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We investigate the restriction of animal movements as a method to control the spread of bluetongue, an infectious disease of livestock that is becoming increasingly prevalent due to the onset of climate change. We derive control policies for the UK that minimise the number of infected farms during an outbreak using Bayesian optimisation and a simulation-based model of BT. Two cases are presented: first, where the region of introduction is randomly selected from England and Wales to find a generalised strategy. This "national" model is shown to be just as effective at subduing the spread of bluetongue as the current strategy of the UK government. Our proposed controls are simpler to implement, affect fewer farms in the process and, in so doing, minimise the potential economic implications. Second, we consider policies that are tailored to the specific region in which the first infection was detected. Seven different regions in the UK were explored and improvements in efficiency from the use of specialised policies presented. As a consequence of the increasing temperatures associated with climate change, efficient control measures for vector-borne diseases such as this are expected to become increasingly important. Our work demonstrates the potential value of using Bayesian optimisation in developing cost-effective disease management strategies.

In 2006, bluetongue (BT), a disease of ruminants, was introduced into northern Europe for the first time and more than two thousand farms across five countries were affected. In 2007, BT affected more than 35,000 farms in France and Germany alone. By contrast, the UK outbreak beginning in 2007 was relatively small, with only 135 farms in southeast England affected. We use a model to investigate the effects of three factors on the scale of BT outbreaks in the UK: (1) place of introduction; (2) temperature; and (3) animal movement restrictions. Our results suggest that the UK outbreak could have been much larger had the infection been introduced into the west of England either directly or as a result of the movement of infected animals from southeast England before the first case was detected. The fact that air temperatures in the UK in 2007 were marginally lower than average probably contributed to the UK outbreak being relatively small. Finally, our results indicate that BT movement restrictions are effective at controlling the spread of infection. However, foot-and-mouth disease restrictions in place before the detection and control of BT in 2007 almost certainly helped to limit BT spread prior to its detection.

There is a solid theoretical basis for expecting climate change to have a considerable effect on the infectious diseases of humans, animals and plants. Vector-borne diseases are the most likely to be affected. It is, however, rare to observe such impacts, as diseases are also influenced by many other drivers, some of which may have stronger effects over shorter time scales than climate change. Nevertheless, there is evidence that our warming climate has already influenced some animal diseases, of which bluetongue is considered a prime example. Bluetongue emerged dramatically in southern Europe after 1998 and in northern Europe from 2006. While the speed and scale of this emergence is a challenge to explain, there is evidence, principally from the development of climate-driven models, that recent climate change has played a significant role. Climate-driven models point to an increase in the risk of bluetongue transmission in Europe in recent decades, caused by an increased suitability of parts of southern Europe for the Afro-tropical biting midge, Culicoides imicola, as well as an increase in the vectorial capacity of indigenous Culicoides vectors in northern Europe. Farm-to-farm transmission models of bluetongue in England and Wales under predicted climatic conditions further suggest that, under high-emission scenarios, the scale of future outbreaks could far exceed those experienced to date. The role of climate change in the developing threat of animal disease is, therefore, likely to be economically and socially costly, unless lower emission targets can be set and followed.

Background: Farm to farm movement of Culicoides midges is believed to play a critical role in the spread of bluetongue (BT), Schmallenberg and other midge-borne diseases. To help understand and predict the spread of diseases carried by midges, there is a need to determine their dispersal patterns, and to identify factors contributing to the direction taken and distance travelled., Methods: The dispersal of Obsoletus Group members was studied on 19 farms around Bala, north Wales. Field-collected Culicoides were trapped in a black-light (OVI) trap and self-marked in the collecting vessel, using micronized fluorescent dust. Culicoides were released at a central farm and OVI traps set on 18 surrounding farms, at distances of 1 to 4 km. The study was repeated using six colours of fluorescent dust over an 18 day period., Results: An estimated 61,062 (95% CI = 56,298-65,830) marked Culicoides were released during the study and 12 (0.02%) Culicoides were recaptured. Of the females recaptured, six were C. obsoletus/scoticus, two C. dewulfi, two C. pulicaris and one C. festivipennis. The male was C. obsoletus. Recaptures occurred 1-2.5 km from the release site, with greatest numbers at 2.5 km. Most recaptures were 2 nights post-release; none were more than 3 nights post-release. Two females were recovered at 1.5 km on the night of release and one male at 1 km two nights post-release. The mean distance travelled (MDT) for males was 1 km, females was 2.21 km, and all recaptured Culicoides was 2.15 km. Recaptures were made both downwind and upwind of the prevailing wind direction during the trapping periods, highlighting possible passive and active dispersal of Culicoides between farms., Conclusions: This is the first study to demonstrate farm to farm movement of the main Palaearctic BT vector species, the Obsoletus Group. Such movement has disease control implications in terms of the vectoral movement of disease between farms. The results suggest that Culicoides control measures applied at an infected farm (trapping or killing Culicoides) will reduce risk of spread to neighbouring farms by lessening the number of Culicoides dispersing from that farm, as well as reducing transmission at the source farm itself.

Climate change is widely expected to cause the emergence and spread of vector-borne diseases, and predictive models are needed so that we can be prepared. We developed a climate-sensitive, predictive, model that describes the risk of bluetongue, an arboviral disease of ruminants, which has emerged dramatically in Europe. Developing the predictive bluetongue model led to the identification of numerous gaps in both the understanding and the availability of data. These mostly pertain to the vectors and their interaction with hosts. Closing these gaps will allow better models, with more precise predictions, to be produced. These research gaps apply to many other arboviral diseases as well. As a consequence, there needs to be an increase in research on the vectors that transmit arboviral diseases. Priorities are the training of a new generation of taxonomists, studies on the field biology of potential vectors, and increased coordination of vector surveillance and recording between countries facing similar threats.

Bluetongue (BT) can cause severe livestock losses and large direct and indirect costs for farmers. To propose targeted control strategies as alternative to massive vaccination, there is a need to better understand how BT virus spread in space and time according to local characteristics of host and vector populations. Our objective was to assess, using a modelling approach, how spatiotemporal heterogeneities in abundance and distribution of hosts and vectors impact the occurrence and amplitude of local and regional BT epidemics. We built a reaction-diffusion model accounting for the seasonality in vector abundance and the active dispersal of vectors. Because of the scale chosen, and movement restrictions imposed during epidemics, host movements and wind-induced passive vector movements were neglected. Four levels of complexity were addressed using a theoretical approach, from a homogeneous to a heterogeneous environment in abundance and distribution of hosts and vectors. These scenarios were illustrated using data on abundance and distribution of hosts and vectors in a real geographical area. We have shown that local epidemics can occur earlier and be larger in scale far from the primary case rather than close to it. Moreover, spatial heterogeneities in hosts and vectors delay the epidemic peak and decrease the infection prevalence. The results obtained on a real area confirmed those obtained on a theoretical domain. Although developed to represent BTV spatiotemporal spread, our model can be used to study other vector-borne diseases of animals with a local to regional spread by vector diffusion.

Mathematical formulations for the basic reproduction ratio (R(0)) exist for several vector-borne diseases. Generally, these are based on models of one-host, one-vector systems or two-host, one-vector systems. For many vector borne diseases, however, two or more vector species often co-occur and, therefore, there is a need for more complex formulations. Here we derive a two-host, two-vector formulation for the R(0) of bluetongue, a vector-borne infection of ruminants that can have serious economic consequences; since 1998 for example, it has led to the deaths of well over 1 million sheep in Europe alone. We illustrate our results by considering the situation in South Africa, where there are two major hosts (sheep, cattle) and two vector species with differing ecologies and competencies as vectors, for which good data exist. We investigate the effects on R(0) of differences in vector abundance, vector competence and vector host preference between vector species. Our results indicate that R(0) can be underestimated if we assume that there is only one vector transmitting the infection (when there are in fact two or more) and/or vector host preferences are overlooked (unless the preferred host is less beneficial or more abundant). The two-host, one-vector formula provides a good approximation when the level of cross-infection between vector species is very small. As this approaches the level of intraspecies infection, a combination of the two-host, one-vector R(0) for each vector species becomes a better estimate. Otherwise, particularly when the level of cross-infection is high, the two-host, two-vector formula is required for accurate estimation of R(0). Our results are equally relevant to Europe, where at least two vector species, which co-occur in parts of the south, have been implicated in the recent epizootic of bluetongue.

Vector-borne diseases are among those most sensitive to climate because the ecology of vectors and the development rate of pathogens within them are highly dependent on environmental conditions. Bluetongue (BT), a recently emerged arboviral disease of ruminants in Europe, is often cited as an illustration of climate's impact on disease emergence, although no study has yet tested this association. Here, we develop a framework to quantitatively evaluate the effects of climate on BT's emergence in Europe by integrating high-resolution climate observations and model simulations within a mechanistic model of BT transmission risk. We demonstrate that a climate-driven model explains, in both space and time, many aspects of BT's recent emergence and spread, including the 2006 BT outbreak in northwest Europe which occurred in the year of highest projected risk since at least 1960. Furthermore, the model provides mechanistic insight into BT's emergence, suggesting that the drivers of emergence across Europe differ between the South and the North. Driven by simulated future climate from an ensemble of 11 regional climate models, the model projects increase in the future risk of BT emergence across most of Europe with uncertainty in rate but not in trend. The framework described here is adaptable and applicable to other diseases, where the link between climate and disease transmission risk can be quantified, permitting the evaluation of scale and uncertainty in climate change's impact on the future of such diseases.

Bluetongue is a notifiable disease of ruminants which, in 2007, occurred for the first time in England. We present the first model for bluetongue that explicitly incorporates farm to farm movements of the two main hosts, as well as vector dispersal. The model also includes a seasonal vector to host ratio and dynamic restriction zones that evolve as infection is detected. Batch movements of sheep were included by modelling degree of mixing at markets. We investigate the transmission of bluetongue virus between farms in eastern England (the focus of the outbreak). Results indicate that most parameters affecting outbreak size relate to vectors and that the infection generally cannot be maintained without between-herd vector transmission. Movement restrictions are effective at reducing outbreak size, and a targeted approach would be as effective as a total movement ban. The model framework is flexible and can be adapted to other vector-borne diseases of livestock.

On four nights in June 2008, light traps were operated for Culicoides biting midges, the vector species for bluetongue virus (BTV), at five sites in Chester Zoo in north-west England. Over 35,000 Culicoides midges, of 25 species, were captured, including high densities inside animal enclosures. Over 94 per cent of all the Culicoides trapped were females of the Obsoletus group, which is implicated as the vector of BTV serotype 8 in northern Europe. The mean catch of this group per trap per night was over 1500, suggesting a potential risk of BTV transmission if the virus is introduced to Chester Zoo in the animals or midges in the summer.

The housing of animals at night was investigated as a possible means of protecting them from attack by Culicoides biting midges (Diptera: Ceratopogonidae), the vectors of bluetongue. Light-trap catches of Culicoides were compared inside and outside animal housing, in the presence and absence of cattle. A three-replicate, 4 x 4 Latin square design was used at four farms in Bala, north Wales, over 12 nights in May and June 2007, and the experiment repeated in October. In the two studies, respectively, >70 000 and >4500 Culicoides were trapped, of which 93% and 86%, respectively, were of the Culicoides obsoletus group. Across the four farms, in May and June, the presence of cattle increased catches of C. obsoletus by 2.3 times, and outside traps caught 6.5 times more insects than inside traps. Similar patterns were apparent in October, but the difference between inside and outside catches was reduced. Catches were strongly correlated with minimum temperature and maximum wind speed and these two variables explained a large amount of night-to-night variation in catch. Outside catches were reduced, to a greater extent than inside catches, by colder minimum temperatures and higher maximum wind speeds. These conditions occur more frequently in October than in May and June, thereby suppressing outside catches more than inside catches, and reducing the apparent degree of exophily of C. obsoletus in autumn. The results suggest that the risk of animals receiving bites from C. obsoletus is reduced by housing at both times of year and the benefit would be greatest on warm, still nights when outside catches are at their greatest.

Bluetongue virus (BTV) exists around the world in a broad band covering much of the Americas, Africa, southern Asia and northern Australia. Historically, it also occasionally occurred in the southern fringes of Europe. It is considered to be one of the most important diseases of domestic livestock. Recently BTV has extended its range northwards into areas of Europe never before affected and has persisted in many of these locations causing the greatest epizootic of bluetongue (BT), the disease caused by BTV, on record. Indeed, the most recent outbreaks of BT in Europe are further north than this virus has ever previously occurred anywhere in the world. The reasons for this dramatic change in BT epidemiology are complex but are linked to recent extensions in the distribution of its major vector, Culicoides imicola, to the involvement of novel Culicoides vector(s) and to on-going climate-change. This paper investigates these recent outbreaks in the European theatre, up to the beginning of 2006, highlights prospects for the future and sets the scene for the following papers in this special issue.

Two experiments were undertaken to estimate the transmission rates of bluetongue virus (BTV) serotype 1 between a biting midge vector, Culicoides sonorensis (Wirth & Jones) (Ceratopogonidae), and a natural host, sheep. In an experiment to measure the transmission rate from vector to host (V-->H), six batches of one, five and 20 intrathoracically infected midges were fed on a total of 18 bluetongue (BT)-naïve sheep. The sheep were then monitored for 21 days for clinical signs of BT, viraemia and antibody response. All sheep fed on by five or 20 midges and five of six sheep fed on by just one midge showed signs of BT, were viraemic and developed antibody. The sixth sheep fed on by a single infected midge did not show signs of BT or have detectable viraemia; it did, however, develop a weak antibody response. A bite from a single infected midge is therefore able to transmit BTV to naïve sheep with 80-100% efficiency. Sheep fed upon by larger numbers of infected midges took less time to reach maximum viraemia and developed stronger antibody responses. Sheep exposed to greater amounts of BTV in feeding midges developed a higher level of viraemia and stronger antibody responses. In a second experiment to measure the transmission rate from host to vector (H-->V), batches of up to 500 uninfected female C. sonorensis fed every 1-2 days on two experimentally infected sheep during the course of infection. Of 3929 engorged midges that were individually titrated after surviving the extrinsic incubation period, only 23 (0.6%) were infected with BTV. Viraemia in the sheep extended for up to 19 days post-inoculation. No infected midges, however, were detected from 14 days post-infection.

Since 1998 bluetongue virus (BTV), which causes bluetongue, a non-contagious, insect-borne infectious disease of ruminants, has expanded northwards in Europe in an unprecedented series of incursions, suggesting that there is a risk to the large and valuable British livestock industry. The basic reproduction number, R(0), provides a powerful tool with which to assess the level of risk posed by a disease. In this paper, we compute R(0) for BTV in a population comprising two host species, cattle and sheep. Estimates for each parameter which influences R(0) were obtained from the published literature, using those applicable to the UK situation wherever possible. Moreover, explicit temperature dependence was included for those parameters for which it had been quantified. Uncertainty and sensitivity analyses based on Latin hypercube sampling and partial rank correlation coefficients identified temperature, the probability of transmission from host to vector and the vector to host ratio as being most important in determining the magnitude of R(0). The importance of temperature reflects the fact that it influences many processes involved in the transmission of BTV and, in particular, the biting rate, the extrinsic incubation period and the vector mortality rate.

Surveillance of Culicoides (Diptera: Ceratopogonidae) biting midges was carried out between 2001 and 2003, at 119 sites within a 50 x 50-km grid distributed across Bulgaria, using light trap collections around the time of peak adult midge abundance. Sentinel and ad hoc serum surveillance of hosts susceptible to bluetongue infection was carried out at around 300 sites between 1999 and 2003. Following the initial incursion of bluetongue virus 9 (BTV-9) into Bourgas province in 1999, affecting 85 villages along the southern border, a further 76 villages were affected along the western border in 2001, with outbreaks extending as far north as 43.6 degrees N. The BTV-9 strain in circulation was found to have a low pathogenicity for Bulgarian sheep populations, with less than 2% of susceptible individuals becoming sick and seroconversions detected up to 30 km from recorded outbreaks in the south. The major Old World vector Culicoides imicola Kieffer was not detected among over 70,000 Culicoides identified in summer collections, suggesting that BTV-9 transmission in Bulgaria was primarily carried out by indigenous European vectors. The most likely candidates, the Palaearctic species complexes - the Culicoides obsoletus Meigen and C. pulicaris L. complexes - were widespread and abundant across the whole country. The C. obsoletus complex represented 75% of all individuals trapped in summer and occurred in high catch sizes (up to 15,000 individuals per night) but was not found across all outbreak sites, indicating that both Palearctic complexes probably played a role in transmission. Within the C. pulicaris complex, only C. pulicaris s.s., C. punctatus Meigen and C. newsteadi Austen were sufficiently abundant and prevalent to have been widely involved in transmission, whilst within the C. obsoletus complex most trapped males were C. obsoletus s.s. Adult vectors were found to be largely absent from sites in west Bulgaria for a period of at least 3 months over winter, which, taken along with the spatiotemporal pattern of outbreaks in the region between years, indicates the virus may be overwintering here by an alternative mechanism - either by covert persistence in the vertebrate host or possibly by persistence in larval stages of the vector.

Bluetongue, a devastating disease of ruminants, has historically made only brief, sporadic incursions into the fringes of Europe. However, since 1998, six strains of bluetongue virus have spread across 12 countries and 800 km further north in Europe than has previously been reported. We suggest that this spread has been driven by recent changes in European climate that have allowed increased virus persistence during winter, the northward expansion of Culicoides imicola, the main bluetongue virus vector, and, beyond this vector's range, transmission by indigenous European Culicoides species - thereby expanding the risk of transmission over larger geographical regions. Understanding this sequence of events may help us predict the emergence of other vector-borne pathogens.

Determining the temporal relationship between climate and epidemics of Culicoides-borne viral disease may allow control and surveillance measures to be implemented earlier and more efficiently. In Israel, outbreaks of bluetongue (BT) have occurred almost annually since at least 1950, with severe episodes occurring periodically. In this paper, the authors model a twenty-year time-series of BT outbreaks in relation to climate. Satellite-derived correlates of low temperatures and high moisture levels increased the number of outbreaks per year. This is the first study to find a temporal relationship between the risk of Culicoides-borne disease and satellite-derived climate variables. Climatic conditions in the year preceding a BT episode, between October and December, coincident with the seasonal peak of vector abundance and outbreak numbers, appeared to be more importantthan spring or early summer conditions in the same year as the episode. Since Israel is an arid country, higher-than-average moisture levels during this period may increase the availability of breeding sites and refuges for adult Culicoides imicola vectors, while cooler-than-average temperatures will increase fecundity, offspring size and survival through adulthood in winter, which, in turn, increases the size of the initial vector population the following year. The proportion of variance in the annual BT outbreak time-series resulting from climate factors was relatively low, at around 20%. This was possibly due to temporal variation in other factors, such as viral incursions from surrounding countries and levels of herd immunity. Alternatively, since most BT virus (BTV) circulation in this region occurs silently, in resistant breeds of local sheep, the level of transmission is poorly correlated with outbreak notification so that strong relationships between BTV circulation and climate, if they exist, are obscured.

During the recent Mediterranean epizootic of bluetongue, an extensive programme of serological and vector (Culicoides biting midges (Diptera: Ceratopogonidae)) surveillance was carried out across Sicily. This paper presents the analysis of 911 light trap catches collected at the times of peak Culicoides abundance (summer to autumn 2000-2002) in 269 sites, in order to produce detailed maps of the spatial distribution of the main European vector, Culicoides imicola Kieffer and that of potential novel vectors. Whereas C. imicola was found at only 12% of sites, potential novel vectors, Culicoides obsoletus group Meigen, Culicoides pulicaris Linnaeus and Culicoides newsteadi Austen were present at over 50% of sites. However, the spatial distribution of C. imicola showed the closest correspondence to that of the 2000 and 2001 bluetongue (BT) outbreaks and its presence and abundance were significant predictors of the probability of an outbreak, suggesting that it was the main vector during these years. Although C. imicola may have played a role in transmission in several sites near Paternó, it was absent from the majority of sites at which outbreaks occurred in 2002 and from all sites in the province of Messina. All three potential novel vectors were widespread across sites at which outbreaks occurred during 2002. Of these, C. newsteadi was an unlikely candidate, as it was significantly less prevalent in outbreak vs. non-outbreak sites in Messina. It is hypothesized that the yearly distribution and intensity of outbreaks is directly attributable to the distribution and abundance of the vectors involved in transmission during each year. When C. imicola operated as the main vector in 2000 and 2001, outbreaks were few in number and were restricted to coastal regions due to low abundance and prevalence of this species. In 2002, it is hypothesized that BTV transmission was handed over to more prevalent and abundant novel vector species, leading to numerous and widespread outbreaks and probably to overwintering of the virus between 2001 and 2002. Based on catch ranges in outbreak vs. non-outbreak sites, it is tentatively suggested that nightly catches of 400 or more C. obsoletus and 150 or more C. pulicaris allow BTV transmission at a site, and provide a strategy for a fuller examination of the relationship between BTV transmission and the abundance and distribution of different vector species.

Bluetongue is an infectious, non-contagious arboviral disease thought to infect all known ruminant species. Since 1998, an unprecedented epizootic of the disease has occurred in the Mediterranean region, resulting in the deaths of over 800,000 sheep to date. Bluetongue virus (BTV) is transmitted by biting midges of which one species, Culicoides imicola, is the major vector in the old world. C. imicola was trapped for 2 years at 87 sites across Portugal and models were developed for predicting the presence and abundance of the midge at these sites. Discriminant analysis was used to identify the best models from 40 temporally Fourier-processed 1 km spatial resolution remotely-sensed variables. The best models correctly predicted presence and absence at 83 of the 87 sites, and abundance at 76 sites. The models were then used to predict C. imicola presence and abundance elsewhere across Europe and north Africa. C. imicola was predicted to be present and in high abundance at the majority of areas affected in the recent bluetongue epizootic, including the Balearics, Sardinia, Corsica, Sicily, areas of mainland Italy, large areas of Greece, western Turkey and northern Algeria and Tunisia.

Surveillance of Culicoides (Diptera: Ceratopogonidae) biting midge vectors was carried out at 87 sites within a 50 x 50 km grid distributed across Portugal, using light trap collections at the time of peak midge abundance. Culicoides imicola (Kieffer) made up 66% of the 55 937 Culicoides in these summer collections. It was highly abundant in the central eastern portion of Portugal, between 37 degrees 5' N and 41 degrees 5' N, and in a band across to the Lisbon peninsula (at around 38 degrees 5' N). Of all the complexes, its distribution was most consistent with that of previous outbreaks of Culicoides-borne disease, suggesting that it may remain the major vector in Portugal. Its distribution was also broadly consistent with that predicted by a recent climate-driven model validating the use of remote sensing datasets for modelling of Culicoides distribution. Adult C. imicola were found to have overwintered at 12 of 20 sites re-surveyed in winter but it did so in very low numbers. Culicoides obsoletus (Meigen) and Culicoides pulicaris (Linnaeus) complex midges were widespread despite their low summer abundance. The observed coincidence of high abundances of C. imicola and high abundances of C. pulicaris in summer lead us to suggest that C. imicola could bring African horse sickness virus or bluetongue virus into contact with C. pulicaris and the latter complex, together with C. obsoletus, could then transmit these viruses across much wider areas of Europe. The fact that adult C. pulicaris are present in high abundances in winter may provide a mechanism by which these viruses can overwinter in these areas.

Bluetongue is an infectious disease of ruminants caused by a virus transmitted by biting midges, one species of which, Culicoides imicola, is the major vector in the Old World. Following an epizootic of African horse sickness,a related disease, in Iberia and Morocco between 1987 and 1991, C imicola was trapped for two years at 44 sites in the affected region and models were developed for predicting the abundance of C imicola at these sites. Discriminant analysis was applied to identify the best model of three levels of abundance from 40 Fourier-processed remotely sensed variables and a digital elevation model. The best model correctly predicted the abundance level at 41 of the 44 sites. The single most important variable was the phase of the annual cycle of the normalised difference vegetation index. The model was used to predict the abundances of C imicola elsewhere around the Mediterranean and predicted high levels of abundance in many areas recently affected by bluetongue, including the Balearics, Sardinia, Sicily, eastern Greece, western Turkey, Tunisia and northern Algeria. The model suggests that eastern Spain, the island of Ibiza, the provinces of Lazio and Puglia in Italy, the Peloponnese and parts of northern Algeria and Libya may be at risk of bluetongue in 2001.

Changes in the distribution and abundance of insects are likely to be amongst the most important and immediate effects of climate change. We review here the risk that climate change poses to the UK's livestock industry via effects on Culicoides biting midges, the vectors of several arboviruses, including those that cause bluetongue (BT) and African horse sickness (AHS). The major old-world vector of BT and AHS viruses, C. imicola, occurs in southern Europe and will spread further north as global temperatures increase. It is unlikely, however, that in the foreseeable future it will reach and become established in the UK. As the distribution of C. imicola moves north, however, it may bring BT and AHS viruses into the range of other Culicoides species that are known to be competent vectors and which occur much further north. Once infected via this 'baton effect', these species may be able to spread the viruses over much of Europe, including the UK. Climate change may increase their vector competence further and will also increase the likelihood of viruses surviving from one year to the next. An additional risk is that the predicted increase in the frequency of short periods of hot temperatures may lead to the creation of novel vector species, by removing the barriers that in colder conditions make them refractory to viral infection., (Copyright 2000 Harcourt Publishers Ltd.)

The biting midge Culicoides imicola Kieffer (Diptera: Ceratopogonidae) is the most important Old World vector of African horse sickness (AHS) and bluetongue (BT). Recent increases of BT incidence in the Mediterranean basin are attributed to its increased abundance and distribution. The phylogenetic status and genetic structure of C. imicola in this region are unknown, despite the importance of these aspects for BT epidemiology in the North American BT vector. In this study, analyses of partial mitochondrial cytochrome oxidase subunit I gene (COI) sequences were used to infer phylogenetic relationships among 50 C. imicola from Portugal, Rhodes, Israel, and South Africa and four other species of the Imicola Complex from southern Africa, and to estimate levels of matrilineal subdivision in C. imicola between Portugal and Israel. Eleven haplotypes were detected in C. imicola, and these formed one well-supported clade in maximum likelihood and Bayesian trees implying that the C. imicola samples comprise one phylogenetic species. Molecular variance was distributed mainly between Portugal and Israel, with no haplotypes shared between these countries, suggesting that female-mediated gene flow at this scale has been either limited or non-existent. Our results provide phylogenetic evidence that C. imicola in the study areas are potentially competent AHS and BT vectors. The geographical structure of the C. imicola COI haplotypes was concordant with that of BT virus serotypes in recent BT outbreaks in the Mediterranean basin, suggesting that population subdivision in its vector can impose spatial constraints on BT virus transmission. [ABSTRACT FROM AUTHOR]