Sanders CJ, Harrup LE, Tugwell LA, Brugman VA, England M, Carpenter S (2017). Quantification of within- and between-farm dispersal of *Culicoides* biting midges using an immunomarking technique. *J Appl Ecol* **54**(5): 1429-1439.

Sanders CJ, Shortall CR, Gubbins S, Burgin L, Gloster J, Harrington R *et al* (2011). Influence of season and meteorological parameters on flight activity of *Culicoides* biting midges. *J Appl Ecol* **48**(6): 1355-1364.

Sanogo ZL, Yaro AS, Dao A, Diallo M, Yossi O, Samake D *et al* (2020). The Effects of High-Altitude Windborne Migration on Survival, Oviposition, and Blood-Feeding of the African Malaria Mosquito, *Anopheles gambiae s.l.* (Diptera: Culicidae). *J Med Entomol*.

Santman-Berends IM, van Wuijckhuise L, Vellema P, van Rijn PA (2010). Vertical transmission of bluetongue virus serotype 8 virus in Dutch dairy herds in 2007. *Vet Microbiol* **141**(1-2): 31-35.

Sappington T, Siegfried B, Guillemaud T (2006). Coordinated Diabrotica Genetics Research: Accelerating Progress on an Urgent Insect Pest Problem. *Am Entomol* **52**: 90-97.

Sarvasova A, Goffredo M, Sopoliga I, Savini G, Kocisova A (2014). *Culicoides* midges (diptera: ceratopogonidae) as vectors of orbiviruses in Slovakia. *Vet Ital* **50**(3): 203-212.

Savini G, MacLachlan NJ, Sanchez-Vizcaino JM, Zientara S (2008). Vaccines against bluetongue in Europe. *Comp Immunol Microbiol Infect Dis* **31**(2-3): 101-120.

Schmitt T, Varga Z (2012). Extra-Mediterranean refugia: The rule and not the exception? *Front Zool* **9**(1): 22.

Schulz C, Breard E, Sailleau C, Jenckel M, Viarouge C, Vitour D *et al* (2016). Bluetongue virus serotype 27: detection and characterization of two novel variants in Corsica, France. *J Gen Virol* **97**(9): 2073-2083.

Schwartz-Cornil I, Mertens PP, Contreras V, Hemati B, Pascale F, Breard E *et al* (2008). Bluetongue virus: virology, pathogenesis and immunity. *Vet Res* **39**(5): 46.

Schwenkenbecher JM, Mordue AJ, Piertney SB (2009). Phylogenetic analysis indicates that *Culicoides dewulfi* should not be considered part of the *Culicoides obsoletus* complex. *Bull Entomol Res* **99**(4): 371-375.

Searle KR, Barber J, Stubbins F, Labuschagne K, Carpenter S, Butler A *et al* (2014). Environmental drivers of *Culicoides* phenology: how important is species-specific variation when determining disease policy? *PLoS One* **9**(11): e111876.

Sedda L, Brown HE, Purse BV, Burgin L, Gloster J, Rogers DJ (2012). A new algorithm quantifies the roles of wind and midge flight activity in the bluetongue epizootic in northwest Europe. *Proc Biol Sci* **279**(1737): 2354-2362.

Sedda L, Morley D, Brown HE (2015). Characteristics of Wind-Infective Farms of the 2006 Bluetongue Serotype 8 Epidemic in Northern Europe. *EcoHealth* **12**(3): 461-467.
Sedda L, Rogers DJ (2013). The influence of the wind in the Schmallenberg virus outbreak in Europe. *Sci Rep* **3**: 3361.

Segelbacher G, Cushman SA, Epperson BK, Fortin MJ, Francois O, Hardy OJ *et al* (2010). Applications of landscape genetics in conservation biology: concepts and challenges. *Conserv Genet* **11**(2): 375-385.

Sellers RF (1980). Weather, host and vector--their interplay in the spread of insect-borne animal virus diseases. *J Hyg (Lond)* **85**(1): 65-102.

Sellers RF, Maarouf AR (1993). Weather factors in the prediction of western equine encephalitis epidemics in Manitoba. *Epidemiol Infect* **111**(2): 373-390.

Sellers RF, Pedgley DE, Tucker MR (1982). Rift Valley fever, Egypt 1977: disease spread by windborne insect vectors? *Vet Rec* **110**(4): 73-77.

Semenza JC, Suk JE (2018). Vector-borne diseases and climate change: a European perspective. *FEMS Microbiol Lett* **365**(2): fnx244.

Semenza JC, Tran A, Espinosa L, Sudre B, Domanovic D, Paz S (2016). Climate change projections of West Nile virus infections in Europe: implications for blood safety practices. *Environ Health 15 Suppl 1*(1): 28.

Sexton JP, McIntyre PJ, Angert AL, Rice KJ (2009). Evolution and Ecology of Species Range Limits. *Annual Review of Ecology Evolution and Systematics* **40**(1): 415-436.

Sghaier S, Hammami S, Goffredo M, Hammami M, Portanti O, Lorusso A *et al* (2017). New species of the genus *Culicoides* (Diptera: Ceratopogonidae) for Tunisia, with detection of Bluetongue viruses in vectors. *Vet Ital* **53**(4): 357-366.

Shao ZY, Mao HX, Fu WJ, Ono M, Wang DS, Bonizzoni M *et al* (2004). Genetic structure of Asian populations of *Bombus ignitus* (Hymenoptera: Apidae). *J Hered* **95**(1): 46-52.

She JX, Autem M, Kotulas G, Pasteur N, Bonhomme F (1987). Multivariate-Analysis of Genetic Exchanges between Solea-Aegyptiaca and Solea-Senegalensis (Teleosts, Soleidae). *Biol J Linn Soc* **32**(4): 357-371.

Shirk AJ, Landguth EL, Cushman SA (2017). A comparison of individual-based genetic distance metrics for landscape genetics. *Mol Ecol Resour* **17**(6): 1308-1317.

Shirk AJ, Wallin DO, Cushman SA, Rice CG, Warheit KI (2010). Inferring landscape effects on gene flow: a new model selection framework. *Mol Ecol* **19**(17): 3603-3619.

Shults P, Ho A, Martin EM, McGregor BL, Vargo EL (2020). Genetic Diversity of *Culicoides stellifer* (Diptera: Ceratopogonidae) in the Southeastern United States Compared With Sequences From Ontario, Canada. *J Med Entomol* **57**(4): 1324-1327.

Sick F, Beer M, Kampen H, Wernike K (2019). *Culicoides* Biting Midges-Underestimated Vectors for Arboviruses of Public Health and Veterinary Importance. *Viruses* **11**(4).
Simo G, Njiokou F, Tume C, Lueong S, De Meeus T, Cuny G et al (2010). Population genetic structure of Central African *Trypanosoma brucei gambiense* isolates using microsatellite DNA markers. *Infect Genet Evol* **10**(1): 68-76.

Simpson SD, Harrison HB, Claereboudt MR, Planes S (2014). Long-distance dispersal via ocean currents connects Omani clownfish populations throughout entire species range. *PLoS One* **9**(9): e107610.

Siozios S, Pilgrim J, Darby AC, Baylis M, Hurst GDD (2019). The draft genome of strain cCpun from biting midges confirms insect *Cardinium* are not a monophyletic group and reveals a novel gene family expansion in a symbiont. *PeerJ* **7**: e6448.

Slade NA, Blair SM (2000). An empirical test of using counts of individuals captured as indices of population size. *J Mammal* **81**(4): 1035-1045.

Slarkin M (1985). Gene Flow in Natural Populations. *Annual Review of Ecology and Systematics* **16**(1): 393-430.

Slatkin M (1993). Isolation by Distance in Equilibrium and Non-Equilibrium Populations. *Evolution* **47**(1): 264-279.

Smouse PE, Long JC, Sokal RR (1986). Multiple-Regression and Correlation Extensions of the Mantel Test of Matrix Correspondence. *Syst Zool* **35**(4): 627-632.

Sokal RR, Jacquez GM, Oden NL, DiGiovanni D, Falsetti AB, McGee E et al (1993). Genetic relationships of European populations reflect their ethnohistorical affinities. *Am J Phys Anthropol* **91**(1): 55-70.

Solano P, Kaba D, Ravel S, Dyer NA, Sall B, Vreysen MJ et al (2010). Population genetics as a tool to select tsetse control strategies: suppression or eradication of *Glossina palpalis gambiensis* in the Niayes of Senegal. *PLoS Negl Trop Dis* **4**(5): e692.

Solano P, Ravel S, Bouyer J, Camara M, Kagbadouno MS, Dyer N et al (2009). The population structure of *Glossina palpalis gambiensis* from island and continental locations in Coastal Guinea. *PLoS Negl Trop Dis* **3**(3): e392.

Solecki AM, Skevington JH, Buddle CM, Wheeler TA (2019). Phylogeography of higher Diptera in glacial and postglacial grasslands in western North America. *BMC Ecol* **19**(1): 53.

Sommer RS, Zachos FE (2009). Fossil evidence and phylogeography of temperate species: 'glacial refugia' and post-glacial recolonization. *J Biogeogr* **36**(11): 2013-2020.

Sork VL (2016). Gene flow and natural selection shape spatial patterns of genes in tree populations: implications for evolutionary processes and applications. *Evol Appl* **9**(1): 291-310.

Spear ER, Coley PD, Kursar TA, Thrall P (2015a). Do pathogens limit the distributions of tropical trees across a rainfall gradient? *J Ecol* **103**(1): 165-174.
Spear SF, Balkenhol N, Fortin MJ, McRae BH, Scribner K (2010). Use of resistance surfaces for landscape genetic studies: considerations for parameterization and analysis. *Mol Ecol* **19**(17): 3576-3591.

Spear SF, Cushman SA, McRae BH (2015b). Resistance Surface Modeling in Landscape Genetics *Landscape Genetics*, pp 129-148.

Sperlova A, Zendulkova D (2011). Bluetongue: a review. *Vet Med (Praha)* **56**(9): 430-452.

Sprygin AV, Fiodorova OA, Babin YY, Elatkin NP, Mathieu B, England ME *et al* (2014). *Culicoides* biting midges (Diptera, Ceratopogonidae) in various climatic zones of Russia and adjacent lands. *J Vector Ecol* **39**(2): 306-315.

Stewart JR, Lister AM (2001). Cryptic northern refugia and the origins of the modern biota. *Trends Ecol Evol* **16**(11): 608-613.

Storfer A, Murphy MA, Evans JS, Goldberg CS, Robinson S, Spear SF *et al* (2007). Putting the "landscape" in landscape genetics. *Heredity* (Edinb) **98**(3): 128-142.

Storfer A, Murphy MA, Spear SF, Holderegger R, Waits LP (2010). Landscape genetics: where are we now? *Mol Ecol* **19**(17): 3496-3514.

Stouthamer R, Breeuwer JA, Hurst GD (1999). *Wolbachia pipientis*: microbial manipulator of arthropod reproduction. *Annu Rev Microbiol* **53**: 71-102.

Sudakaran S, Kost C, Kaltenpoth M (2017). Symbiont Acquisition and Replacement as a Source of Ecological Innovation. *Trends Microbiol* **25**(5): 375-390.

Suk JE, Semenza JC (2011). Future infectious disease threats to Europe. *Am J Public Health* **101**(11): 2068-2079.

Sumner T, Orton RJ, Green DM, Kao RR, Gubbins S (2017). Quantifying the roles of host movement and vector dispersal in the transmission of vector-borne diseases of livestock. *PLoS Comput Biol* **13**(4): e1005470.

Sun EC, Huang LP, Xu QY, Wang HX, Xue XM, Lu P *et al* (2016). Emergence of a Novel Bluetongue Virus Serotype, China 2014. *Transbound Emerg Dis* **63**(6): 585-589.

Tabachnick WJ (2004). *Culicoides* and the global epidemiology of bluetongue virus infection. *Vet Ital* **40**(3): 144-150.

Taberlet P, Cheddadi R (2002). Quaterny refugia and persistence of biodiversity. *Science* **297**.

Taberlet P, Fumagalli L, Wust-Saucy AG, Cosson JF (1998). Comparative phylogeography and postglacial colonization routes in Europe. *Mol Ecol* **7**(4): 453-464.

Tajima F (1989). Statistical method for testing the neutral mutation hypothesis by DNA polymorphism. *Genetics* **123**(3): 585-595.
Takezaki N, Nei M (1996). Genetic distances and reconstruction of phylogenetic trees from microsatellite DNA. *Genetics* **144**(1): 389-399.

Takken W, Verhulst N, Scholte EJ, Jacobs F, Jongema Y, van Lammeren R (2008). The phenology and population dynamics of *Culicoides* spp. in different ecosystems in The Netherlands. *Prev Vet Med* **87**(1-2): 41-54.

Talavera G, Dinca V, Vila R (2013). Factors affecting species delimitations with the GMYC model: insights from a butterfly survey. *Methods Ecol Evol* **4**(12): 1101-1110.

Talavera S, Munoz-Munoz F, Duran M, Verdun M, Soler-Membrives A, Oleaga A *et al* (2015). *Culicoides* Species Communities Associated with Wild Ruminant Ecosystems in Spain: Tracking the Way to Determine Potential Bridge Vectors for Arboviruses. *PLoS One* **10**(10): e0141667.

Talavera S, Munoz-Munoz F, Verdun M, Pages N (2017). Morphology and DNA barcoding reveal three species in one: description of *Culicoides cryptipulicaris* sp. nov. and *Culicoides quasipulicaris* sp. nov. in the subgenus *Culicoides*. *Med Vet Entomol* **31**(2): 178-191.

Talbot B, Garant D, Rioux Paquette S, Mainguy J, Pelletier F (2012). Lack of genetic structure and female-specific effect of dispersal barriers in a rabies vector, the striped skunk (*Mephitis mephitis*). *PLoS One* **7**(11): e49736.

Tatem AJ, Hay SI, Rogers DJ (2006). Global traffic and disease vector dispersal. *Proc Natl Acad Sci U S A* **103**(16): 6242-6247.

Taylor PD, Fahrig L, Henein K, Merriam G (1993). Connectivity Is a Vital Element of Landscape Structure. *Oikos* **68**(3): 571.

Temizel EM, Yesilbag K, Batten C, Senturk S, Maan NS, Mertens PPC *et al* (2009). Epizootic hemorrhagic disease in cattle, Western Turkey. *Emerg Infect Dis* **15**(2): 317-319.

Tibayrenc M (1998). Genetic epidemiology of parasitic protozoa and other infectious agents: the need for an integrated approach. *Int J Parasitol* **28**(1): 85-104.

Tibayrenc M, Ayala FJ (2002). The clonal theory of parasitic protozoa: 12 years on. *Trends Parasitol* **18**(9): 405-410.

Torina A, Caracappa S, Mellor PS, Baylis M, Purse BV (2004). Spatial distribution of bluetongue virus and its *Culicoides* vectors in Sicily. *Med Vet Entomol* **18**(2): 81-89.

Tsutsui T, Hayama Y, Yamakawa M, Shirafuji H, Yanase T (2011). Flight behavior of adult *Culicoides oxystoma* and *Culicoides maculatus* under different temperatures in the laboratory. *Parasitol Res* **108**(6): 1575-1578.

Turner MG (2005). Landscape Ecology: What Is the State of the Science? *Annu Rev Ecol Evol Syst* **36**(1): 319-344.
Tzedakis PC, Emerson BC, Hewitt GM (2013). Cryptic or mystic? Glacial tree refugia in northern Europe. *Trends Ecol Evol* **28**(12): 696-704.

Uezu A, Metzger JP, Vielliard JME (2005). Effects of structural and functional connectivity and patch size on the abundance of seven Atlantic Forest bird species. *Biol Conserv* **123**(4): 507-519.

Valade R, Kenis M, Hernandez-Lopez A, Augustin S, Mari Mena N, Magnoux E *et al* (2009). Mitochondrial and microsatellite DNA markers reveal a Balkan origin for the highly invasive horse-chestnut leaf miner *Cameraria ohridella* (Lepidoptera, Gracillariidae). *Mol Ecol* **18**(16): 3458-3470.

Van Strien MJ, Keller D, Holderegger R (2012). A new analytical approach to landscape genetic modelling: least-cost transect analysis and linear mixed models. *Mol Ecol* **21**(16): 4010-4023.

Venail R (2014). Sensibilité aux insecticides et évaluation préliminaire des méthodes de lutte antivectorielle disponibles contre les *Culicoides* (Diptera : Ceratopogonidae) paléarctiques, vecteurs de virus émergents d'intérêt en santé animale. Université De Montpellier II Sciences Et Techniques Du Languedoc.

Venail R, Balenghien T, Guis H, Tran A, Setier-Rio ML, Delécolle JC *et al* (2012). Assessing Diversity and Abundance of Vector Populations at a National Scale: Example of *Culicoides* Surveillance in France After Bluetooth Virus Emergence *Arthropods as Vectors of Emerging Diseases*. Vol. 3, pp 77-102.

Venturi G, Di Luca M, Fortuna C, Remoli ME, Riccardo F, Severini F *et al* (2017). Detection of a chikungunya outbreak in Central Italy, August to September 2017. *Euro Surveill* **22**(39): 17-00646.

Veronesi E, Henstock M, Gubbins S, Batten C, Manley R, Barber J *et al* (2013). Implicating *Culicoides* biting midges as vectors of Schmallenberg virus using semi-quantitative RT-PCR. *PLoS One* **8**(3): e57747.

Viennet E, Garros C, Gardes L, Rakotoarivony I, Allene X, Lancelot R *et al* (2013). Host preferences of Palaearctic *Culicoides* biting midges: implications for transmission of orbiviruses. *Med Vet Entomol* **27**(3): 255-266.

Vignal A, Milan D, SanCristobal M, Eggen A (2002). A review on SNP and other types of molecular markers and their use in animal genetics. *Genet Sel Evol* **34**(3): 275-305.

Votypka J, Synek P, Svobodova M (2009). Endophagy of biting midges attacking cavity-nesting birds. *Med Vet Entomol* **23**(3): 277-280.

Vreysen MJ, Balenghien T, Saleh KM, Maiga S, Koudougou Z, Cecchi G *et al* (2013). Release-recapture studies confirm dispersal of *Glossina palpalis gambiensis* between river basins in Mali. *PLoS Negl Trop Dis* **7**(4): e2022.

Wagner HH, Fortin MJ (2005). Spatial analysis of landscapes: Concepts and statistics. *Ecology* **86**(8): 1975-1987.
Wagner HH, Fortin MJ (2015). Basics of Spatial Data Analysis: Linking Landscape and Genetic Data for Landscape Genetic Studies. *Landscape Genetics: Concepts, Methods, Applications*: 77-98.

Walker AR (1977). Seasonal Fluctuations of *Culicoides* Species (Diptera-Ceratopogonidae) in Kenya. *Bull Entomol Res* **67**(2): 217-233.

Walton TE (2004). The history of bluetongue and a current global overview. *Vet Ital* **40**(3): 31-38.

Wan ZM (2014). New refinements and validation of the collection-6 MODIS land-surface temperature/emissivity product. *Remote Sens Environ* **140**: 36-45.

Wang IJ (2013). Examining the full effects of landscape heterogeneity on spatial genetic variation: a multiple matrix regression approach for quantifying geographic and ecological isolation. *Evolution* **67**(12): 3403-3411.

Waples RS, Do C (2008). ldne: a program for estimating effective population size from data on linkage disequilibrium. *Mol Ecol Resour* **8**(4): 753-756.

Waples RS, Gaggiotti O (2006). What is a population? An empirical evaluation of some genetic methods for identifying the number of gene pools and their degree of connectivity. *Mol Ecol* **15**(6): 1419-1439.

Watson TM, Saul A, Kay BH (2000). *Aedes notoscriptus* (Diptera: Culicidae) survival and dispersal estimated by mark-release-recapture in Brisbane, Queensland, Australia. *J Med Entomol* **37**(3): 380-384.

Wattier R, Engel CR, Saumitou-Laprade P, Valero M (1998). Short allele dominance as a source of heterozygote deficiency at microsatellite loci: experimental evidence at the dinucleotide locus Gv1CT in *Gracilaria gracilis* (Rhodophyta). *Mol Ecol* **7**(11): 1569-1573.

Weir BS, Cockerham CC (1984). Estimating F-Statistics for the Analysis of Population Structure. *Evolution* **38**(6): 1358-1370.

Weir BS, Goudet J (2017). A Unified Characterization of Population Structure and Relatedness. *Genetics* **206**(4): 2085-2103.

Wenk CE, Kaufmann C, Schaffner F, Mathis A (2012). Molecular characterization of Swiss Ceratopogonidae (Diptera) and evaluation of real-time PCR assays for the identification of *Culicoides* biting midges. *Vet Parasitol* **184**(2-4): 258-266.

Werner D, Groschupp S, Bauer C, Kampen H (2020). Breeding Habitat Preferences of Major *Culicoides* Species (Diptera: Ceratopogonidae) in Germany. *Int J Environ Res Public Health* **17**(14).

Werren JH, Baldo L, Clark ME (2008). *Wolbachia*: master manipulators of invertebrate biology. *Nat Rev Microbiol* **6**(10): 741-751.
Westbrook JK, Isard SA (1999). Atmospheric scales of biotic dispersal. *Agricultural and Forest Meteorology* **97**(4): 263-274.

White BP, Pilgrim EM, Boykin LM, Stein ED, Mazor RD (2014). Comparison of four species-delimitation methods applied to a DNA barcode data set of insect larvae for use in routine bioassessment. *Freshwater Science* **33**(1): 338-348.

White TA, Perkins SE, Heckel G, Searle JB (2013). Adaptive evolution during an ongoing range expansion: the invasive bank vole (*Myodes glareolus*) in Ireland. *Mol Ecol* **22**(11): 2971-2985.

Whittaker RJ, Field R (2000). Tree species richness modelling: an approach of global applicability? *Oikos* **89**(2): 399-402.

Widmer A, Lerer C (2001). Glacial refugia: sanctuaries for allelic richness, but not for gene diversity. *Trends Ecol Evol* **16**(6).

Widmer A, Schmid-Hempel P, Estoup A, Scholl A (1998). Population genetic structure and colonization history of Bombus terrestris s.l. (Hymenoptera : Apidae) from the Canary Islands and Madeira. *Heredity* **81**: 563-572.

Wiens JA, Stenseth NC, Vanhorne B, Ims RA (1993). Ecological Mechanisms and Landscape Ecology. *Oikos* **66**(3): 369-380.

Wilson AJ, Mellor PS (2009). Bluetongue in Europe: past, present and future. *Philos Trans R Soc Lond B Biol Sci* **364**(1530): 2669-2681.

Wilson GA, Rannala B (2003). Bayesian inference of recent migration rates using multilocus genotypes. *Genetics* **163**(3): 1177-1191.

Winter DJ (2012). MMOD: an R library for the calculation of population differentiation statistics. *Mol Ecol Resour* **12**(6): 1158-1160.

Wittmann EJ, Mello PS, Baylis M (2002). Effect of temperature on the transmission of orbiviruses by the biting midge, *Culicoides sonorensis*. *Med Vet Entomol* **16**(2): 147-156.

Wold S, Esbensen K, Geladi P (1987). Principal Component Analysis. *Chemometrics Intellig Lab Syst* **2**(1-3): 37-52.

Wotton KR, Gao B, Menz MHM, Morris RKA, Ball SG, Lim KS *et al* (2019). Mass Seasonal Migrations of Hoverflies Provide Extensive Pollination and Crop Protection Services. *Curr Biol* **29**(13): 2167-2173 e2165.

Wright IM (2014). Serological and genetic characterisation of putative new serotypes of bluetongue virus and epizootic haemorrhagic disease virus isolated from an alpaca MSc Thesis thesis, University Potchefstroom.

Wright S (1931). Evolution in Mendelian Populations. *Genetics* **16**(2): 97-159.
Wright S (1943). Isolation by Distance. *Genetics* **28**(2): 114-138.

Yang RC (1998). Estimating Hierarchical F-Statistics. *Evolution* **52**(4): 950-956.

Yang ZH (2015). The BPP program for species tree estimation and species delimitation. *Current Zoology* **61**(5): 854-865.

Yeruham I, Perl S, Braverman Y (2004). Seasonal allergic dermatitis in sheep associated with *Ctenocephalides* and *Culicoides* bites. *Vet Dermatol* **15**(6): 377-380.

Young SG, Carrel M, Kitchen A, Malanson GP, Tamerius J, Ali M *et al* (2017). How's the Flu Getting Through? Landscape genetics suggests both humans and birds spread H5N1 in Egypt. *Infect Genet Evol* **49**: 293-299.

Zeller KA, McGarigal K, Whiteley AR (2012). Estimating landscape resistance to movement: a review. *Landscape Ecology* **27**(6): 777-797.

Zhang DX, Hewitt GM (1997). Insect mitochondrial control region: A review of its structure, evolution and usefulness in evolutionary studies. *Biochemical Systematics and Ecology* **25**(2): 99-120.

Zhang DX, Hewitt GM (2003). Nuclear DNA analyses in genetic studies of populations: practice, problems and prospects. *Mol Ecol* **12**(3): 563-584.

Zhang J, Kapli P, Pavlidis P, Stamatakis A (2013). A general species delimitation method with applications to phylogenetic placements. *Bioinformatics* **29**(22): 2869-2876.

Zientara S, Sailleau C, Viarouge C, Hoper D, Beer M, Jenckel M *et al* (2014). Novel bluetongue virus in goats, Corsica, France, 2014. *Emerg Infect Dis* **20**(12): 2123-2125.

Zientara S, Sánchez-Vizcaíno JM. (2013). *Veterinary Microbiology, Vol. 165*.

Zimmer J-y, Haubruge É, Francis F (2014a). Synthèse bibliographique: l' écologie larvaire des *Culicoides* (Diptera: Ceratopogonidae). *Biotechnol Agron Soc Environ* **18**: 301-312.

Zimmer JY, Brostaux Y, Haubruge E, Francis F (2014b). Larval development sites of the main *Culicoides* species (Diptera: Ceratopogonidae) in northern Europe and distribution of coprophilic species larvae in Belgian pastures. *Vet Parasitol* **205**(3-4): 676-686.
# Résumé

Depuis 2006, des épizooties importantes de fièvre catarrhale ovine et de virus (FCO) de Schmallenberg (SB) sont observées dans la région ouest paléarctique, ces deux virus étant transmis par des moucherons hématophages autochtones du genre *Culicoides* (Diptera : Ceratopogonidae). Ces virus affectent les bovins, les ovins, les caprins et les ruminants sauvages. La dispersion des *Culicoides* correspond à ce que les écologues appellent dispersion stratifiée ou « stratified dispersal ». Elle résulte de la combinaison de dispersion active de proche en proche, sur de courtes distances, et de sauts sur de longues distances via le vent de manière passive ou semi-passive, permettant des sauts de dispersion de plusieurs centaines de kilomètres. *Culicoides obsoletus* est une espèce vectrice importante pour les virus de la FCO et de SB et abondante en région paléarctique. Si la dispersion des *Culicoides* par le vent au-dessus des masses d'eaux est bien établie, il est aussi important d'évaluer la dispersion au-dessus des terres et de déterminer l'influence des facteurs biotiques et abiotiques influant sur celle-ci. L'objectif de ce travail de thèse est de déterminer par l'observation des patrons de distribution des fréquences alléliques comment la dispersion a pu façonner l'arrangement spatial de la diversité génétique et comment elle est influencée par les paysages et par les caractéristiques des habitats qui peuvent favoriser ou limiter les mouvements des *Culicoides*. Pour cela, deux plans d'échantillonnage ont été utilisés : un échantillonnage populationnel à l'échelle de l'Europe (de la Scandinavie aux îles Canaries, à la Turquie), et un échantillonnage individuel à l'échelle française. L'utilisation complémentaire de marqueurs mitochondriaux et nucléaires à ces différentes échelles spatiales nous ont permis (i) de clarifier la distribution et de réviser le schéma taxonomique du Complexe Obsoletus/Scoticus, (ii) de mettre en évidence des flux de gènes important permettant d'homogénéiser la structure génétique des populations à l'échelle d'un pays et d'un continent, (iii) de quantifier l'impact d'un large éventail de facteurs environnementaux et météorologiques sur la dispersion des *Culicoides* et de mettre en évidence l'importance de la densité d'hôtes, (iv) et de proposer un scénario présentant l'histoire évolutive récente des populations européennes de *C. obsoletus*. L'hypothèse retenue est une recolonisation post-glaciaire récente depuis le refuge glaciaire des Balkans et un refuge plus au nord-est. Ces populations auraient recolonisé ensemble l'ouest de l'Europe, engendrant une admixture des populations. Nos résultats démontrent une capacité de dispersion de *C. obsoletus* à l'intérieur des terres très importante, qui doit être prise en compte pour les travaux à venir sur la compétence des vecteurs et la modélisation épidémiologique de la transmission des virus transmis.

**Mots-clés :** *Culicoides*, dispersion, génétique des populations, microsatellites, gènes mitochondriaux.

## Emerging diseases and animal health: genetic diversity and dispersal of a biting midge species vector of bluetongue and Schmallenberg viruses in the Palearctic region, *Culicoides obsoletus* (Diptera: Ceratopogonidae)

Since 2006, important outbreaks of bluetongue virus (BTV) and Schmallenberg virus (SBV) have been reported in the West Palearctic region. Both viruses are transmitted by hematophagous biting midges of the genus *Culicoides* (Diptera: Ceratopogonidae). These viruses affect wild cattle, sheep, goats and wild ruminants. The dispersion of *Culicoides* corresponds to what ecologists call stratified dispersal. It results from the combination of active dispersal from near to near, over short distances, and jumps over long distances via the wind in a passive or semi-passive way, allowing dispersal jumps of several hundred kilometers. *Culicoides obsoletus* is a recognized vector species for BTV and SBV, abundant in the Palearctic region. While the dispersal of *Culicoides* by wind over water bodies is well established, it is also important to assess the dispersion over land and determine the influence of biotic and abiotic factors influencing it. The objective of this thesis is to determine by observing the pattern of distributions of allelic frequencies how dispersion may have shaped the spatial arrangement of genetic diversity and how it is influenced by landscapes and habitat characteristics that can promote or limit the movements of *Culicoides*. Two sampling strategies were used: an Europe-wide population sampling (from Scandinavia to the Canary Islands, to Turkey), and an individual sampling at the French level. The complementary use of mitochondrial and nuclear markers at these different spatial scales has allowed us (i) to clarify the distribution and revise the taxonomic scheme of the Obsoletus/Scoticus Complex, (ii) to highlight important gene flows that homogenize the genetic structure of populations at the country and continent level, (iii) to quantify the impact of a wide range of environmental and meteorological factors on *Culicoides* dispersal and highlight the importance of host density, (iv) and to propose a scenario for the recent evolutionary history of European populations of *C. obsoletus*. The hypothesis is a recent post-glacial recolonization from the Balkan ice refuge and a refuge further northeast. These populations would have recolonized together western Europe, resulting in admixture of populations. Our results demonstrate a very important inland dispersal capability of *C. obsoletus*, which must be considered for future works on vector competence and epidemiological modelling of transmitted virus transmission.

**Key-words :** *Culicoides*, dispersal, population genetics, microsatellites, mitochondrial genes.