A review of species diversity and distribution of Culicoides
Latreille, 1809 (Diptera: Ceratopogonidae) in Turkey
BILAL DIK1, SUKRAN YAGCI2 & YVONNE-MARIE LINTON3
1Selcuk University, Faculty of Veterinary Medicine, Department of Parasitology, Konya, Turkey,2Ankara University, Faculty of Veterinary Medicine, Department of Parasitology, Ankara, Turkey,
and 3Department of Entomology, Natural History Museum, Cromwell Road, London, UK
(Accepted 29 September 2006)
AbstractThis paper reviews the species composition and distribution of Culicoides biting midges in Turkey forthe first time. The Culicoides fauna of Turkey is diverse, with 57 reported species collected in 54localities across the Marmara, Aegean, Mediterranean, and Central, Eastern and SoutheasternAnatolian Regions of Turkey. No records are available for the Black Sea Region. The mostwidespread species include Culicoides cataneii Clastrier, C. circumscriptus Kieffer, C. festivipennisKieffer, C. gejgelensis Dzhafarov, C. longipennis Khalaf, C. maritimus Kieffer, C. nubeculosus (Meigen),C. obsoletus (Meigen), C. odiatus Austen, C. pulicaris (Linnaeus), C. puncticollis (Becker), C. saevusKieffer, and C. shaklawensis Khalaf, whereas C. denisoni Boorman, C. dewulfi Goetghebuer, C. fagineusEdwards, C. minutissmus (Zetterstedt), C. montanus Schakirzjanova, C. semimaculatus Clastrier, C.slovacus Orszagh, and C. turanicus Gutsevich and Smatov are represented from only one locality. Onlysingle specimens represent the country records of C. dewulfi and C. seifadinei. The potential of TurkishCulicoides in zoonotic arbovirus, protozoa, and nematode transmission is discussed.
Keywords: Biting midges, Culicoides species, distribution, faunal list, Turkey, vector
Introduction
Bluetongue is an infectious, non-contagious, and often lethal disease in domestic ruminants
including sheep, goats, and cattle that is endemic in tropical latitudes worldwide.
Bluetongue is transmitted via the bites of some Culicoides biting midges and thus it follows
that the disease is present only where populations of competent vectors exist. Range
expansion of the major Old World vector, Culicoides imicola Kieffer, is believed to be a
key factor in the changes in bluetongue epidemiology in the Mediterranean basin
(Mellor and Wittman 2002). Culicoides imicola s.s. has expanded from Afrotropical zones
and now exists in stable populations in countries surrounding the Mediterranean,
including Turkey, below 40uN latitude. The first outbreak of bluetongue in Turkey
(serotype 4) was reported from Aydın province in western Turkey in 1977. The disease
overwintered and the outbreak lasted through to 1981 (Yonguc et al. 1982; Jennings et al.
Correspondence: Bilal Dik, Selcuk University, Faculty of Veterinary Medicine, Department of Parasitology, 42031 Campus,
Konya, Turkey. Email: [email protected]
Published 4 December 2006
Journal of Natural History, 2006; 40(32–34): 1947–1967
ISSN 0022-2933 print/ISSN 1464-5262 online # 2006 Taylor & Francis
DOI: 10.1080/00222930601046634
1983). The Imicola Group comprises some eight described species (with an additional four
remaining to be formally described), including C. imicola s.s., the major Afrotropical and
Mediterranean vector of bluetongue (Meiswinkel and Linton 2003). Despite several
molecular studies on the composition of this complex in the Mediterranean Basin
(including Italy, Greece, north Africa and Israel), only C. imicola s.s. has been detected
to date (Sebastiani et al. 2001; Linton et al. 2002; Dallas et al. 2003), suggesting that
the species acting as a vector of bluetongue in Turkey is also likely to comprise only
C. imicola s.s.
Prior to the 1980s outbreaks of bluetongue in Europe were restricted to the
Mediterranean Basin and were short and sporadic. In contrast, from 1998 an
unprecedented continuous outbreak of bluetongue has affected central Europe and the
countries of the Mediterranean Basin resulting in the deaths of over 800,000 sheep, and
making this the longest and most economically devastating outbreak of bluetongue in living
history. Countries affected in this outbreak include: Algeria, Bulgaria, Bosnia-Herzegovina,
Cyprus, Croatia, the French island of Corsica, Greece (and the Greek islands of Kos,
Rhodes, Chios, Evia, Lesbos, Skiathos, Skopelos, and Samos), Israel, Italy (and the Italian
islands of Sicily and Sardinia), Kosovo, Macedonia, Morocco, the Spanish islands of
Majorca and Menorca, Tunisia, Turkey, and Yugoslavia (see OIE World Organisation for
Animal Health website, http://www.oie.int).
Common European farmyard species including the C. obsoletus and C. pulicaris species
complexes have long been suspected to act as arbovirus vectors due to their widespread
abundance, and associated high densities around domestic livestock. Members of the C.
obsoletus group are almost impossible to differentiate as adult females and thus references in
the literature to C. obsoletus almost always refer to the C. obsoletus group, which includes C.
chiopterus (Meigen), C. dewulfi Goetghebuer, C. montanus Shakirzjanova, C. obsoletus
(Meigen), C. scoticus Downes and Kettle, and up to two further as yet undescribed species
(Gomulski et al. 2005). At the same time that De Liberato et al. (2005) isolated BTV-2
from non-engorged parous C. obsoletus females caught alongside C. imicola, BTV-2 and
BTV-9 serotypes were isolated from mixed pools of C. obsoletus and C. scoticus collected
from bluetongue outbreak zones in Italy, where no C. imicola was reported (Savini et al.
2005). Carpenter et al. (2006) showed differential bluetongue vector capacity in
populations of C. obsoletus s.l. and Culicoides pulicaris s.l. in the UK, but showed that in
the most susceptible populations of infection prevalence was comparable with that of C.
imicola, using the same viral strain and feeding method.
Given the socioeconomic and veterinary importance of the genus Culicoides and the
obvious pressures to identify the vectors of the ongoing bluetongue outbreak, this paper
attempts to succinctly review published works (written both in English and Turkish) to
comprehensively document the species composition and distribution of Culicoides biting
midges in Turkey for the first time.
Results and discussion
Species composition
Early literature on the Culicoides fauna of Turkey prior to the 1977–81 bluetongue outbreak
is very scanty. The first documented records of Culicoides in Turkey came in the form of two
new species descriptions by Kieffer (1918): C. albonotatus Kieffer later became a junior
synonym of C. fascipennis (Staeger), whilst C. nadayanus Kieffer was subsequently
synonomized with C. circumscriptus Kieffer. A single male specimen of Culicoides parroti
1948 B. Dik et al.
Kieffer from Turkey was reported in a textbook of British haematophagous insects
(Edwards et al. 1939). Later, Oytun (1945) wrote a short paper highlighting the potential
biomedical importance and systematics of the genus Culicoides but he did not report which
species were present in Turkey. In later studies of blood-sucking flies, Mimioglu (1961)
reported collecting a number of Culicoides species from southeastern Turkey, but did not
identify them to species. In 1965, Unat reconfirmed the presence of C. circumscriptus and C.
parroti and Leclercq (1966) reported C. punctatus (Meigen) for the first time, feeding in
high numbers on a horse in Kusadası on the Aegean coast.
Serious efforts to document and identify Culicoides species in Turkey began with Navai’s
PhD thesis (1977), where she documented the presence of 19 Culicoides species from the
Aegean (AE), Mediterranean (ME) and South-Eastern Anatolian (SO) Regions in Turkey
(Table I; Figure 1). With the exception of C. circumscriptus, all 18 others comprised new
country records as follows: C. bulbostylus Khalaf, C. dewulfi Goetghebuer, C. festivipennis
Kieffer (as C. odibilis Austen), C. furcillatus Callot, Kremer and Paradis, C. ibericus
Dzhafarov, C. kurensis Dzhafarov, C. longipennis Khalaf, C. maritimus Kieffer, C.
minutissmus (Zetterstedt) (as C. tugaicus Dzhafarov), C. montanus Schakirzjanova, C.
nubeculosus (Meigen), C. obsoletus (Meigen), C. odiatus Austen, C. pictipennis (Staeger),
C. pulicaris (Linnaeus), C. puncticollis (Becker), C. saevus Kieffer, and C. schultzei
(Enderlein) (Table I). Although a description is written in the original text (Navai 1977,
p 30), Navai inadvertently missed out the single male specimen of C. dewulfi found in
Adana, Turkey from her summary table of species distributions (Navai 1977, p 121–122,
Table 6) and this error has been perpetuated by all later authors who cite, incorrectly, that
18 rather than 19 species were found in this study (e.g. Jennings et al. 1983).
Culicoides surveillance was undertaken in the Aegean (Aydın, Cakmak, Nazilli, and
Salihaga; and Mediterranean Region (Antalya) in direct response to the bluetongue
outbreak of 1978–81 (Jennings et al. 1983). Light traps set in the outbreak zone revealed C.
imicola for the first time, amongst 18 species collected. Besides C. imicola, a further five
were previously unreported in Turkey (C. dzhafarovi Remm, C. gejgelensis Dzhafarov, C.
newsteadi Austen, C. subneglectus Vimmer, and C. vidourlensis Callot, Kremer, Molet and
Bach) (Table I) (Jennings et al. 1983). Dik (1989) collected 14,098 Culicoides comprising
35 species around the city of Konya, in the Central Anatolian Region (CE, Figure 1).
Seventeen of these were new country records as follow: C. achrayi Kettle and Lawson, C.
cataneii Clastrier, C. duddingstoni Kettle and Lawson, C. fagineus Edwards, C. flavipulicaris
Dzhafarov, C. heliophilus Edwards, C. kibunensis Tokunaga, C. pallidicornis Kieffer, C.
picturatus Kremer and Deduit, C. pumilus (Winnertz), C. riethi Kieffer, C. scoticus Downes
and Kettle, C. shaklawensis Khalaf, C. simulator Edwards, C. tauricus Gutsevich, C.
truncorum Edwards, and C. vexans (Staeger) (Table I) (Dik 1989). Collecting in southern
Turkey, Burgu et al. (1992) and Dik (1993) reported 13 and 22 species, respectively,
including new country records for C. sahariensis Kieffer and C. subneglectus Vimmer (Burgu
et al. 1992), and C. denisoni Boorman (Dik 1993) (Table I). In 1994, Yılmaz reported 42
species from the Elazıg province (Eastern Anatolian Region (EA), Figure 1)—the highest
diversity in any publication on Turkish Culicoides—including seven new country records for
C. azerbajdzhanicus Dzhafarov, C. badooshensis Khalaf, C. kolymbiensis Boorman, C.
reconditus Campbell and Pelham-Clinton, C. sejfadinei Dzhafarov, C. semimaculatus
Clastrier, and C. slovacus Orszagh (Table I). Dik (1996) reported C. salinarius Kieffer
and C. turanicus Edwards as new country records amongst 31 species collected from the
Aegean Region (AE) of western Turkey (Aydın, Denizli, Izmir, Kutahya, and Mugla
provinces), completing the current Turkish species list of 57 species. Dik (1997) discussed
Species diversity and distribution of Culicoides 1949
Table I. Faunal list and known distribution of biting midges of the genus Culicoides (Diptera: Ceratopogonidae) of
Turkey based on published data (Kieffer 1918; Edwards et al. 1939; Leclercq 1966; Navai 1977; Jennings et al.
1983; Dik 1989; Burgu et al. 1992; Dik 1993, 1996; Yılmaz 1994; Eren et al. 1995; Yagcı et al. 1999; Eren and
Inci 2002; Tilki and Dik 2003; Uslu 2003).
Culicoides species Locality Map Specimens Source
C. achrayi Kettle and
Lawson, 1955
Baskil EA 7 122R, 13„ Yılmaz 1994
Elazıg EA 22 1R Yılmaz 1994
Kazan CE 34 3R Eren et al. 1995
Kadınhanı CE 31 1R Dik 1989
Keban EA 35 64R, 45„ Yılmaz 1994
Kovancılar EA 37 40R, 8„ Yılmaz 1994
Maden EA 40 1R Yılmaz 1994
Sivrice EA 50 1R Yılmaz 1994
C. azerbajdzhanicus
Dzhafarov, 1962
Baskil EA 7 47R, 33„ Yılmaz 1994
Elazıg EA 22 3R Yılmaz 1994
Keban EA 35 43R, 76„ Yılmaz 1994
Kovancılar EA 37 18R, 43„ Yılmaz 1994
C. badooshensis Khalaf, 1961 Elazıg EA 22 1R, 1„ Yılmaz 1994
Kovancılar EA 37 20R, 13„ Yılmaz 1994
Maden EA 40 2„ Yılmaz 1994
Salihaga AE 46 1R Dik 1996
Soke AE 51 1R Dik 1996
C. bulbostylus Khalaf, 1961 Diyarbakır SO 20 6R, 11„ Navai 1977
Salihaga AE 44 2R Dik 1996
C. cataneiia Clastrier, 1957 Adana ME 1 1R Dik 1993
Aksehir CE 2 2R Dik 1989
Baskil EA 7 21R, 1„ Yılmaz 1994
Beysehir CE 10 1R Dik 1989
Elazıg EA 22 3R Yılmaz 1994
Gemlik MA 25 3R Eren and Inci 2002
Golbası CE 27 5R Eren et al. 1995
Hadim CE 28 2R Dik 1989
Kadınhanı CE 31 1R Dik 1989
Kazan CE 34 8R Eren et al. 1995
Keban EA 35 29R, 6„ Yılmaz 1994
Konya CE 36 7R, 1„ Uslu 2003
Kovancılar EA 37 14R, 4„ Yılmaz 1994
Maden EA 40 6R Yılmaz 1994
Soke AE 51 1R Dik 1996
Simav AE 49 1R Dik 1996
Umurlu AE 54 4R Yagcı et al. 1999
S. Turkeya ME ?b Burgu et al. 1992
C. circumscriptusa Kieffer, 1918 Adana ME 1 1R; 21R, 4„ Navai 1977; Dik 1993
Syn: C. nadayanus Aksehir CE 2 170b Dik 1989
Kieffer, 1918 Anamur ME 3 4R, 2„ Dik 1993
Syn: C. edwardsi Ankara CE 4 3b Jennings et al. 1983
Goetghebuer, 1921 Antalya ME 5 129b; 3R Jennings et al. 1983;
Syn: C. algarum Dik 1993
Kieffer, 1924 Baskil EA 7 283R, 26„ Yılmaz 1994
Syn: C. pictidorsum (as variety of Bergama AE 8 21R, 9„ Dik 1996
C. salicola) Kieffer, 1924 Beysehir CE 10 9b Dik 1989
Syn: C. albonotatus Bornova AE 11 31R, 17„; 44R, Navai 1977; Dik 1996
Vimmer, 1932 53„
1950 B. Dik et al.
Culicoides species Locality Map Specimens Source
Syn: C. albosignatus Bozkır CE 12 32b Dik 1989
Vimmer, 1932 Cigli AE 16 10R, 2„ Navai 1977
Syn: C. polymaculatus Cihanbeyli CE 14 73b Dik 1989
Vimmer, 1932 Cumra CE 10 133b Dik 1989
Syn: C. pulcher Denizli AE 19 2R Dik 1996
Zihali-Sebess, 1934 Elazıg EA 22 61R, 48„ Yılmaz 1994
Syn: C. kirovabadicus Eregli CE 24 15b Dik 1989
Dzhafarov, 1964 Gemlik MA 25 125R, 26„ Eren and Inci 2002
Gokcekısık CE 26 1R Kieffer 1918
Golbası CE 27 61R, 9„ Eren et al. 1995
Hadim CE 28 5b Dik 1989
Iskenderun ME 29 5R Tilki and Dik 2003
Izmir AE 30 51R, 77„ Navai 1977
Karaman CE 33 25b Dik 1989
Kazan CE 34 320R, 20„ Eren et al. 1995
Keban EA 35 112R, 43„ Yılmaz 1994
Kovancılar EA 37 764R, 385„ Yılmaz 1994
Konya CE 36 7b; 409b; 736R,
614„
Jennings et al. 1983;
Dik 1989; Uslu 2003
Kuyucak AE 39 4R Dik 1996
Maden EA 40 10R, 3„ Yılmaz 1994
Mugla AE 42 2R Dik 1996
Pirinclik SO 45 7R, 9„ Navai 1977
Salihaga AE 46 2b; 62R, 5„ Jennings et al. 1983;
Dik 1996
Serik ME 47 37R, 6„ Dik 1993
Seydisehir CE 48 107b Dik 1989
Simav AE 49 37R, 2„ Dik 1996
Sivrice EA 50 11R, 1„ Yılmaz 1994
Soke AE 51 8R Dik 1996
Tarsus ME 52 15R, 5„ Dik 1993
Tire AE 53 1R Navai 1977
Umurlu AE 54 54R, 4„ Yagcı et al. 1999
S. Turkeya ME ?b Burgu et al. 1992
C. denisoni Boorman, 1988 Serik ME 47 3R Dik 1993
C. dewulfi Goetghebuer, 1936 Adana ME 1 1„ Navai 1977
Syn: C. pseudochipterus
Downes and Ketle, 1952
C. duddingstoni Kettle Beysehir CE 10 1„ Dik 1989
and Lawson, 1955 Eregli CE 24 2„ Dik 1989
Karaman CE 33 1„ Dik 1989
Salihaga AE 46 1R, 1„ Dik 1996
Simav AE 49 1„ Dik 1996
C. dzhafarovi Remm, 1967 Adana ME 1 4R Dik 1993
Syn: C. dzhafavovi Callot, Anamur ME 3 2R Dik 1993
Kremer, Molet and
Bach, 1968
Antalya ME 5 13b; 108R Jennings et al. 1983;
Dik 1993
Aksehir CE 2 5R Dik 1989
Baskil EA 7 601R, 263„ Yılmaz 1994
Beysehir CE 10 6R Dik 1989
Cihanbeyli CE 14 20R Dik 1989
Elazıg EA 22 227R, 112„ Yılmaz 1994
Eregli CE 24 8R Dik 1989
Table I. Continued.
Species diversity and distribution of Culicoides 1951
Culicoides species Locality Map Specimens Source
Golbası CE 27 51R, 4„ Eren et al. 1995
Hadim CE 28 6R Dik 1989
Kazan CE 34 46R Eren et al. 1995
Keban EA 35 124R, 130„ Yılmaz 1994
Kovancılar EA 37 392R, 266„ Yılmaz 1994
Kuyucak AE 39 1R Dik 1996
Maden EA 40 58R, 25„ Yılmaz 1994
Mezitli ME 41 2R Dik 1993
Salihaga AE 46 1b; 5R Jennings et al. 1983;
Dik 1996
Serik ME 47 13R Dik 1993
Seydisehir CE 48 7R Dik 1989
Simav AE 49 8R Dik 1996
Sivrice EA 50 224R, 67„ Yılmaz 1994
Tarsus ME 52 16R Dik 1993
Umurlu AE 54 13R Yagcı et al. 1999
C. fagineus Edwards, 1939 Hadim CE 28 1R, 1„ Dik 1989
C. fascipennis (Staeger, 1839) Beysehir CE 10 1R Dik 1989
Syn: C. distictus Kieffer, Dinek SO 20 1R Kieffer 1918
1916 Serik ME 47 1R Dik 1993
Syn: C. albonotatus Kieffer, 1918
Syn: C. turficola Kieffer, 1925
C. festivipennis Kieffer, 1914 Antalya ME 5 3„ Jennings et al. 1983
Syn: C. odibilis Austen, Aksehir CE 2 7b Dik 1989
1921 Baskil EA 7 33R, 48„ Yılmaz 1994
Syn: C. winnertzi Bornova AE 11 8R, 7„; 2R Navai 1977; Dik 1996
Edwards, 1926 Bozkır CE 12 1b Dik 1989
Diyarbakır SO 21 1R Navai 1977
Elazıg EA 22 5R, 1„ Yılmaz 1994
Eregli CE 24 1b Dik 1989
Gemlik MA 25 36R Eren and Inci 2002
Golbası CE 27 3R Eren et al. 1995
Hadim CE 28 3b Dik 1989
Karaman CE 33 2b Dik 1989
Kazan CE 34 8R, 3„ Eren et al. 1995
Keban EA 35 3R, 1„ Yılmaz 1994
Konya CE 36 4b; 360R, 270„ Dik 1989; Uslu 2003
Kovancılar EA 37 144R, 113„ Yılmaz 1994
Maden EA 40 4R, 7„ Yılmaz 1994
Pirinclik SO 45 2R Navai 1977
Simav AE 49 22R, 39„ Dik 1996
Sivrice EA 50 3R Yılmaz 1994
Tire AE 53 1R Navai 1977
Umurlu AE 54 1R Yagcı et al. 1999
C. flavipulicaris Dzhafarov,
1964
Hadim CE 28 3R Dik 1989
Konya CE 36 3R, 6„ Uslu 2003
Seydisehir CE 48 4R Dik 1989
Simav AE 49 1R Dik 1996
Table I. Continued.
1952 B. Dik et al.
Culicoides species Locality Map Specimens Source
C. furcillatus Callot,
Kremer and Paradis, 1962
Baskil EA 7 175R, 8„ Yılmaz 1994
Bornova AE 11 1„ Navai 1977
Elazıg EA 22 1R Yılmaz 1994
Keban EA 35 29R, 7„ Yılmaz 1994
Maden EA 40 3R Yılmaz 1994
Sivrice EA 50 2R Yılmaz 1994
C. gejgelensisa Dzhafarov,
1964
Antalya ME 5 2„, 2R Jennings et al. 1983;
Dik 1993
Aksehir CE 2 64R Dik 1989
Baskil EA 7 95R, 50„ Yılmaz 1994
Bergama AE 8 2R Dik 1996
Beysehir CE 10 1R Dik 1989
Bozkır CE 12 2R Dik 1989
Cihanbeyli CE 14 8R Dik 1989
Elazıg EA 22 37R, 14„ Yılmaz 1994
Gemlik MA 25 51R, 3„ Eren and Inci 2002
Golbası CE 27 2R Eren et al. 1995
Hadim CE 28 22R Dik 1989
Kadınhanı CE 31 3R Dik 1989
Kazan CE 34 34R, 1„ Eren et al. 1995
Keban EA 35 90R, 120„ Yılmaz 1994
Konya CE 36 114R, 70„ Uslu 2003
Kovancılar EA 37 185R, 94„ Yılmaz 1994
Kuyucak AE 39 2R Dik 1996
Maden EA 40 6R, 1„ Yılmaz 1994
Salihaga AE 46 3R Dik 1996
Serik ME 47 1R Dik 1993
Seydisehir CE 48 1R Dik 1989
Simav AE 49 5R Dik 1996
Sivrice EA 50 2R Yılmaz 1994
Soke AE 51 5R Dik 1996
Tarsus ME 52 2R Dik 1993
Umurlu AE 54 13R Yagcı et al. 1999
S. Turkeya ME ?b Burgu et al. 1992
C. heliophilus Edwards, 1921 Aksehir CE 2 1R Dik 1989
Baskil EA 7 237R, 2„ Yılmaz 1994
Hadim CE 28 3R Dik 1989
Kadınhanı CE 31 1R Dik 1989
Keban EA 35 13R Dik 1989
Maden EA 40 2R Yılmaz 1994
Sivrice EA 50 2R Yılmaz 1994
C. ibericus Dzhafarov, 1964 Antalya ME 5 1R; 42R Jennings et al. 1983;
Dik 1993
Baskil EA 7 4R Yılmaz 1994
Bornova AE 11 1R Navai 1977
Elazıg EA 22 5R Yılmaz 1994
Hadim CE 28 3R Dik 1989
Keban EA 35 1R Yılmaz 1994
Serik ME 47 7R Dik 1993
C. imicolaa Kieffer, 1913 Adana ME 1 15R, 2„ Dik 1993
Syn: C. pallidipennis Carter,
Ingram and Macfie 1920
Anamur ME 3 5R, 1„ Dik 1993
Antalya ME 5 95b; 1R Jennings et al. 1983;
Dik 1993
Table I. Continued.
Species diversity and distribution of Culicoides 1953
Culicoides species Locality Map Specimens Source
Syn: C. iraqensis Khalaf,
1957
Aydın AE 6 13b Jennings et al. 1983
Bornova AE 11 1R Dik 1996
Syn: C. minutus Sen and
Das Gupta, 1959
Cakmak AE 15 16b Jennings et al. 1983
Iskenderun ME 29 53R, 3„ Tilki and Dik 2003
Syn: C. pseudoturgidus
Das Gupta, 1962
Mezitli ME 41 23R, 1„ Dik 1993
Nazili AE 44 13b Jennings et al. 1983
Salihaga AE 46 479b Jennings et al. 1983
Tarsus ME 52 10R Dik 1993
Umurlu AE 54 9R Yagcı et al. 1999
S. Turkeya ME ?b Burgu et al. 1992
C. kibunensis Tokunaga, 1937 Baskil EA 7 48R, 43„ Yılmaz 1994
Syn: C. cubitalis
Edwards, 1939
Elazıg EA 22 25R, 33„ Yılmaz 1994
Gemlik MA 25 21R Eren and Inci 2002
Syn: C. ponkikiri Kono
and Takahasi, 1940
Golbası CE 27 10R Eren et al. 1995
Keban EA 35 51R, 67„ Yılmaz 1994
Syn: C. sitinohensis
Okada, 1941
Hadim CE 28 3R Dik 1989
Kazan CE 34 30R Eren et al. 1995
Konya CE 36 3R Uslu 2003
Kovancılar EA 37 4R, 3„ Yılmaz 1994
Maden EA 40 10R, 19„ Yılmaz 1994
Salihaga AE 46 3R, 2„ Dik 1996
Sivrice EA 50 6R, 1„ Yılmaz 1994
C. kolymbiensis Boorman, 1988 Baskil EA 7 110R, 78„ Yılmaz 1994
Elazıg EA 22 9R, 3„ Yılmaz 1994
Keban EA 35 44R, 11„ Yılmaz 1994
Kovancılar EA 37 70R, 60„ Yılmaz 1994
C. kurensis Dzhafarov, 1960 Antalya ME 5 348b Jennings et al. 1983
Bespınar SO 9 12R Navai 1977
Pirinclik SO 45 5„ Navai 1977
Salihaga AE 46 1b Jennings et al. 1983
C. longipennisa Khalaf, 1957 Adana ME 1 2R; 2R Navai 1977; Dik 1993
Syn: C. flavisimilis
Dzhafarov, 1964
Anamur ME 3 4R Dik 1993
Antalya ME 5 245b; 6R Jennings et al. 1983;
Dik 1993
Baskil EA 7 475R, 223„ Yılmaz 1994
Bornova AE 11 5R; 2R Navai 1977; Dik 1996
Cakmak AE 15 7b Jennings et al. 1983
Cihanbeyli CE 14 1b Dik 1989
Elazıg EA 22 126R, 29„ Yılmaz 1994
Gemlik MA 25 20R Eren and Inci 2002
Hadim CE 28 4b Dik 1989
Iskenderun ME 29 5012R, 145„ Tilki and Dik 2003
Kazan CE 34 5R Eren et al. 1995
Keban EA 35 236R, 139„ Yılmaz 1994
Konya CE 36 14R, 10„ Uslu 2003
Kovancılar EA 37 836R, 254„ Yılmaz 1994
Kuyucak AE 39 1R Dik 1996
Maden EA 40 77R, 48„ Yilmaz 1994
Nazilli AE 44 4b Jennings et al. 1983
Salihaga AE 46 23b; 35R, 8„ Jennings et al. 1983;
Dik 1996
Serik ME 47 25R, 1„ Dik 1993
Simav AE 49 20R, 8„ Dik 1996
Table I. Continued.
1954 B. Dik et al.
Culicoides species Locality Map Specimens Source
Sivrice EA 50 9R, 3„ Yılmaz 1994
Tarsus ME 52 35R, 12„ Dik 1993
Umurlu AE 54 17R Yagcı et al. 1999
S. Turkeya ME ?b Burgu et al. 1992
C. maritimus Kieffer, 1924 Ankara CE 4 1b Jennings et al. 1983
Syn: C. submaritimus
Dzhafarov, 1962
Antalya ME 5 51b Jennings et al. 1983
Aydın AE 6 1b Jennings et al. 1983
Cakmak AE 15 1b Jennings et al. 1983
Bornova AE 11 10R, 4„ Navai 1977
Bozkır CE 12 22b Dik 1989
Cumra CE 18 23b Dik 1989
Gemlik MA 25 25R, 3„ Eren and Inci 2002
Kadınhanı CE 31 2b Dik 1989
Keban EA 35 1R Yılmaz 1994
Konya CE 36 171b; 6„ Dik 1989; Uslu 2003
Maden EA 40 7R, 8„ Yılmaz 1994
Pirinclik SO 45 2R, 4„ Navai 1977
Salihaga AE 46 1R Dik 1996
Seydisehir CE 48 7b Dik 1989
C. minutissmus (Zetterstedt), 1855 Bornova AE 11 2R, 9„ Navai 1977
Syn: C. nanulus Kieffer, 1919
Syn: C. albihalter Kieffer, 1919
Syn: C. bychowskyi Dzhafarov, 1964
Syn: C. tugaicus Dzhafarov, 1960
C. montanus Schakirzjanova, 1962 Diyarbakır SO 21 3R Navai 1977
C. newsteadi Austen, 1921 Aksehir CE 2 6b Dik 1989
Syn: C. biclavatus Kieffer, 1924 Ankara CE 4 1b Jennings et al. 1983
Syn: C. halophilus Kieffer, 1924 Antalya ME 5 67b; 1R Jennings et al. 1983;
Syn: C. edwardsi (as variety of
C. pulicaris) Goetghebuer, 1933
Dik 1993
Baskil EA 7 24R, 6„ Yılmaz 1994
Syn: C. edwardsianus
(as variety of C. pulicaris)
Goetghebuer, 1933
Beysehir CE 10 19b Dik 1989
Cakmak AE 15 2b Jennings et al. 1983
Cihanbeyli CE 14 31b Dik 1989
Cumra CE 18 16b Dik 1989
Elazıg EA 22 16R, 1„ Yılmaz 1994
Eregli CE 24 21b Dik 1989
Hadim CE 28 12b Dik 1989
Kadınhanı CE 31 28b Dik 1989
Karaman CE 33 17b Dik 1989
Kazan CE 34 3R Eren et al. 1995
Keban EA 35 11R Yılmaz 1994
Konya CE 36 2b Jennings et al. 1983
Kovancılar EA 37 146R, 10„ Yılmaz 1994
Maden EA 40 5R, 1„ Yılmaz 1994
Nazilli AE 44 14b Jennings et al. 1983
Salihaga AE 46 68b Jennings et al. 1983
Seydisehir CE 48 78b Dik 1989
Sivrice EA 50 23R, 3„ Yılmaz 1994
Soke AE 51 5R Dik 1996
Umurlu AE 54 7R Yagcı et al. 1999
Table I. Continued.
Species diversity and distribution of Culicoides 1955
Culicoides species Locality Map Specimens Source
C. nubeculosus(Meigen), 1830 Adana ME 1 405R, 62„ Dik 1993
Syn: C. puncticollis
Goetghebuer, 1912
Antalya ME 5 3R Dik 1993
Bergama AE 8 1R, 4„ Dik 1996
Syn: C. punctaticollis
Goetghebuer, 1912
Bornova AE 11 1R, 8„ Navai 1977
Bozkır CE 12 2b Dik 1989
Cihanbeyli CE 14 153b Dik 1989
Cumra CE 18 12b Dik 1989
Denizli AE 19 1R Dik 1996
Eregli CE 24 45b Dik 1989
Elazıg EA 22 94R, 119„ Yılmaz 1994
Gemlik MA 25 6R Eren and Inci 2002
Kadınhanı CE 31 15b Dik 1989
Karaman CE 33 41b Dik 1989
Konya CE 36 41R, 39„ Uslu 2003
Kovancılar EA 37 16R, 10„ Yılmaz 1994
Maden EA 40 1R Yılmaz 1994
Mezitli ME 41 18R, 1„ Dik 1993
Seydisehir CE 48 6b Dik 1989
Simav AE 49 71R, 26„ Dik 1996
Sivrice EA 50 3R, 4„ Yılmaz 1994
Tarsus ME 52 32R, 2„ Dik 1993
C. obsoletusa (Meigen), 1818 Adana ME 1 1R Dik 1993
Syn: Ceratopogon varius
(Winnertz), 1852
Aksehir CE 2 7b Dik 1989
Antalya ME 5 1R; 2R Jennings et al. 1983;
Syn: C. obscuripes (as variety
of C. varius) Santos Abreu, 1918
Dik 1993
Aydın AE 6 1R Jennings et al. 1983
Syn: C. lacteinervis Kieffer, 1919 Baskil EA 7 5R, 2„ Yılmaz 1994
Syn: C. rivicola Kieffer, 1921 Bornova AE 11 3R, 1„ Navai 1977
Syn: C. clavatus Kieffer, 1921 Beysehir CE 10 1b Dik 1989
Syn: C. heterocerus Kieffer, 1921 Elazıg EA 22 3R Yılmaz 1994
Syn: C. pegobius Kieffer, 1922 Eregli CE 24 1b Dik 1989
Syn: C. kabyliensis Kieffer, 1922 Gemlik MA 25 10R Eren and Inci 2002
Syn: C. concitus Kieffer, 1922 Golbası CE 27 46R, 3„ Eren et al. 1995
Syn: C. intermedius Okada, 1941 Iskenderun ME 29 3R Tilki and Dik 2003
Syn: C. intermedius
(as variety of C. obsoletus)
Okada, 1941
Kadınhanı CE 31 3b Dik 1989
Karaman CE 33 2b Dik 1989
Kazan CE 34 109R Eren et al. 1995
Syn: C. sintrensis
Cambournac, 1956
Keban EA 35 2R, 2„ Yılmaz 1994
Kovancılar EA 37 10R Yılmaz 1994
Syn: C. seimi Shevchenko, 1967 Kuyucak AE 39 3R Dik 1996
Maden EA 40 51R, 1„ Yılmaz 1994
Mugla AE 42 2R Dik 1996
Nazilli AE 44 1R Jennings et al. 1983
Salihaga AE 46 2R; 16R, 1„ Jennings et al. 1983;
Dik 1996
Serik ME 47 4R Dik 1993
Seydisehir CE 48 3b Dik 1989
Simav AE 49 12R, 1„ Dik 1996
Tarsus ME 52 1R Dik 1993
Umurlu AE 54 9R Yagcı et al. 1999
S. Turkeya ME ?b Burgu et al. 1992
Table I. Continued.
1956 B. Dik et al.
Culicoides species Locality Map Specimens Source
C. odiatusa Austen, 1921 Adana ME 1 25R 7„ Dik 1993
Syn: C. niger (as variety of
C. pallidicornis) Root and
Hoffman, 1937
Aksehir CE 2 3b Dik 1989
Anamur ME 3 5R,1„ Dik 1993
Antalya ME 5 2304b; 3R Jennings et al. 1983;
Syn: C. indistinctus Khalaf, 1961 Dik 1993
Syn: C. lailae Khalaf, 1961 Aydın AE 6 14b Jennings et al. 1983
Syn: C. kureksthaicus
Dzhafarov, 1962
Baskil EA 7 455R, 150„ Yılmaz 1994
Bergama AE 8 5R Dik 1996
Syn: C. conicus Remm
and Zhagolev, 1968
Bespınar SO 9 1R Navai 1977
Beysehir CE 10 3b Dik 1989
Bornova AE 11 22R, 1„; 1R Navai 1977; Dik 1996
Cakmak AE 15 89b Jennings et al. 1983
Cigli AE 16 1R Navai 1977
Cihanbeyli CE 14 22b Dik 1989
Cumra CE 18 5b Dik 1989
Denizli AE 19 4R Dik 1996
Diyarbakır SO 21 1„ Navai 1977
Eregli CE 24 3b Dik 1989
Elazıg EA 22 73R, 23„ Yılmaz 1994
Golbası CE 27 26R Eren et al. 1995
Hadim CE 28 87b Dik 1989
Kazan CE 34 32R, 1„ Eren et al. 1995
Keban EA 35 237R, 252„ Yılmaz 1994
Konya CE 36 10R, 2„ Uslu 2003
Kovancılar EA 37 773R, 327„ Yılmaz 1994
Maden EA 40 151R, 33„ Yılmaz 1994
Mezitli ME 41 1R Dik 1993
Mugla AE 42 2R Dik 1996
Nazilli AE 44 27b Jennings et al. 1983
Pirinclik SO 45 3R Navai 1977
Salihaga AE 46 1981b; 54R, 1„ Jennings et al. 1983;
Dik 1996
Serik ME 47 19R, 2„ Dik 1993
Seydisehir CE 48 17b Dik 1989
Simav AE 49 2R Dik 1996
Sivrice EA 50 24R, 14„ Yılmaz 1994
Soke AE 51 1R Dik 1996
Tarsus ME 52 5R, 2„ Dik 1993
Tire AE 53 8R, 1„ Navai 1977
S. Turkeya ME ?b Burgu et al. 1992
C. pallidicornis Kieffer, 1919 Baskil EA 7 283R, 25„ Yılmaz 1994
Syn: C. susae Kieffer, 1919 Elazıg EA 22 3R, 1„ Yılmaz 1994
Syn: C. dileucus Kieffer, 1921 Keban EA 35 172R, 174„ Yılmaz 1994
Syn: C. brunneiscutellatus
(as variety of C. pallidicornis)
Zilahi-Sebess, 1933
Kovancılar EA 37 8R, 8„ Yılmaz 1994
Hadim CE 28 11R Dik 1989
Maden EA 40 4R Yılmaz 1994
Syn: C. bruneoscutellatus
(as variety of C. pallidicornis)
Zilahi-Sebess, 1934
Sivrice EA 50 1R, 1„ Yılmaz 1994
Syn: C. niger Root and
Hoffman, 1937
C. parrotic Kieffer, 1922 Ankara CE 4 1„ Edwards et al. 1939
Baskil EA 7 7R, 3„ Yılmaz 1994
Elazıg EA 22 10R, 7„ Yılmaz 1994
Table I. Continued.
Species diversity and distribution of Culicoides 1957
Culicoides species Locality Map Specimens Source
Konya CE 36 2R, 1„ Uslu 2003
Maden EA 40 1R, 1„ Yılmaz 1994
Nazilli AE 44 1R Jennings et al. 1983
Kovancılar EA 37 5R, 8„ Yılmaz 1994
Sivrice EA 50 2R, 1„ Yılmaz 1994
Simav AE 49 4R Dik 1996
C. pictipennis (Staeger), 1839 Baskil EA 7 1„ Yılmaz 1994
Syn: Ceratopogon arcuatus
(Winnertz), 1852
Bornova AE 11 28R, 46„ Navai 1977
Cigli AE 16 2R Navai 1977
Syn: C. guttularis Kieffer, 1919 Elazıg EA 22 12R, 4„ Yılmaz 1994
Syn: C. maculatus
(as variety of C. pictipennis)
Zilahi-Sebess, 1936
Maden EA 40 61R, 41„ Yılmaz 1994
Pirinclik SO 45 2R, 18„ Navai 1977
Sivrice EA 50 1R Yılmaz 1994
Syn: C. subgrisescens
Dzhafarov, 1964
Tire AE 53 1R, 2„ Navai 1977
Syn: C. achkamalicus
Dzhafarov, 1964
Syn: C. luganicus
Shevchenko, 1972
C. picturatus Kremer and
Deduit, 1961
Aksehir CE 2 13R Dik 1989
Baskil EA 7 365R, 13„ Yilmaz 1994
Elazıg EA 22 90R, 10„ Yılmaz 1994
Keban EA 35 35R, 20„ Yılmaz 1994
Hadim CE 28 89R Dik 1989
Kadınhanı CE 31 57R Dik 1989
Kovancılar EA 37 10R, 3„ Yılmaz 1994
Maden EA 40 115R, 14„ Yılmaz 1994
Sivrice EA 50 252R Yılmaz 1994
C. pulicarisa (Linnaeus), 1758 Anamur ME 3 1R Dik 1993
Syn: C. setosinervis Kieffer, 1913 Aksehir CE 2 78b Dik 1989
Syn: C. pullatus Kieffer, 1915 Baskil EA 7 27R, 5„ Yılmaz 1994
Syn: C. stephensi Carter, 1916 Beysehir CE 10 3b Dik 1989
Syn: C. cinerellus Kieffer, 1919 Bornova AE 11 16R, 10„ Navai 1977
Syn: C. quinquepunctatus
Goetghebuer, 1921
Cigli AE 16 7R 3„ Navai 1977
Cihanbeyli CE 14 6b Dik 1989
Syn: C. flaviplumus Kieffer, 1924 Elazıg EA 22 11R Yılmaz 1994
Syn: C. sawamotoi Kono and
Takahasi, 1940
Gemlik MA 25 2R Eren and Inci 2002
Kadınhanı CE 31 5b Dik 1989
Hadim CE 28 27b Dik 1989
Keban EA 35 15R, 21„ Yılmaz 1994
Konya CE 36 7b; 1R Dik 1989; Uslu 2003
Maden EA 40 11R Yılmaz 1994
Kazan CE 34 2R, 1„ Eren et al. 1995
Kovancılar EA 37 71R, 8„ Yılmaz 1994
Pirinclik SO 45 5R, 4„ Navai 1977
Seydisehir CE 48 4b Dik 1989
Simav AE 49 1R Dik 1996
Sivrice EA 50 3R, 1„ Yılmaz 1994
Soke AE 51 1„ Dik 1996
Umurlu AE 54 1R Yagcı et al. 1999
S. Turkeya ME ?b Burgu et al. 1992
Table I. Continued.
1958 B. Dik et al.
Culicoides species Locality Map Specimens Source
C. pumilus (Winnertz), 1852 Antalya ME 5 1R Dik 1993
Aksehir CE 2 2R Dik 1989
Baskil EA 7 66R, 46„ Yılmaz 1994
Cihanbeyli CE 14 2R Dik 1989
Elazıg EA 22 19R, 9„ Yılmaz 1994
Hadim CE 28 5R Dik 1989
Kadınhanı CE 31 1R Dik 1989
Keban EA 35 1261R, 1000„ Yılmaz 1994
Kovancılar EA 37 7R, 7„ Yılmaz 1994
Maden EA 40 28R, 6„ Yılmaz 1994
Seydisehir CE 48 1R Dik 1989
Sivrice EA 50 7R Yılmaz 1994
C. punctatus (Meigen), 1804 Ankara CE 4 3b Jennings et al. 1983
Syn: C. punctatus Latreille, 1809 Antalya ME 5 38b; 3R Jennings et al. 1983;
Syn: C. ocellaris Kieffer, 1921 Dik 1993
Syn: C. kasachstanicus
(as subspecies of C. pulicaris)
Shakirzjanova, 1963
Aydın AE 6 1b Jennings et al. 1983
Aksehir CE 2 786b Dik 1989
Baskil EA 7 5R, 4„ Yılmaz 1994
Syn: C. remmi
Damian-Georgescu, 1972
Beysehir CE 10 197b Dik 1989
Bornova AE 11 2R Dik 1996
Bozkır CE 12 41b Dik 1989
Cakmak AE 15 62b Jennings et al. 1983
Cihanbeyli CE 14 152b Dik 1989
Cumra CE 18 46b Dik 1989
Elazıg EA 22 1R Yılmaz 1994
Eregli CE 24 153b Dik 1989
Gemlik MA 25 74R, 5„ Eren and Inci 2002
Golbası CE 27 246R, 14„ Eren et al. 1995
Hadim CE 28 38b Dik 1989
Kadınhanı CE 31 107b Dik 1989
Karaman CE 33 89b Dik 1989
Kazan CE 34 412R, 10„ Eren et al. 1995
Keban EA 35 12R, 4„ Yılmaz 1994
Konya
Kovancılar
CE 36 1b; 272b Jennings et al. 1983;
Dik 1989
Kusadası EA 37 132R, 16„ Yılmaz 1994
Kuyucak AE 38 Numerous R Leclercq 1966
Maden AE 39 3R Dik 1996
Salihaga EA 40 10R, 3„ Yılmaz 1994
Serik AE 46 9b; 13R Jennings et al. 1983;
Dik 1996
Seydisehir ME 47 6R Dik 1993
Simav CE 48 317b Dik 1989
Sivrice AE 49 4R Dik 1996
Umurlu EA 50 1002R, 73„ Yılmaz 1994
AE 54 7R, 1„ Yagcı et al. 1999
C. puncticollisa (Becker, 1903) Adana ME 1 6R, 3„; 7R Jennings et al. 1983;
Syn: Ceratopogon algecirensis (as
form of pulicaris)(Strobl), 1900
Dik 1993
Antalya ME 5 4b Jennings et al. 1983
Syn: Ceratopogon algecirensis (as
form of pulicaris)(Strobl), 1900
Aydın AE 6 5b Jennings et al. 1983
Baskil EA 7 5R Yılmaz 1994
Syn: C. impressus Kieffer, 1918 Bespınar SO 9 10R, 5„ Navai 1977
Syn: C. distigma Kieffer, 1922 Bornova AE 11 1„; 1R Navai 1977; Dik 1996
Syn: C. donatieni Kieffer, 1922 Cigli AE 16 2R, 1„ Jennings et al. 1983
Table I. Continued.
Species diversity and distribution of Culicoides 1959
Culicoides species Locality Map Specimens Source
Syn: C. sciniphes Kieffer, 1925 Cihanbeyli CE 14 624b Dik 1989
Syn: C. bipunctatus Vimmer, 1932 Cumra CE 18 60b Dik 1989
Syn: C. tripunctatus Vimmer, 1932 Elazıg EA 22 17R, 4„ Yılmaz 1994
Syn: C. wenigi Vimmer, 1932 Eregli CE 24 25b Dik 1989
Syn: C. flavitarsis Vimmer, 1932 Golbası CE 27 2R, 2„ Eren et al. 1995
Syn: C. griseovittatus Vimmer,
1932
Kadınhanı CE 31 4b Dik 1989
Karaman CE 33 2b Dik 1989
Syn: C. luteosignatus Vimmer,
1932
Kazan CE 34 2768R, 671„ Eren et al. 1995
Keban EA 35 5R Yılmaz 1994
Syn: C. vavrai Vimmer, 1932 Konya CE 36 137R, 35„ Uslu 2003
Kovancılar EA 37 70R, 70„ Yılmaz 1994
Konya CE 36 44b Jennings et al. 1983
Nazilli AE 44 1b Jennings et al. 1983
Salihaga AE 46 1b Jennings et al. 1983
Seydisehir CE 48 3b Dik 1989
Simav AE 49 220R, 78„ Dik 1996
Sivrice EA 50 21R, 3„ Yılmaz 1994
Soke AE 51 13R Dik 1996
Umurlu AE 54 8R Yagcı et al. 1999
S. Turkeya ME ?b Burgu et al. 1992
C. reconditus Campbell and
Pelham- Clinton, 1960
Baskil EA 7 8R, 5„ Yılmaz 1994
Maden EA 40 10R, 5„ Yılmaz 1994
Syn: C. riouxi Callot and
Kremer, 1961
Kazan CE 34 8R Eren et al. 1995
Keban EA 35 1R Yılmaz 1994
Kovancılar EA 37 2R Yılmaz 1994
Sivrice EA 50 1R Yılmaz 1994
C. riethi Kieffer, 1914 Bergama AE 8 1R Dik 1996
Syn: C. cordatus Kieffer, 1921 Bozkır CE 12 19b Dik 1989
Syn: C. crassiforceps Kieffer, 1924 Cihanbeyli CE 14 41b Dik 1989
Cumra CE 18 489b Dik 1989
Kadınhanı CE 31 5b Dik 1989
Karaman CE 33 320b Dik 1989
Kazan CE 34 2R Eren et al. 1995
Konya CE 36 7327b; 8R, 43„ Dik 1989; Uslu 2003
Mut ME 43 1„ Dik 1996
Salihaga AE 46 1R Dik 1996
Seydisehir CE 48 2b Dik 1989
Tarsus ME 52 1R Dik 1993
C. saevus Kieffer, 1922 Adana ME 1 2R Dik 1993
Syn: C. drenskii Zilahi-Sebess,
1934
Baskil EA 7 111R, 10„ Yılmaz 1994
Bespınar SO 9 10R Navai 1977
Syn: C. puncticeps Goetghebuer,
1934
Bornova AE 11 16R, 3„ Navai 1977
Cihanbeyli CE 14 7R Dik 1989
Syn: C. micromaculithorax
Khalaf, 1957
Eregli CE 24 2R Dik 1989
Cigli AE 16 12R 3„ Navai 1977
Diyarbakır SO 21 4R Navai 1977
Elazıg EA 22 7R, 1„ Yılmaz 1994
Hadim CE 28 1R Dik 1989
Kadınhanı CE 31 1R Dik 1989
Keban EA 35 328R, 42„ Yılmaz 1994
Konya CE 36 2R Dik 1989
Kovancılar EA 37 101R, 42„ Yılmaz 1994
Maden EA 40 6R, 1„ Yılmaz 1994
Table I. Continued.
1960 B. Dik et al.
Culicoides species Locality Map Specimens Source
Mezitli ME 41 1R Dik 1993
Mut ME 42 43R Dik 1993
Pirinclik SO 45 4R, 2„ Navai 1977
Salihaga AE 46 21„ Jennings et al. 1983
Serik ME 47 3R Dik 1993
Simav AE 49 1R Dik 1996
Soke AE 51 2R Dik 1996
Tarsus ME 52 2R Dik 1993
Tire AE 53 7R Navai 1977
C. sahariensisa Kieffer, 1923 Mugla AE 42 1R Dik 1996
Syn: C. baghdadensis Khalaf,
1957
Simav AE 49 3R, 2„ Dik 1996
Salihaga AE 46 1R Dik 1996
Syn: C. coluzzii Callot, Kremer
and Bailly-Choumara, 1970
Tarsus ME 52 1R Dik 1993
S. Turkeya ME ?b Burgu et al. 1992
C. salinarius Kieffer, 1914 Bornova AE 11 1R Dik 1996
Syn: C. halobius Kieffer, 1914 Konya CE 36 1R Uslu 2003
Syn: C. meinerti Kieffer, 1915
Syn: C. punctatidorsum Kieffer, 1924
C. schultzeiad (Enderlein), 1908 Adana ME 1 4R; 259R, 13„ Navai 1977; Dik 1993
Anamur ME 3 16R Dik 1993
Antalya ME 5 1251b Jennings et al. 1983
Aydın AE 6 1b Jennings et al. 1983
Baskil EA 7 1„ Yılmaz 1994
Eregli CE 24 2R Dik 1989
Iskenderun ME 29 48R, 24„ Tilki and Dik 2003
Konya CE 36 2R, 1„ Uslu 2003
Kuyucak AE 39 7R, 5„ Dik 1996
Mezitli ME 41 340R, 24„ Dik 1993
Nazilli AE 44 435b Jennings et al. 1983
Salihaga AE 46 734b Jennings et al. 1983
Tarsus ME 52 22R, 6„ Dik 1993
Umurlu AE 54 2R Yagcı et al. 1999
S. Turkeya ME ?b Burgu et al. 1992
C. scoticus Downes and Kettle, 1952 Hadım CE 28 1R Dik 1989
Salihaga AE 46 2R Dik 1996
C. sejfadinei Dzhafarov, 1958 Baskil EA 7 32R, 4„ Yılmaz 1994
Syn: C. flavidus Dzhafarov, 1959 Keban EA 35 1R, 1„ Yılmaz 1994
Kovancılar EA 37 14R, 16„ Yılmaz 1994
Maden EA 40 1R Yılmaz 1994
Sivrice EA 50 1R Yılmaz 1994
C. semimaculatus Clastrier, 1958 Maden EA 40 1R Yılmaz 1994
C. shaklawensisa Khalaf, 1957 Aksehir CE 2 6R Dik 1989
Syn: C. caspius Gutsevich, 1959 Baskil EA 7 22R, 13„ Yılmaz 1994
Elazıg EA 22 15R, 10„ Yılmaz 1994
Gemlik MA 25 3R Eren and Inci 2002
Kazan CE 34 1R, 1„ Eren et al. 1995
Keban EA 35 47R, 30„ Uslu 2003
Konya CE 36 119R, 140„ Uslu 2003
Kovancılar EA 37 312R, 78„ Yılmaz 1994
Maden EA 40 5R, 1„ Yılmaz 1994
Salihaga AE 46 3R Dik 1996
Table I. Continued.
Species diversity and distribution of Culicoides 1961
Culicoides species Locality Map Specimens Source
Umurlu AE 54 1R Yagcı et al. 1999
S. Turkeya ME ?b Burgu et al. 1992
C. simulator Edwards, 1939 Aksehir CE 2 44b Dik 1989
Baskil EA 7 9R, 7„ Yılmaz 1994
Bozkır CE 12 7b Dik 1989
Elazıg EA 22 9R, 1„ Yılmaz 1994
Golbası CE 27 7R Eren et al. 1995
Kazan CE 34 11R Eren et al. 1995
Keban EA 35 2R Yılmaz 1994
Maden EA 40 257R, 60„ Yılmaz 1994
Kovancılar EA 37 15R, 9„ Yılmaz 1994
Konya CE 36 234b Dik 1989
Seydisehir CE 48 2b Dik 1989
Sivrice EA 50 1R Yılmaz 1994
C. slovacus Orszagh, 1969 Baskil EA 7 2R Yılmaz 1994
C. subfasciipennis Kieffer, 1919 Aksehir CE 2 64R Dik 1989
Syn: C. analis (as variety of C.
subfasciipennis) Kieffer, 1925
Antalya ME 5 6R, 1„ Jennings et al 1983
Baskil EA 7 9R Yılmaz 1994
Beysehir CE 10 5R Dik 1989
Cihanbeyli CE 14 7R Dik 1989
Elazıg EA 22 2R Yılmaz 1994
Hadim CE 28 37R Dik 1989
Kadınhanı CE 31 55R Dik 1989
Keban EA 35 7R Yılmaz 1994
Konya CE 36 2R; 1R, 1„ Dik 1989; Uslu 2003
Kuyucak AE 39 1R Dik 1996
Maden EA 40 4R, 1„ Yılmaz 1994
Salihaga AE 46 1R Dik 1996
Seydisehir CE 48 1R Dik 1989
Simav AE 49 3R Dik 1996
Sivrice EA 50 1R Yılmaz 1994
C. subneglectusa Vimmer, 1932 Baskil EA 7 1R, 2„ Yılmaz 1994
Elazıg EA 22 2R, 1„ Yılmaz 1994
Keban EA 35 8R, 5„ Yılmaz 1994
Kovancılar EA 37 178R, 330„ Yılmaz 1994
Maden EA 40 1R, 2„ Yılmaz 1994
S. Turkeya ME ?b Burgu et al. 1992
C. tauricus Gutsevich, 1959 Aksehir CE 2 4R Dik 1989
Antalya ME 5 1R Dik 1993
Baskil EA 7 53R, 1„ Yılmaz 1994
Bergama AE 8 1R, 1„ Dik 1996
Serik ME 47 83R Dik 1993
Elazıg EA 22 3R Yılmaz 1994
Hadim CE 28 10R Dik 1989
Keban EA 35 6R Yılmaz 1994
Kovancılar EA 37 45R, 1„ Yılmaz 1994
Maden EA 40 2R Yılmaz 1994
Simav AE 49 2R Dik 1996
Sivrice EA 50 2R Yılmaz 1994
C. truncorum Edwards, 1939 Baskil EA 7 17R, 1„ Yılmaz 1994
Syn: C. sylvarum Callot
and Kremer, 1961
Beysehir CE 10 1R Dik 1989
Elazıg EA 22 24R Yılmaz 1994
Table I. Continued.
1962 B. Dik et al.
the morphology and ecological parameters of the Turkish fauna in relation to disease
transmission. Total numbers of reported species per study are given in relation to currently
accepted nomenclature (Borkent and Wirth 1997).
Although other authors have published Culicoides studies and added valuable data to
species distributions across Turkey (Table I; Figure 1), their studies did not report novel
Culicoides: Eren et al. (1995) (Ankara province, Central Anatolian Region (CE), 19 spp.),
Yagcı et al. (1999) (Aydin, Aegean Region (AE), 14 spp.), Eren and Inci (2002) (Bursa
province, Marmara Region (MA), 12 spp.), Tilki and Dik (2003) (Iskenderun,
Mediterranean Region (ME), 5 spp.), and Uslu (2003) (Konya province, Central
Anatolian Region (CE), 18 spp.) (Table I; Figure 1).
Species distribution and relative abundance
Culicoides sampling in Turkey has been sporadic and is far from comprehensive. Adult
Culicoides have been reported from six of the seven recognized Regions of Turkey
(Figure 1). No records are available on the Culicoides fauna of the Black Sea Region in the
north of the country. Distribution records included in this review cover the Central
Anatolian Region (15 localities in four of 15 provinces), Aegean Region (16 localities in five
of eight provinces), Mediterranean Region (11 localities in five of eight provinces), Eastern
Anatolian Region (six localities in one of 15 provinces) and Southeastern Anatolian
Culicoides species Locality Map Specimens Source
Keban EA 35 5R Yılmaz 1994
Golbası CE 27 46R, 3„ Eren et al. 1995
Hadim CE 28 2R Dik 1989
Kovancılar EA 37 1R Yılmaz 1994
Maden EA 40 17R Yılmaz 1994
Simav AE 49 1R Dik 1996
Sivrice EA 50 222R Yılmaz 1994
C. turanicus Gutsevich and
Smatov, 1971
Salihaga AE 46 3R Dik 1996
C. vexans (Staeger), 1839 Aksehir CE 2 1R Dik 1989
Syn: Palpomyia pungens
(Kieffer, 1901)
Kadınhanı CE 31 2R Dik 1989
Keban EA 35 1R Yılmaz 1994
Syn: C. perpungens Kieffer, 1919 Maden EA 40 1„ Yılmaz 1994
Syn: C. ajbassovi Shakirzjanova,
1962
Sivrice EA 50 1R Yılmaz 1994
C. vidourlensis Callot, Kremer,
Molet and Bach, 1968
Baskil EA 7 106R, 10„ Yılmaz 1994
Elazıg EA 22 66R, 50„ Yılmaz 1994
Keban EA 35 137R, 78„ Yılmaz 1994
Kovancılar EA 37 14R, 2„ Yılmaz 1994
Maden EA 40 1„ Yılmaz 1994
Salihaga AE 46 1R Dik 1996
Serik ME 47 1Re Dik 1993
Sivrice EA 50 2R Yılmaz 1994
aBurgu et al. (1992) reported several species from the southern Turkish localities of Aydın, Antalya, Ceylanpınar,
Cukurova, Erdemli, and Turkoglu. Exact locality data and numbers/sex of samples were not detailed. bSex
unspecified. cLocality originally listed as Koktshe Kissik in Kieffer (1918). dSpecies identified as C. schultzei may
comprise a species complex. eIdentification verified after publication.
Table I. Continued.
Species diversity and distribution of Culicoides 1963
Region (five localities in two of seven provinces), and a single locality (in one of 12
provinces) of the Marmara Region (Kieffer 1918; Edwards et al. 1939; Leclercq 1966;
Navai 1977; Jennings et al. 1983; Dik 1989, 1993, 1996; Burgu et al. 1992; Yılmaz 1994;
Eren et al. 1995; Yagcı et al. 1999; Eren and Inci 2002; Tilki and Dik 2003; Uslu 2003)
(Figure 1).
Culicoides circumscriptus, C. festivipennis, C. maritimus, and C. pulicaris are the most
widespread species, being found in all six provinces sampled, with C. cataneii, C. gejgelensis,
C. longipennis, C. nubeculosus, C. obsoletus, C. odiatus, C. puncticollis, C. saevus, and C.
shaklawensis being reported in five provinces, respectively (Table I). The least common
species are C. denisoni, C. dewulfi, C. fagineus, C. minutissmus, C. montanus, C. semima-
culatus, C. slovacus, and C. turanicus which are represented from only one locality (Table I).
Only a single male and a single female specimen represent C. dewulfi and C. semimaculatus,
respectively (Navai 1977; Yılmaz 1994). Culicoides azerbajdzhanicus and C. sejfadinei were
collected from four and five localities, respectively, all within the Eastern Anatolian Region
(Yılmaz 1994). Culicoides imicola has been confirmed in the Aegean (Aydın, Bornova,
Cakmak, Nazilli, Salihaga, and Umurlu) (Jennings et al. 1983; Dik 1993, 1996; Yagcı et al.
1999) and the Mediterranean (Adana, Anamur, Antalya, Iskenderun, Mezitli, and Tarsus)
(Burgu et al. 1992; Dik 1993; Tilki and Dik 2003) regions of Turkey. Although the species
was not detected elsewhere, care should be taken in the interpretation of this and other
species distributions, given the random sampling strategies reflected in this paper.
As the records of Culicoides presented here have been collected by various methods
(pootering, baffle traps, UV light traps, etc.), no inference of relative abundance can be
drawn. However, the following 19 species appear to be locally abundant at some sites in
Turkey: C. circumscriptus, C. dzhafarovi, C. festivipennis, C. gejgelensis, C. imicola, C.
kolymbiensis, C. longipennis, C. newsteadi, C. odiatus, C. pallidicornis, C. picturatus, C. pumilus,
C. punctatus, C. riethi, C. saevus, C. schultzei group, C. shaklawensis, C. subneglectus, and C.
vidourlensis (Table I). Little is known on the preferred breeding sites and other ecological
parameters of Culicoides species in Turkey.
Veterinary importance of Turkish Culicoides
Protozoa. Three Turkish Culicoides have been implicated in the transmission of zooparasitic
protozoa. Leucocytozoonosis is an economically important disease of poultry. In SE Asia,
C. circumscriptus and C. festivipennis (as C. odibilis) have been shown to transmit Leucocy-
tozoon caulleryi, although the latter species is not an effective vector (Morii and Kitaoka
1968). The transmission of Haemoproteus desseri and Trypanosoma bakeri to parrots, and
Hepatocystis brayi to squirrels via the bites of C. nubeculosus has been implicated experi-
mentally, but not confirmed in nature (Miltgen et al. 1976; Miltgen and Landau 1982).
Nematoda. At least two Turkish Culicoides species are proven vectors of Onchocerciasis to
domestic animals. Culicoides nubeculosus is an efficient vector of Onchocerca reticulata and O.
cervicalis to horses (Steward 1933; Moignoux 1952) and O. gutturosa to cattle (Bain 1979),
and C. oxystoma of the C. schultzei group transmits O. gibsoni to cattle (Buckley 1938).
Culicoides nubeculosus has also been shown to be capable of experimental transmission of the
Eufilarial nematodes, Eufilaria delicata and E. bartlettae, to blackbirds (Bain 1980).
Arboviruses. Culicoides imicola is clearly involved in bluetongue transmission in Turkey
(Jennings et al. 1983), but other common species including those in the C. obsoletus and C.
1964 B. Dik et al.
pulicaris species complexes may also play a criticial role in Turkey, as in other countries in
the Mediterranean Basin (De Liberato et al. 2005; Savini et al. 2005; Carpenter et al.
2006). Due to its high association with domestic livestock, members of the C. schultzei
group have also been implicated as potential bluetongue and African horse sickness virus
vectors in South Africa (Venter et al. 1996). In other regions of its range (Africa, Middle
East, and in other areas of Mediterranean Europe), Culicoides imicola is also a competent
vector of other arboviruses including African horse sickness virus, Akabane virus, Bovine
ephemeral fever, Equine encephalitis, and the Sabo and Shamonda viruses (Yonguc et al.
Figure 1. Outline map of Turkey detailing the collection sites of Culicoides listed in Table I. Collection sites are
listed numerically in alphabetical order and indicated on the map as follows: 1, Adana (36u589N, 35u229E); 2,
Aksehir (38u219N, 31u259E); 3, Anamur (36u049N, 32u499E); 4, Ankara (40u009N, 32u529E); 5, Antalya
(36u549N, 30u419E); 6, Aydın (37u499N, 27u499E); 7, Baskil (38u349N, 38u499E); 8, Bergama (39u079N,
27u109E); 9, Bespınar (37u509N, 41u359E); 10, Beysehir (37u419N, 31u439E); 11, Bornova (38u279N, 27u139E);
12, Bozkır (37u119N, 32u149E); 13, Ceylanpınar (36u509N, 40u039E); 14, Cihanbeyli (38u399N, 32u559E); 15,
Cakmak (37u479N, 29u009E); 16, Cigli (38u299N, 27u049E); 17, Cukurova (36u509N, 35u209E); 18, Cumra
(37u349N, 32u469E); 19, Denizli (37u509N, 29u089E); 20, Dinek (37u479N, 39u049E); 21, Diyarbakır (37u589N,
40u109E); 22, Elazıg (38u389N, 39u059E); 23, Erdemli (36u369N, 34u189E); 24, Eregli (37u319N, 34u019E); 25,
Gemlik (40u269N, 29u099E); 26, Gokcekısık (39u389N, 30u239E); 27, Golbası (39u489N, 32u489E); 28, Hadim
(36u599N, 32u279E); 29, Iskenderun (36u349N, 36u109E); 30, Izmir (38u279N, 27u109E); 31, Kadınhanı
(38u149N, 32u129E); 32, Kahramanmaras (37u389N, 36u489E); 33, Karaman (37u119N, 33u089E); 34, Kazan
(40u139N, 32u419E); 35, Keban (38u479N, 38u449E); 36, Konya (37u559N, 32u129E); 37, Kovancılar (38u439N,
39u519E); 38, Kusadası (37u519N, 27u159E); 39, Kuyucak (37u559N, 28u279E); 40, Maden (38u239N, 39u409E);
41, Mezitli (36u489N, 34u379E); 42, Mugla (37u139N, 28u169E); 43, Mut (36u389N, 33u269E); 44, Nazilli
(37u549N, 28u199E); 45, Pirinclik (38u129N, 49u209E); 46, Salihaga (37u549N, 29u049E); 47, Serik (36u559N,
31u069E); 48, Seydisehir (37u259N, 31u509E); 49, Simav (39u059N, 28u589E); 50, Sivrice (38u279N, 39u189E); 51,
Soke (37u459N, 27u249E); 52, Tarsus (36u559N, 34u539E); 53, Tire (38u059N, 27u449E); 54, Umurlu (37u519N,
27u579E).
Species diversity and distribution of Culicoides 1965
1982; Sellers and Pedgley 1985; Dik 1988/89), and therefore could also be involved in local
transmission of other arboviruses to livestock in Turkey. Mellor et al. (1990) isolated
African Horse Sickness (AHS) virus serotype 4 from mixed C. obsoletus and C. pulicaris
pools in Spain during the 1988 AHS outbreak and C. pulicaris s.l. (including C. pulicaris s.s.
and C. punctatus, both present in Turkey) was shown to be an effective vector of bluetongue
(BTV-1 and BTV-4) in the laboratory (Jennings and Mellor 1988). Further studies on the
species composition and vector competencies of proven/potential vector complexes
(Imicola, Obsoletus, Pulicaris, and Schultzei species complexes) in Turkey is required.
Acknowledgements
The authors are grateful to Dr John Boorman (UK) and Mr Rudy Meiswinkel (Italy) for
taxonomic advice.
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