administrative regions, e.g. Essex, Aude; 3. geographic areas e.g. Danube delta, Vardar valley; 4. sections of a country e.g. northern Italy, southern Sweden; 5. countries e.g. Switzerland, Estonia i.e. present in country with no further details.

These maps do not represent snapshots of current distribution as they are compiled from data accumulated over several decades. However they represent the best estimate of the current distribution of these species. It is fully appreciated that with very old records there have certainly been geographic changes since the records were collected and hence there must be doubt as to their current validity. In addition, some old records such as those relating to An. maculipennis/ beklemishevi, are almost impossible to unravel.

Clearly there is an urgent need for the production of more precise national records to allow European maps to be refined. A major aim of this article is therefore to stimulate further interest in the production of detailed country distribution maps for the whole of Europe. Utrio (1979) produced distribution maps for Finland using a 50km grid system but it is suggested that the more precise 10 km grid system, as used for Britain (Snow et al., 1998) and Poland (Kubica-Biernat, 1999) be adopted.

Information on species distribution is dynamic and continuously changing and hence the authors would be grateful to have omissions and errors communicated to them either personally or through the pages of the Bulletin. Once these data have been refined, it is anticipated that they will allow comparisons and correlations to be made between the distribution of individual species and geographic features, and help in assessing the validity of outlying and discontinuous records. They will also permit prediction of the presence of species in countries where they are not currently located and allow monitoring of the effects of predicted climate change on the distribution of mosquitoes and of the human and animal diseases that they transmit.

It is clearly not possible to consider distribution without reference to taxonomy, as sound identification is required to make certain that distributional data is valid. For Anopheles there are still a number of concerns and these are highlighted in the sections on individual species. For example, in many countries there is a need to unravel the Maculipennis Complex. This is seen particularly in Ireland and Switzerland. It is most likely that the species in Ireland are An. messeae and An. atroparvus and in Switzerland it is these two species together with An. maculipennis s.s. However this needs further investigation. Such holes in our current knowledge are a barrier to the fuller understanding of the distribution of Anopheles mosquitoes in Europe.

In the following listings only those records pertaining to the European areas of Russia and Turkey are included.

Anopheles (Anopheles) algeriensis Theobald, 1903

The main distribution encompasses the Mediterranean countries and Balkans, with an eastward extension into Iraq, Iran and, northwards through the Caucasus, into middle Asia, with some isolated populations outside this range. Within Europe there are records from Albania (Bates, 1941), Austria (Moog, 1995), Britain (Anglesey in Wales, Norfolk in England) (Snow et al., 1998), Bulgaria (Gecheva, 1998), Croatia (Neretva Delta, Zadar area, northern Dalmatia and Island of Pag) (Adamovic, 1983; 1984), Estonia (Saaremaa Island) (Remm, 1957), France (Aude, Bouche-de-Rhone, Charente-Maritime, Gard, Gironde, Herault, Loire-Atlantique, Morbihan, Pyrenees-Orientales and Corsica (Moussiegt, 1986; Schaffner, 1998), Germany (Krefeld and Frankfurt am Main areas) (Mohrig, 1969), Greece (Attica, Epirus, Lamia, Macedonia, Pelepponese, Drama River in Thrace, Kavalla, Vermion, Crete) (Waterston, 1918; Pandazis, 1935; Peus, 1954; Samanidou-Voyadjoglou & Darsie, 1993), Hungary (Balaton) (Mihalyi, 1955; Toth & Saringer, 1997), Ireland (Clare) (Ashe et al., 1991), Italy (northern and southern Italy, Sicily, Sardinia) (Sabatini et al., 1981; Coluzzi & Sabatini, 1995; Romi et al., 1997), Moldova (Gutsevich et al., 1971; Zubchuk, 1989), Portugal (Boca do Rio - Algarve) (Ramos et al., 1982; Ribeiro et al., 1988; Ribeiro & Ramos, 1999), Russia (Sulak River of Dagestan, Volga and Don basins, Caucasus) (Gutsevich et al., 1971; Gornostaeva, 2000), Spain (Alicante, Almeria, Granada, Murcia, Majorca, Balearics) (Encinas Grandes, 1982), Ukraine (south of about 51°N) (Gutsevich, 1989).

Anopheles (Anopheles) atroparvus van Thiel, 1927

This species has been recorded from Austria (Moog, 1995), Belgium (Gosseries & Goddeeris, 1991), Britain (Essex Rivers, Thames Estuary, Romney Marsh, Pevensey Levels, lower reaches of Sussex rivers, Hayling Island, Dorset, Devon, Berkshire, Cheshire, Surrey, all in England and Anglesey in Wales) (Snow et al., 1998), Belarus (generally west of the Dnieper River) (Sergeyeva, 1937; Artemiev, 1980; Stegnii, 1991), Bulgaria (Gecheva, 1998), Boznia-Herzegovina (Brcko, Macva, Srem) (Adamovic, 1979; 1982), Croatia (Istria, northern Dalmatia, Kuarne Island) (Adamovic, 1986a; Adamovic & Paulus, 1987), Czech Republic (Moravia) (Minar & Halgos, 1997), Denmark (Jutland, Falster, Langeland, Lolland, Sjaelland) (Wesenberg-Lund. 1921; Bruce-Chwatt & de Zulueta, 1980; Nielsen & Pedersen, 1997), France (Allier, Aude, Bouche-du-Rhone, Calvados, Charente-Maritime, CorrÀ ze, FinistÀ re, Gard, Haute-Garonne, Hérault, Indre, Indre-et-Loire, Loire-Atlantique, Manche, Morbihan, Pyrénées-Orientales, Savoie, Var, Vendée) (Moussiegt, 1986; Schaffner, 1998), Germany (all coastal and estuarine plains and inland from Eisleben, Frankfurt-am-Main, Goslar, Leipzig, Magdeburg, Schwarzwald and Thuringian lakes) (Mohrig, 1969), Hungary (Balaton) (Mihalyi, 1955; Toth & Saringer, 1997), Ireland (probable – see maculipennis), Italy (Po delta and peninsular as far south as the Volturno River; replaces An. labranchiae in the north) (Ramsdale & Coluzzi, 1975; Coluzzi & Sabatini, 1995; Romi et al., 1997), Latvia (Spungis, 2000), Lithuania (Pakalniskis, 1999), Macedonia (Skopje) (Weyer, 1942; Lepes & Vitanovic, 1962), Moldova (Zubchuk, 1989; Stegnii, 1991), Netherlands (de Meijere, 1939; 1946; 1950), Poland (Bydgoszcz, Gdansk, Katowice, Koszalin, Szczecin, Wejherowo/ Darlowa) (Skierska, 1963; Okroy-Rysop et al., 1991; Kubica-Biernat, 1997, 1999), Portugal (widespread in Algarve, Alentejo, Beira Alta, Arrábida, Beira Baixa, Esremadura) (Ramos et al., 1977/78; Ribeiro et al., 1989, 1996; Ribeiro & Ramos, 1999), Romania (Danube delta, Dobrudja plain, Wallachia) (Nicolescu, 1995), Russia (north of the Caucasus from the Caspian to the Black and Azov Seas) (Bates et al., 1949; Gutsevich et al., 1971; Artemiev, 1980; Stegnii, 1991; Gornostaeva, 2000), Slovakia Plain of Morava and Danube rivers (Jalili, 1995; Minar & Halgos, 1997; Jalili et al., 2000), Slovenia (Ljubliana) (Adamovic & Paulus, 1988), Spain (Albacete, Alicante, Almeria, Asturias, Avila, Badajos, Balearics (uncertain) Barcelona, Burgos, Caceres, Cadiz, Castellón, Ciudad Real, Cordoba, Cuenca, Granada, Huelva, Huesca, Jaen, Leon, Lerida, Madrid, Malaga, Murcia, Navarra, Orense, Palencia, Pontevedra, Salamanca, Santander, Segovia, Sevilla, Soria, Teruel, Toledo, Valencia, Valladolid, Zamora, Zaragoza) (Encinas Grandes, 1982; Lucientes, 1996; Eritja et al., 1998), Sweden (Skane, Blekinge, Gotland, Vastergotland (Ekblom, 1938; Jaenson et al., 1986; Dahl, 1997), Ukraine (Dnepropetrovsk, Crimea) (Gutsevich et al., 1971; Artemiev, 1980; Zarechnaya et al., 1989), Yugoslavia (Tamnic area, Potisje, Karst area of East Serbia, Tisa, Zlatica and Tarnts rivers of Pannonian plain (Belgrade area) (Adamovic, 1984, 1986b, 1987; Bozicic, 1985, 1987; Srdic et al., 1986; Adamovic & Paulus, 1987).

There are no records from Albania, Greece, Malta, Turkey, Sardinia, Sicily, Norway, Finland, Estonia, northern Russia, Sweden north of about 57ºN, and Switzerland. Briegel (1998) lists only maculipennis s.l. for Switzerland but Fouque et al. (1991) found three species of the Maculipennis Complex in the Magadino Plain, without putting names to them). A common mosquito in the Spanish south coastal Barbate and Tarifa areas, it has not been detected in Morocco on the other side of the 15-km wide Straits of Gibraltar.

Anopheles (Anopheles) beklemishevi Stegnii & Kabanova, 1976

The distribution of An. beklemishevi includes the boreal coniferous forest (taiga) region of Sweden and Finland (Jaenson et al., 1986; Utrio, 1975; 1978; 1979) and extends through the taiga belt and northern plain of European Russia into Siberia (Novikov & Alekseev, 1989; Stegnii, 1991; Gornostaeva, 2000). The southern distribution boundary of An. beklemishevi in European Russia is at about 60°N (Stegnii, 1991).

Anopheles (Anopheles) claviger (Meigen, 1804)

This species and Anopheles petragnani are sibling species and there may be confusion in identification, especially in older literature. An. claviger appears to be a northern mosquito which has penetrated into the Mediterranean by exploiting cold water issuing from springs or contained in underground cisterns and wells (Coluzzi et al., 1965). The distribution of An. claviger covers virtually the whole of Europe, and extends east to Afghanistan, southeast into the Middle East and south and southwest into North Africa. It may be absent from the Balearics, Sardinia and Malta and has not been specifically reported from Moldova, Belarus (although Gutsevich et al. (1971) indicate its presence throughout the European region of the former USSR) and Luxembourg. European records are from:

Albania (Bates, 1941), Austria (Moog, 1995), Belgium (Gosseries & Goddeeris, 1991), Boznia-Herzegovina (border area with Croatia and Yugoslavia, Malovan, Kova¹ i) (Paulus & Adamovic, 1983, 1985), Britain (throughout England, Scotland, Wales and Channel Isles) (Ramsdale, 1991; Snow et al., 1998), Belarus (indicated by Gutsevich et al., 1971), Bulgaria (Gecheva, 1998), Croatia (Neretva Delta, Lika, North Dalmatia and Island of Pag, Zagreb, Kamenmost, Istrian Peninsula [Livade, Cepic, Luka Plomion, Putera, Secovije]) (Adamovic, 1983, 1984; Adamovic & Paulus, 1985a, 1985b, 1987; Paulus & Adamovic, 1985), Czech Republic (Trebon Basin, Melnik, Hrade Kralove, Podebrady) (Rettich, 1973, 1979, 1982; Rettich et al., 1978; Minar & Halgos, 1997), Denmark (Neilsen & Pedersen, 1997), Estonia (Remm, 1957), Finland southern Finland, Lemland, (Utrio, 1975; 1976; 1979; Dahl, 1997), France (Ain, Aisne, Allier, Hautes-Alpes, Alpes Maritimes, Ardennes, Ariège, Aube, Aude, Aveyron, Bouche-du-Rhône, Calvados, Charente-Maritime, Cote-d'Or, Dordogne, Drôme, Eure-et-Loir, Gard, Haute-Garonne, Gironde, Hérault, Ille-et-Vilaine, Indre-et-Loire, Isère, Landes, Loire, Haute-Loire, Loire-Atlantique, Loiret, Lot, Maine-et-Loire, Manche, Marne, Meurthe-et-Moselle, Meuse, Morbihan, Moselle, Nord, Oise, Pas-de-Calais, Puy-de-Dôme, Pyrénées Atlantiques, Haut-Pyrénées, Pyrénées-Orientales, Bas-Rhin, Haut-Rhin, Rhône, Haute-Saône, Savoie, Seine-Maritime, Seine-et-Marne, Deux-Sèvres, Somme, Tarn, Var, Vaucluse, Vendée, Vienne, Yonne, Yvelines, territoire de Belfort, région parisienne and Corsica (Moussiegt, 1986; Schaffner, 1998), Germany (Frankfurt am Main, Krefeld, Munich, Mannheim, Black Forest, Meiningen, Bad Naucheim, Erlangen, Sontheim, Leipzig, Halle, Oberschwaben, Nettegebiet, Nordhausen, Baverian Alps, Maulbronn, Fichtelgebirge, Spessart, Erfurt, Neckarsulm, Schleiz, Wiermar, Plagefenn, Tiefenwerder, GØ ttingen, Greifswald, Bolter Kanal, Tò bingen, Upper Rhine) (Mohrig, 1969; Becker & Kaiser, 1995; Becker, 1997), Greece (north, central and south Greece [Thrace, Macedonia, Phthiotis, Attiki, north-west and central Peloponnese] (Lividas & Sphangos, 1941; Samanidou-Voyadjoglou & Darsie, 1993; Samanidou-Voyadjoglou, 1997), Hungary (Near Homorud/ Hoduna, Drava region, Lake Balaton) (Toth, 1992, 1995; Toth & Saringer, 1997), Italy (north, south, Sicily) (Coluzzi & Sabatini, 1995; Romi et al., 1997; Zamburlini, 1998), Ireland (Carlow, Clare, Cork, Down, Dublin, Galway, Kerry, Kildare, Laois, Limerick, Mayo, Offaly, Westmeath, Wexford, Wicklow) (Ashe et al., 1991), Latvia (Spungis, 2000), Lithuania (Pakalniskis, 1989), Macedonia (Drachevo) (Weyer, 1942), Moldova (indicated by Gutsevich et al., 1971), Netherlands (de Meijere, 1939, 1946, 1950), Norway (Trandum) (Mehl et al., 1983; Dahl, 1997), Poland (common throughout country; Warsaw, Mazovia, Zarnowieckie Lake, Gdansk, Wilkowska valley, Wolin Island) (Skierska, 1963, 1974; Wegner, 1979, 1982, 1991; Okroy-Rysop et al., 1991; Wegner et al., 1993; Kubica-Biernat, 1997, 1999), Portugal (Serra da Arrábida, Beiras) (Ribeiro et al., 1977, 1988, 1996; Ribeiro & Ramos, 1999), Romania (Salva) (Ciolpan et al., 1988; Nicolescu, 1995), Russia (European Russia north to St. Petersburg region) (Gutsevich et al., 1971; Gornostaeva, 2000), Slovakia (plain of Morava and Danube rivers, Moravia flood plain on Slovak-Austrian border, nr. Bratislava) (Jalili & Labuda, 1994; Jalili, 1995; Minar & Halgos, 1997; Jalili & Degma, 1998; Jalili et al., 2000), Slovenia (Ljubliana, close to border with Croatia and Italy) (Trpis & Tovornik, 1958; Adamovic & Paulus, 1988; Tovornik, 1990), Spain (Asturias, Badajoz, Barcelona, Caceres, Cadiz, Castellon, Ciudad Real, Córdoba, Cuenca, Granada, Jaen, Madrid, Malaga, Murcia, Salamanca, Santander, Segovia (Encinas Grandes, 1982; Lucientes, 1996; Eritja et al., 1998), Sweden (Oland, Skane, Halland, Dalarna) (Dahl, 1975, 1977a; Jaenson et al., 1986), Switzerland (Briegel, 1973, 1998), Turkey (Tekirdag Province) (Postiglione et al., 1973), Ukraine (Depropetrovsk region, Crimea) (Gutsevich et al., 1971; Zarechnaya et al., 1989), Yugoslavia (border area with Boznia-Herzegovina and Croatia, Vojvodina, Karst area, Temnic, Potisje, Vitkovac, Belusic, Planinica, Seliste, Bacina) (Paulus & Adamovic, 1983; Bozicic, 1985; Adamovic, 1986b, 1987; Srdic et al., 1986).

Anopheles (Anopheles) hyrcanus (Pallas, 1771)

The Hyrcanus Group comprises a large number of species, including the Far Eastern malaria vector An. sinensis, and has a wide Oriental and Palaearctic distribution. This Group has been intensively studied in the Oriental Region and the contiguous extreme eastern part of the Palaearctic, from where no less than 27 species have been described. Comparable studies have not been made elsewhere in the Palaearctic Region, where a number of forms differing in certain aspects of adult morphology and behaviour occur (Livadas & Sphangos, 1941; Ward, 1972; Postiglione et al., 1973; Critescu et al., 1975; Gutsevich, 1976). These include the named forms, flerowi, mahmouti, marzinovski, mesopotamiae, pictus, popovi and pseudopictus, treated by Knight & Stone (1977) as synonyms of the type form. An. pseudopictus, described in Europe from an unknown location in Italy (Grassi, in Grassi et al., 1899), has since been elevated to species status by Glick (1992) based on a study outside Europe.

Laboratory crosses between An. lesteri (a member of the Hyrcanus Group) from the Philippines and 'An. hyrcanus' from southern France, southern Turkey and Sri Lanka indicated that each population belonged to a different species (Ross Institute, unpublished reports to W.H.O., 1976, 1977). The same work showed a population of ‘An. hyrcanus’ from southern France to be conspecific with another from southern Anatolia. At least two Anatolian populations are morphologically distinguishable (Postiglione et al., 1973). Until it is more thoroughly investigated, ‘An hyrcanus’ in the western Palaearctic, including Europe, must necessarily continue to be treated as a single entity. However, it is already evident that existing biological information refers to an aggregate of species and not to a single polytypic species.

The European distribution of the western forms, 'An. hyrcanus', includes Albania (Bates, 1941), Bulgaria (Gecheva, 1998), Croatia (Dalmatian coast) (Merdic, 1997), France (Ain, Bouches-du Rhone, Gard, Haut-Garonne, Herault, Corsica) (Moussiegt, 1986; Schaffner, 1998), Greece (Epirus, Lake Doira, Macedonia, Peloponnese, Thessaly, Kavalla in Thrace, Rhodes, Crete) (Pandazis, 1935; Peus, 1954; Samanidou-Voyadjoglou & Darsie, 1993), Hungary (Lake Balaton, Dráva) (Mihalyi, 1955; Toth, 1995; Toth & Saringer, 1997), Italy (north, south, Sicily, Sardinia) (Coluzzi & Sabatini, 1995; Romi et al., 1997), Macedonia (Lake Doiran) (Weyer, 1942), Moldova (Kishinev) (Gutsevich et al., 1971), Romania (Tulcea, Danube Delta) (Critescu et al., 1975; Nicolescu, 1995), Russia (lower Don and Volga basins, Caucasus up to 1600m) (Gutsevich et al., 1971; Gornostaeva, 2000), Spain (Barcelona, Huesca, Teruel, Valencia) (Encinas Grandes, 1982; Eritja et al., 1998), Ukraine (south of 50°N) (Gutsevich et al., 1971), Yugoslavia (Durmitor Massif, Stara Planina, Novi Sad) (Bozicic, 1987; Bozicic-Luthrop & Vujic, 1996; Petric, 1999).

Anopheles (Anopheles) labranchiae Falleroni, 1926

An. labranchiae is a North African species, abundant and widespread within, north and west of the Atlas mountain ranges. A few parts of southern Europe have been colonised, probably within historical times. This was made possible by climatic amelioration since the end of the last glaciation coupled with the more recent creation of suitable larval conditions by human activities, notably forest clearance (de Zulueta, 1973). The European distribution is principally coastal; with records from Croatia (Kvarneric Islands, Zadar area, north Dalmatia) (Adamoviº , 1984; 1986a), France (Corsica only according to Schaffner (1998) although Moussiegt (1986) lists a record from Haute-Garonne), Italy (Sardinia, Sicily, Italian Peninsula from Calabria north to Tuscany in a discontinuous manner along both coasts – replaced by An. atroparvus towards north) (Hackett & Missiroli, 1935; Coluzzi & Sabatini, 1995; Romi et al., 1997; Romi, 1999).

Despite the proximity of Morocco, where it is abundant, it does not occur on the Spanish side of the 15-km wide Straights of Gibraltar, where An. atroparvus is common. It was previously recorded from the most arid regions of the provinces of Alicante and Murcia in Spain from which it was eradicated during the course of anti-malarial operations (Blazquez & de Zulueta, 1980; Encinas Grandes, 1982). Records from Bulgaria, (Gecheva, 1998), the Czech Republic and Slovakia (Minar & Kramar, 1987; Minar & Halgos, 1997) are considered erroneous due to the former practices of considering An. maculipennis s.l. as a single polytypic species, and/or atroparvus being a subspecies of labranchiae.

Anopheles (Anopheles) maculipennis Meigen, 1818

At first thought to have a very wide European and Asian distribution, the discovery of the apparently allopatric An. beklemishevi led to the realisation that records of An. maculipennis from much of northern European Russia and Scandinavia, and northern and central Asia probably refer to the former species (White, 1978). As currently understood, the distribution of An. maculipennis covers the greater part of Europe. It has not been recorded in Britain (Cranston et al., 1987), the southern part of the Iberian Peninsula (Encinas Grandes, 1982; Ribeiro et al., 1989), Ireland (only maculipennis s.l. recorded - almost certainly atroparvus and/or messeae as in Britain) (Ashe et al., 1991), Sardinia (Coluzzi & Sabatini, 1995; Romi et al., 1997), much of Norway (Dahl, 1977a) and northern Sweden (Ekblom, 1938). A record from the Balearic Islands may have referred to specimens of An. subalpinus, which is present in eastern Spain (Romeo Viamonte, 1950).

An. maculipennis is widely distributed over most of the continent, being recorded from: Albania (Bates, 1941), Austria (Moog, 1995), Belarus (Mogilev region) (Trukhan et al., 1988; Stegnii, 1991), Belgium (Gosseries & Goddeeris, 1991), Bosnia-Herzegovina (Malovan, Kovaci, border area with Croatia and Yugoslavia) (Paulus & Adamovic, 1983, 1985), Bulgaria (Gecheva, 1998), Croatia (Zagreb, Istrian Peninsula, border area with Bosnia-Herzegovina and Yugoslavia) (Adamovic & Paulus, 1983, 1988; Paulus & Adamovic, 1985), Czech Republic (Trebon basin) (Rettich et al., 1978; Minar & Halgos, 1997), Denmark (Nielsen & Pedersen, 1997), Estonia (Remm, 1957), Finland (Vihanti) (Lumiaho & Itamies, 1981; Dahl, 1997), France (all departements except Cher, Doubs, Eure, Indre, Indre-et-Cher, Lozère, Haute-Marne, Mayenne, Nièvre, Orne, Sarthe, Vosges; present in Corsica) (Moussiegt, 1986; Schaffner, 1998), Germany (upper Rhine valley, Thuringia, southern Germany) (Mohrig, 1969; Becker & Kaiser, 1995), Greece (Boeotia, Epirus, Lamia Plain, Macedonia, Peleponese, Thessaly and Thrace, but recent information is sparse) (Barber, 1935; Hackett & Missiroli, 1935; Shannon, 1935; Sakellariou & Lane, 1977; Samanidou-Voyadjoglou & Darsie, 1993; Samanidou-Voyadjoglou, 1997), Hungary (Drava, Ortilos) (Mihalyi, 1955; Toth, 1995), Italy (widespread in peninsula, present but rare in Sicily and absent from Sardinia (Coluzzi & Sabatini, 1995: Romi et al., 1997), Latvia (Stegnii, 1991; Spungis, 2000), Lithuania (Pakalniškis, 1989; Stegnii, 1991), Luxembourg (Schaffner, 1999), Macedonia (Skopje, Vor Matka, Katlanova, Drachova, Isdeklave, Longovarde) (Weyer, 1942), Moldova (Stegnii, 1991), Netherlands (de Meijere, 1939, 1946, 1950), Norway (Dahl, 1997), Poland (north-west Poland, Gdansk area, Wejherowo/ Darlowo area (Lukasiak, 1959; Skierska, 1963; Okroy-Rysop et al., 1991; Kubica-Biernat, 1997, 1999), Portugal (Beiras) (Ribeiro et al., 1989; Ribeiro & Ramos, 1999), Romania (Nicolescu, 1995), Russia (south of about latitude 60^ON) (Novikov & Alekseev, 1989; Stegnii, 1991; Gornostaeva, 2000), Spain (Asturias, Guadalajara, Guipúzcoa, León, Lerida, Lugo, Orense, Palencia, Santander, Tarragona) (Encinas Grandes, 1982; Eritja et al., 1998), Sweden (south of about 60^ON) (Jaenson et al., 1986), Switzerland (uncertain – see under messeae) (Briegel, 1973), Slovakia (Minar & Halgos, 1997; Jalili et al., 2000), Slovenia (Ljubliana, nr. San Dorligo della Valle on Italian border, nr. Lendava, nr. Krizsevci) (Trpis & Tovornik, 1958; Tovornic, 1983; Adamovic & Paulus, 1988), Turkey (Edirne, Kirklareli, Tekirdag, Istanbul provinces) (Postiglione et al., 1973), Ukraine (Zarechnaya et al., 1989), Yugoslavia (border area with Croatia and Boznia-Herzegovina, Potisje region and Vrsac Mountains (Vojvodina), Morava river to Stara Planina (Nis, Pirot, Trnovac), Gruza, Pozega, Ivanjica, Levac, Tamnic areas) (Paulus & Adamovic, 1983; Adamovic, 1986a 1987; Bozicic, 1985, 1988).

Anopheles (Anopheles) marteri Senevet & Prunelle, 1927

This species is present in north-western Africa, southern Europe, Asia Minor, Cyprus, Middle East and Tadzhikstan. European records are from Albania (Shkumbini Valley) (Bates, 1941), Bulgaria (Mount Beles) (Bozkov, 1966; Gecheva 1998), France (Corsica) (Langeron & Galliard, 1933; Schaffner, 1998), Greece (Mount Pangeon, Ypati, Kallidromon, Mount Iti, Parnassos, Delphi) (Shannon, 1933; Hadjinicolaou, 1938; Livadas & Sphangos, 1941; Samanidou-Voyadjoglou & Darsie, 1993), Italy (Calabria, central-southern Apennines, Sicily, Sardinia) (Sabatini et al., 1981; Coluzzi & Sabatini, 1995; Romi et al., 1997), Portugal (north-western Tras os Montes) (Ribeiro et al., 1985; Ribeiro & Ramos, 1999), Spain (Cuenca, Granada, Jaen, Murcia) (Encinas Grandes, 1982; Eritja et al., 1998).

Anopheles (Anopheles) melanoon Hackett, 1934

An. melanoon is known to occur in mainland Italy (most prevalent in the north and in hilly areas of central and southern Italy) (Coluzzi & Sabatini, 1995; Romi et al., 1997). Records from Sardinia predate the 1950s and its existence there now is considered doubtful (Marchi & Munstermann, 1987; Romi, 1999).

First regarded as a variety of messeae and subsequently as con-specific with subalpinus (Hackett & Lewis, 1935), melanoon was regarded as having an extensive southern European distribution (White, 1978). The realisation that these taxa are all separate species (Bullini et al., 1980; Cianchi et al., 1987), leaves many records open to question.

An. melanoon was recorded in the south of continental France and in Corsica (Moussiegt, 1986; Schaffner, 1998) during the time when subalpinus was considered to be conspecific. It is unclear which records refer to melanoon, but Saliers et al. (1978) found subalpinus to be abundant in southern France. Amongst eggs laid by mosquitoes collected near Toulouse were some resembling An. melanoon (Suzzoni-Blatger, 1989); however iso-enzyme analysis of this population detected An. maculipennis, An. messeae and An. atroparvus but not An. melanoon (Suzzoni-Blatger et al., 1990). A unique record of An. melanoon from Switzerland is considered questionable by Briegel (1999). The situation is further complicated by the tendency of An. subalpinus to occasionally oviposit a dark, melanoon-like egg batch (Coluzzi, 2000).

There are two identifications from the Iberian Peninsula separated by 26 years and almost 600 km. One, from an inland rural area in Portugal (Serra do Barrosa) was made on six eggs laid by a dying captive female (Ribeiro et al., 1980; Ribeiro & Ramos, 1999); the other, from the port of Valencia in Spain, was identified on chromosomal evidence from preserved material (Frizzi, 1953).

In the eastern Mediterranean, melanoon and subalpinus were not recorded separately in Albania and there is only a single report of the melanoon form during many years of mosquito collecting by entomologists of the Rockefeller Foundation and the Greek National Malaria Control Organisation in Greece (Lividas & Sphangos, 1941). Occasional reports of melanoon in Asia Minor were found to be due to examination of wetted eggs with obscured deck patterns (Postiglione et al., 1973). It is included in the checklist of mosquitoes from European Russia from where subalpinus is also recorded (Gornostaeva, 2000).

Anopheles (Anopheles) messeae Falleroni, 1926

Widespread in Europe with the south-eastern limits of the distribution passing through Bulgaria (Gecheva, 1998), Romania (Nicolescu, 1995), Moldova (Zubchuk, 1989) and along the Black Sea coast of Ukraine and Russia (Stegnii, 1991). Outside Europe, the range extends eastward into Asiatic Russia, where it has an extensive distribution (Stegnii, 1991).

This species is present in Albania (the vicinity of the mountain lakes Malik, Ohrid and Prespa, through which the frontiers with Greece and/or Macedonia run (Bates, 1941; Adamoviº , 1980), Austria (Moog, 1995), Belarus (Stegnii, 1991), Belgium (Gosseries & Goddeeris, 1991), Bosnia-Herzegovina (Malovan, Kovaci, border area with Croatia and Yugoslavia) (Paulus & Adamovic, 1983, 1985), Britain (England only - Devon, Cambridgeshire, Suffolk, Surrey, Sussex, Norfolk, Northumberland) (Snow et al., 1998), Bulgaria (Gecheva, 1998), Croatia (Sava River, border area with Bosnia-Herzegovina and Yugoslavia, Locvicici, Kamenmost) (Paulus & Adamovic, 1983, 1985; Adamovic & Paulus, 1988; Merdic, 1997), Czech Republic (Trebon basin, Moravia flood plain) (Rettich et al., 1978; Minar & Halgos, 1997), Denmark (Nielsen & Pedersen, 1997), Estonia (Stegnii, 1991), Finland (Kumlinge, Tvarminne, Lohja, Joroinen, Rovaniemi, Sodankyla, Kittilä, Kolari) (Ulmanen & Brummer-Korvenkontio, 1971; Dahl, 1997), France (Ain, Bouches-du-Rhône, Haute-Garonne, Indre, Indre-et-Loire, Isère, Loire-Atlantique, Pyrénées-Orientales, Savoie, Vendée and Corsica (Rioux, 1958; Moussiegt, 1986; Schaffner, 1998), Germany (widely distributed, Upper Rhine valley) (Mohrig, 1969; Becker & Kaiser, 1995), Greece (limited to the vicinity of the mountain lakes Malik and Prespa. After the description of subalpinus (Hackett & Lewis, 1935) it was soon realised that messeae was confined to this small high altitude frontier area and, as pointed out by Lividas & Sphangos (1941), early records of messeae from elsewhere were, in reality, instances of subalpinus which is widespread in the country. More recently doubts were expressed about the absence of An. messeae and An. atroparvus (Samanidou-Voyadjoglou & Darsie, 1993; Samanidou-Voyadjoglou, 1997), but no evidence of the presence of either species has been found. Hungary (Dr´ va) (Mihalyi, 1955; Toth, 1995), Ireland (probable – see maculipennis), Italy (widespread in northern Italy and central Apennines) (Coluzzi & Sabatini, 1995; Romi et al., 1997), Latvia (Stegnii, 1991; Spungis, 2000), Lithuania (Stegnii, 1991), Macedonia (Skopje, Vor Matka, Katbarvo, Isdeklave, Longovade, Vardar Valley) (Weyer, 1942; Adamovic, 1980), Moldova (Zubchuk, 1989), Netherlands (de Meijere 1939, 1946, 1950), Norway (Dahl, 1997), Poland (Wejherowo/ Darlowo area, northern Poland (Okroy-Rysop et al., 1991; Kubica-Biernat, 1997, 1999), Romania (Danube Delta, Danube Plain, Dobrudja (references in Nicolescu, 1995), Russia (widespread from the plains bordering the Caucasus and Black Sea north to the White Sea) (Novikov & Alekseev, 1989; Stegnii, 1991; Gornostaeva, 2000), Sweden (Skåne, Blekinge, Småland, Gotland, Vastergotland, Bohusian, Narke, Sodermanland, Varmland, Dalarna, Angermanland, Vasterbotten, Norrbotten) (Dahl, 1977a, b; Jaenson et al., 1986; Jaenson & Ameneshewa, 1991), Slovakia (plain of Morava and Danube rivers) (Jalili & Labuda, 1994; Jalili, 1995; Minar & Halgos, 1997; Jalili et al., 2000), Slovenia (Ljubljana) (Tovornic, 1983), Switzerland (Briegel (1997) lists only maculipennis s.l. but Fouque et al. (1991) state that they found three species of the Maculipennis Complex in the Magadino Plain, without naming them.), Ukraine (Dnepropetrovsk) (Zarechnaya et al., 1989; Stegnii, 1991), Yugoslavia (border area with Bosnia-Herzegovina and Croatia, Morava river to Stara Planina area (Nis, Pirot, Trnovac), Potisje region (Vojvodina), Gruza, Levac, Tamnic (Adamovic, 1980, 1986b, 1987; Paulus & Adamovic, 1983; Adamovic & Paulus, 1984; Bozicic, 1985).

Anopheles (Anopheles) petragnani Del Vecchio, 1939

This species and Anopheles claviger are members of a Complex and there may be confusion in identification, especially in older literature.

An. petragnani is a Mediterranean species with larvae able to tolerate higher water temperatures than An. claviger (Coluzzi, 1962). For reasons not readily apparent, An. petragnani appears to be confined to the countries surrounding the western Mediterranean. In Europe it is recorded from France (southern provinces and Corsica only) (Coluzzi et al., 1965; Schaffner, 1998), Italy (north [Liguria], central southern regions, Sicily, Sardinia) (Coluzzi et al., 1965; Romi et al., 1997), Portugal (Arrabida, north-western Alentejo, western Algarve, Beira Alto, Beira Baixo Provinces) (Ramos et al., 1977/78; Pires et al., 1982; Ribeiro et al., 1988, 1996), Spain (Huesca, Salamanca, Balearic Islands) (Encinas Grandes, 1982; Eritja et al., 1998).

Anopheles (Anopheles) plumbeus Stephens, 1828

An. plumbeus is widely distributed throughout Europe, with a range extending from Asiatic Russia and contiguous countries, and the Middle East as far south as Iraq and Saudi Arabia, to Europe and North Africa. Because of its larval habitats it is under-recorded throughout its wide distribution. European records are from:

Albania (Bates, 1941), Austria (Moog, 1995), Belarus (Trukhan & Pakholkina, 1984), Belgium (Gosseries & Goddeeris, 1991), Boznia-Herzegovina (near border with Croatia, Srednja Posavina and Bijeldina in Save Valley, Imotsko, Mostar) (Paulus & Adamovic, 1983, 1985; Adamovic & Paulus, 1984), Britain (widely distributed throughout England, Scotland and Wales) (Snow et al., 1998), Bulgaria (Gecheva, 1998), Croatia (Hrvatsaka, Papuk highlands, Istrian peninsula, Zadar area, Osilek, Srednja Posavina in Save Valley, Velebit Planina, Zagreb) (Trpis & Tovornik, 1958; Adamovic, 1984; Paulus & Adamovic, 1983; Adamovic & Paulus, 1984, 1985a, 1987; Merdic, 1995), Czech Republic (Melnik, Moravia flood plain, Podbrady, Trebon Basin) (Rettich et al., 1978; Rettich, 1982; Minar & Halgos, 1997) Denmark (Neilsen & Pedersen, 1997), Estonia (Remm, 1957), France (Ain, Aisne, Alpes Maritimes, Ardennes, Ariège, Aveyron, Bouche-du-Rhône, Calvados, Charente-Maritime, Cote-d'Or, Dordogne, Drôme, Gard, Haute-Garonne, Hérault, Indre-et-Loire, Isère, Loire, Loire-Atlantique, Marne, Haut-Marne, Oise, Puy-de-Dôme, Pyrénées Atlantiques, Haut-Pyrénées, Pyrénées-Orientales, Bas-Rhin, Haut-Rhin, Rhône, Savoie, Seine-et-Marne, Deux-Sèvres, Var, Vendée, Vienne, Yvelines, territoire de Belfort, région parisienne and Corsica (Moussiegt, 1986; Schaffner, 1998), Germany (altmark, Lower and Upper Bavaria, Bodensee, Upper Danube valley, Hamburg, Harz, Hessen, Leipzig, Mecklenburg, Rhineland, Schleswig Holstein, Thuringenwald, Nord Rhein-Westfalen, Wurttemberg) (Mohrig, 1969; Becker & Kaiser, 1995), Greece (Corfu, Macedonia, Thrace) (Pandazis, 1935; Stephanides, 1938; Samanidou-Voyadjoglou & Darsie, 1993), Hungary (Balaton) (Mihályi, 1955; Toth & Saringer, 1997), Italy (North, South, Sicily, Sardinia) (Coluzzi & Sabatini, 1995; Romi et al., 1997), Ireland (Clare, Cork, Down, Dublin, Kerry, Kildare, Waterford, Wicklow) (Ashe et al., 1991), Lithuania (Pakalniškis, 1989), Macedonia (Struma Valley, Kumanovo, Leë nica, Sië evo, Skopje, Veles) (Martini, 1929-31; Weyer, 1942; Trpis & Tovornik, 1958), Netherlands (de Meijere, 1939, 1946, 1950), Poland (Bialowieza, Krakow) (Skierska. 1963, 1977; Kubica-Biernat, 1999), Portugal (scanty but general distribution; Beiras) (Ribeiro et al, 1988, 1989; Ribeiro & Ramos, 1999), Romania (Danube delta, Danube plain, northern Moldova, Salva) (Ciolpan et al., 1988; Nicolescu, 1995), Russia (Moscow, Caucasus) (Gutsevich et al., 1971; Gornostaeva, 2000), Slovakia (nr. Bratislava; plains of Morava and Danube rivers) (Jalila & Labuda, 1994; Jalili, 1995; Minar & Halgos, 1997; Jalili & Degma, 1998; Jalili et al., 2000), Slovenia (Ljubliana) (Tovornik, 1983; Adamovic & Paulus, 1988), Spain (Asturias, Barcelona, Cadiz, Cuenca, Madrid, Salamanca, Valencia, Zaragoza) (Encinas Grandes, 1982; Lucientes, 1996; Eritja et al., 1998), Sweden (Skane) (Natvig, 1948; Dahl, 1997), Switzerland (Adlikon, Bern, Genf, Lausanne, Tessin, Wallisellen) (Briegel, 1973), Turkey (Postiglione et al., 1973; Ramsdale, 1999), Ukraine (Crimea) (Gutsevich et al., 1971), Yugoslavia (Vrsac highlands, Karst area, border area with Croatia and Boznia-Herzegovina, Stara Planina, Temnic area, Morova Valley to Stara Planina and the Bulgarian border, Vodjvodina Plain, Drina/ Sava Plain, Decane) (Alkovic, 1949; Trpis & Tovornik, 1958; Paulus & Adamovic, 1983; Adamovic, 1986b, 1987; Bozicic, 1988; Srdic et al., 1986; Bozicic-Luthrop & Vujic (1996).

Anopheles (Anopheles) sacharovi Favre, 1903

Formerly the range of An. sacharovi was considered to extend from southern Europe to China. Since the recognition of the validity of An. martinius (Stegnii & Kabanova, 1976), and redefinition of the Maculipennis Complex (White, 1978), only the distribution west of the Caspian Sea is regarded as referring to An. sacharovi (Stegnii, 1991). This comprises the plains south of the Caucasus Mountains, Iran, the Middle East, Cyprus, Turkey and southern Europe. The European distribution includes Albania (coastal plains) (Bates, 1941), Bulgaria (Gecheva, 1998), Croatia (south Dalmatian coast) (Simic & Zivkovic, 1958), France (Corsica only) (Aitken, 1954; Schaffner, 1998), Greece (coastal and inland plains in Attica, Epirus, Euboea, Macedonia, Peloponnese, Thessaly, Thrace, Cyclades and Crete; very abundant throughout country) (Livadas & Sphangos, 1941; Hadjinicolaou & Betzios, 1973; Samanidou-Voyadjoglou, 1997), Macedonia (villages in the Vardar Valley near the Greek border, Skopje, Gevgelija) (Weyer, 1942; Lepes & Vitanovic, 1962), Russia (Dagestan) (Gutsevich et al., 1971; Stegnii, 1991; Gornostaeva, 2000), Turkey (Edirne, Kirklareli and Tekirdag Provinces) (Gökberk, 1961; Postiglione et al., 1973), Yugoslavia (Montenegro coast) (Simic & Zivkovic, 1958).

In Romania it was formerly found in Costanza, Tulcea and the Danube Delta but was eradicated during the anti-malaria campaigns (Lupascu & Duport, 1968; Nicolescu, 1995). Before the campaign of malaria control in Italy, it was present in several coastal areas of the peninsula and in Sardinia (Aitken, 1954; Coluzzi & Sabatini, 1995). It is now believed to have been eradicated from Italy (Romi et al., 1997) although the presence of residual foci cannot be totally discounted (Romi, 1999).

Anopheles (Anopheles) subalpinus Hackett & Lewis, 1935

There is confusion in the literature regarding this species and An. melanoon which, for a long time, were considered to be con-specific. The discovery that these taxa are separate species (Bullini et al., 1980; Cianchi et al., 1987) leaves some records open to question. Further, many records of messeae prior to the description of subalpinus (Hackett & Lewis, 1935) actually refer to the latter species.

The distribution of An. subalpinus is thought to extend through southern Europe from Iberia to Asia Minor, where it is common on the central Plateau (Postiglione et al., 1973), and the coastal plain of the Caspian Sea. Albania (old records indicate that it is widespread in the plains) (Hackett & Lewis, 1935; Bates, 1941), Bulgaria (as maculipennis melanoon) (Gecheva, 1998), Croatia (Dalmation coastal plains and islands; Kamenmost, Imotski) (Adamovic, 1984; Paulus & Adamovic, 1985), France (Ain, Bouche-de-Rhone, Gard, Herault, Indre-et-Loire, Corsica) (Moussiegt, 1986; Schaffner, 1998), Greece (Epirus, Macedonia, Thessaly, Thrace) (Livadas & Sphangos, 1941, as melanoon subalpinus; Samanidou-Voyadjoglou & Darsie, 1993, as melanoon subalpinus), Italy (uncertain, but probably north and south in peninsula; absent in Sicily and Sardinia) (Coluzzi & Sabatini, 1995; Romi et al., 1997), Macedonia (Skopje, Vor Matka, Katlanovo, Demin, Kapin, Orizari, Stojakova) (Weyer, 1942; Lepes & Vitanovic, 1962; Adamovic, 1984), Portugal (Escarigo in Serra da Marofa) (Ramos et al., 1982; Ribeiro & Ramos, 1999), Russia (Gornostaeva, 2000), Spain (as melanoon and melanoon subalpinus) Alicante, Barcelona, Castellón, Tarragona, Valencia (Encinas Grandes, 1982), Turkey (Edirne, Kirklareli, Tekirdag, Istanbul Provinces) (Postiglione et al., 1973), Yugoslavia (Montenegro coast) (Simic & Zivkovic, 1958).

Anopheles (Cellia) cinereus Theobald, 1901

An. cinereus is principally an Afrotropical species, with a distribution extending from the Cape Province of South Africa through the southern and eastern savannahs to the Horn of Africa and the south-western corner of the Arabian Peninsula (Gillies & de Meillon, 1968). There is one 1939-45 war-time record far to the north, from Jordan (Lumsden, 1944). Toward the north-eastern extremity of the East African distribution there is a north-western extension through the southern Sudan, the Saharan highlands and the western desert to the Mediterranean. It is the most widely distributed mosquito in the Maghrebian countries of North Africa, being found from the southern desert to the Atlantic and Mediterranean coasts. It occurs in the Canary Islands (Baez & Fernandez, 1980) and the most easterly Mediterranean record is from Cyrenaica in Libya (Ramsdale, 1990). It does not occur in Egypt, or in the Middle East and south-eastern Europe, where the larval niche is filled by An superpictus.

In Europe there are records from Portugal as An. cinereus hispaniola (Alto Douro) (Ribeiro et al., 1980; Ribeiro & Ramos, 1999) and Spain (Alicante, Almeria, Avila, Badajoz, Caceres, Cadiz, Cuidad Real, Cordoba, Cuenca, Granada, Huelva, Jaen, Madrid, Malaga, Murcia and Seville) (Encinas Grandes, 1982; Eritja et al., 1998). Isolated records from Greece: Epirus (Lividas, 1931) and Drama (Pandazis, 1935) are considered erroneous and probably referred to An. superpictus. Variations in adult coloration of both species, and especially of the arrangement of banding of the female palps, have led to instances of confusion between Anopheles cinereus and An. superpictus (Ramsdale, 1990).

Anopheles (Cellia) multicolor Cambouliu, 1902

A desertic species common in North Africa and with a distribution extending through the Sahara and the Arabian Peninsula to Pakistan. The only European records are from the Spanish Provinces of Murcia and Almeria (Encinas Grandes, 1982), the most arid region of the Iberian Peninsula. As in the case of An. labranchiae, An. multicolor may have been eradicated during the course of anti-malaria operations and its current status is uncertain.

Anopheles (Cellia) sergentii (Theobald, 1907)

The distribution of the anthropophilic An. sergentii sergentii extends from Pakistan, through the Middle East, the Arabian Peninsula, the Sahara, Atlas and coastal regions of North Africa to the Atlantic and the Canary Islands. That of the zoophilic An. sergentii macmahoni extends from the Horn of Africa, through the savannah belt bordering the southern edges of the Sahara into Mali and northwards into the Saharan oases, where some morphological intergrading occurs (Gilles & de Meillon, 1968; Ramsdale & de Zuluetta, 1983; Senevet & Andarelli, 1955). Apart from a single historical Bulgarian record reported by Markov & Morov (1929), the presence in Pantelleria (an island off the south coast of Sicily) of an apparently anthropophilic, but morphologically intermediate form (d’Alessandro & Sacca, 1967), is the sole record of An. sergentii north of the African coast.

Anopheles (Cellia) superpictus Grassi, 1899

The distribution extends from northern Pakistan and Afghanistan, through the Caucasus into southern Tadzikhstan and middle Asia, and through the Middle East and Asia Minor to southeastern Europe. European records are from Albania (coastal valleys) (Bates, 1941), Bulgaria (Gecheva, 1998), Croatia (Neratva Delta, North Dalmatia and Island of Pag) (Adamovic, 1983; 1984), France (Corsica only) (Moussiegt, 1986; Schaffner, 1998), Greece (Athens, Epirus, Etolia, Euboea, Macedonia, Peleponnese, Thessaly, Thrace, Corfu, Crete, Cyclades) (Waterston, 1918; Pandazis, 1935; Stephanides, 1937, 1938; Livadas & Sphangos, 1941; Peus, 1954; Samanidou-Voyadjoglou & Darsie, 1993), Italy (scattered foci in south and in Sicily, Calabria [Ionian and Tyrrhenian coasts]; not Sardinia) (Coluzzi & Sabatini, 1995; Romi et al., 1997), Macedonia (Greek border area) (Lepes & Vitanovic, 1962), Russia (Dagestan, Caucasus) (Gutsevich et al., 1971; Gornostaeva, 2000), Turkey (Edirne, Kirklareli, Tekirdag Provinces) (Postiglione et al., 1973), Yugoslavia (Karst area of East Serbia, Pukovat) (Adamovic, 1987). There are four doubtful records from continental Spain (Cordoba, Huelva, Murcia) and the Canary Islands (Encinas Grandes, 1982).