Schmallenberg virus in Culicoides Latreille (Diptera: Ceratopogonidae) populations in France during 2011-2012 outbreak.

Following the emergence of the Schmallenberg virus (SBV) in 2011 in Germany and its rapid spread in Europe, Culicoides (Diptera: Ceratopogonidae) collected through the French surveillance network were analysed in order to record the presence of virus genome into species diversity collected, to assess the minimum infectious rates (MIR) and the virus circulation dynamics in Culicoides populations. Two vector activity periods were selected (2011, August to October, 53 sites and 2012, June to October, 35 sites) corresponding to 704 night collections. A total of 29,285 individual midges covering at least 50 species were tested either in pools of maximum 50 females or individually (for Culicoides obsoletus/Culicoides scoticus) using real-time RT-PCR. Nine species were found SBV positive (C. obsoletus, C. scoticus, Culicoides chiopterus, Culicoides dewulfi, Culicoides imicola, Culicoides pulicaris, Culicoides newsteadi, Culicoides lupicaris and Culicoides nubeculosus) with overall MIR ranging from 0.2% to 4.2%. While the Culicoides nubeculosus laboratory strain is generally considered to have only low vector competence for viruses, interestingly, field-caught C. nubeculosus specimens were found positive twice for SBV. The first SBV-positive pool was recorded in August 2011 in north-eastern France, dating the virus circulation in France 5 months earlier than the first recorded congenital malformations and 2 months earlier than the former recorded date based on retrospective serological data. The MIR were maximum in October 2011, and in July 2012 according to dates of virus arrival in the studied areas. Moreover, our study also showed that virus circulation could be locally intense with infection rate (IR) reaching up to 16% for C. obsoletus/C. scoticus in July 2012 in one site of western France. This retrospective study demonstrates the importance of large-scale analysis to describe the spatio-temporal dynamics of virus circulation.

First record of Stegomyia albopicta (= Aedes albopictus) in Morocco: a major threat to public health in North Africa?

The Asian tiger mosquito Stegomyia albopicta (= Aedes albopictus) (Diptera: Culicidae), native to Asian forests, is a nuisance mosquito and is responsible for the transmission of arboviruses of public health importance, such as dengue, chikungunya and Zika viruses. It has colonized parts of all continents, except Antarctica, over the past 30-40 years. However, to date, the only records of S. albopicta in North Africa refer to occasional collections in 2010 and 2014 in Algeria. In early September 2015, S. albopicta larvae and adults were collected in a district of Rabat, Morocco. Morphological identification was confirmed by molecular analysis. This is the first record of this invasive mosquito in Morocco. A national surveillance programme will be implemented in 2016 to establish its geographical distribution in Morocco and to instigate control measures to prevent the establishment of new populations and the transmission of arboviruses.

Foraging range of arthropods with veterinary interest: New insights for Afrotropical Culicoides biting midges (Diptera: Ceratopogonidae) using the ring method.

The identification of blood meal source of arthropod vector species contributes to the understanding of host-vector-pathogen interactions. The aim of the current work was to identify blood meal source in Culicoides biting midge species, biological vectors of internationally important arboviruses of livestock and equids, using a new ecological approach. We examined the correlation between blood meal source identified in engorged Culicoides females collected in a suction light trap and the available vertebrate hosts along four rings (200, 500, 1000 and 2000 m) centered at the trap site and described the foraging range of the three main vector species of veterinary interest present in the study area, Culicoides imicola, Culicoides kingi and Culicoides oxystoma. The study was performed in four sites localized in the Niayes region of Senegal (West Africa) where recent outbreaks of African horse sickness occurred. Blood meal source identification was carried out by species-specific multiplex PCRs with genomic DNA extracted from the abdomen of engorged females collected during nine night collections for twenty-six collections. The four most abundant hosts present in the studied area (horse, cattle, goat and sheep) were surveyed in each ring zone. The blood meal source varied according to Culicoides species and host availability in each site. C. oxystoma and C. imicola females mainly fed on horses readily available at 200 m maximum from the trap location whereas females of C. kingi fed mainly on cattle, at variable distances from the traps (200 to 2000 m). C. oxystoma may also feed on other vertebrates. We discuss the results in relation with the transmission of Culicoides-borne arboviruses and the species dispersion capacities.

Colonization of the Mediterranean basin by the vector biting midge species Culicoides imicola: an old story.

Understanding the demographic history and genetic make-up of colonizing species is critical for inferring population sources and colonization routes. This is of main interest for designing accurate control measures in areas newly colonized by vector species of economically important pathogens. The biting midge Culicoides imicola is a major vector of orbiviruses to livestock. Historically, the distribution of this species was limited to the Afrotropical region. Entomological surveys first revealed the presence of C. imicola in the south of the Mediterranean basin by the 1970s. Following recurrent reports of massive bluetongue outbreaks since the 1990s, the presence of the species was confirmed in northern areas. In this study, we addressed the chronology and processes of C. imicola colonization in the Mediterranean basin. We characterized the genetic structure of its populations across Mediterranean and African regions using both mitochondrial and nuclear markers, and combined phylogeographical analyses with population genetics and approximate Bayesian computation. We found a west/east genetic differentiation between populations, occurring both within Africa and within the Mediterranean basin. We demonstrated that three of these groups had experienced demographic expansions in the Pleistocene, probably because of climate changes during this period. Finally, we showed that C. imicola could have colonized the Mediterranean basin in the Late Pleistocene or Early Holocene through a single event of introduction; however, we cannot exclude the hypothesis involving two routes of colonization. Thus, the recent bluetongue outbreaks are not linked to C. imicola colonization event, but rather to biological changes in the vector or the virus.

A spatiotemporal model to assess the introduction risk of African horse sickness by import of animals and vectors in France.

BACKGROUND: African horse sickness (AHS) is a major, Culicoides-borne viral disease in equines whose introduction into Europe could have dramatic consequences. The disease is considered to be endemic in sub-Saharan Africa. Recent introductions of other Culicoides-borne viruses (bluetongue and Schmallenberg) into northern Europe have highlighted the risk that AHS may arrive in Europe as well. The aim of our study was to provide a spatiotemporal quantitative risk model of AHS introduction into France. The study focused on two pathways of introduction: the arrival of an infectious host (PW-host) and the arrival of an infectious Culicoides midge via the livestock trade (PW-vector). The risk of introduction was calculated by determining the probability of an infectious animal or vector entering the country and the probability of the virus then becoming established: i.e., the virus's arrival in France resulting in at least one local equine host being infected by one local vector. This risk was assessed using data from three consecutive years (2010 to 2012) for 22 regions in France. RESULTS: The results of the model indicate that the annual risk of AHS being introduced to France is very low but that major spatiotemporal differences exist. For both introduction pathways, risk is higher from July to October and peaks in July. In general, regions with warmer climates are more at risk, as are colder regions with larger equine populations; however, regional variation in animal importation patterns (number and species) also play a major role in determining risk. Despite the low probability that AHSV is present in the EU, intra-EU trade of equines contributes most to the risk of AHSV introduction to France because it involves a large number of horse movements. CONCLUSION: It is important to address spatiotemporal differences when assessing the risk of ASH introduction and thus also when implementing efficient surveillance efforts. The methods and results of this study may help develop surveillance techniques and other risk reduction measures that will prevent the introduction of AHS or minimize AHS' potential impact once introduced, both in France and the rest of Europe.

A generic weather-driven model to predict mosquito population dynamics applied to species of Anopheles, Culex and Aedes genera of southern France.

An accurate understanding and prediction of mosquito population dynamics are needed to identify areas where there is a high risk of mosquito-borne disease spread and persistence. Simulation tools are relevant for supporting decision-makers in the surveillance of vector populations, as models of vector population dynamics provide predictions of the greatest risk periods for vector abundance, which can be particularly helpful in areas with a highly variable environment. We present a generic weather-driven model of mosquito population dynamics, which was applied to one species of each of the genera Anopheles, Culex, and Aedes, located in the same area and thus affected by similar weather conditions. The predicted population dynamics of Anopheles hyrcanus, Culex pipiens, and Aedes caspius were not similar. An. hyrcanus was abundant in late summer. Cx. pipiens was less abundant but throughout the summer. The abundance of both species showed a single large peak with few variations between years. The population dynamics of Ae. caspius showed large intra- and inter-annual variations due to pulsed egg hatching. Predictions of the model were compared to longitudinal data on host-seeking adult females. Data were previously obtained using CDC-light traps baited with carbon dioxide dry ice in 2005 at two sites (Marais du Viguerat and Tour Carbonnière) in a favourable temperate wetland of southern France (Camargue). The observed and predicted periods of maximal abundance for An. hyrcanus and Cx. pipiens tallied very well. Pearson's coefficients for these two species were over 75% for both species. The model also reproduced the major trends in the intra-annual fluctuations of Ae. caspius population dynamics, with peaks occurring in early summer and following the autumn rainfall events. Few individuals of this species were trapped so the comparison of predicted and observed dynamics was not relevant. A global sensitivity analysis of the species-specific models enabled us to identify the parameters most influencing the maximal abundance of mosquitoes. These key parameters were almost similar between species, but not with the same contributions. The emergence of adult mosquitoes was identified as a key process in the population dynamics of all of the three species considered here. Parameters associated with adult emergence therefore need to be precisely known to achieve accurate predictions. Our model is a flexible and efficient tool that predicts mosquito abundance based on local environmental factors. It is useful to and already used by a mosquito surveillance manager in France.

First overview of the Culicoides Latreille (Diptera: Ceratopogonidae) livestock associated species of Reunion Island, Indian Ocean.

This study establishes the first faunistic inventory of livestock associated Culicoides (Diptera: Ceratopogonidae) species of Reunion Island (Indian Ocean), where bluetongue and epizootic hemorrhagic disease are regularly recorded. Single night-catches were performed at 41 sites using light suction traps at altitudes ranging from 0 to 1525 m, from March to April 2005. Five species were recorded: Culicoides imicola, Culicoides bolitinos, Culicoides enderleini, Culicoides grahamii, and Culicoides kibatiensis, among which at least the first three species are known to be involved in virus transmission to ruminants and equids. This is the first record of C. bolitinos, C. kibatiensis, and C. enderleini on the island. C. imicola was the most abundant species along the sea coast. C. bolitinos was more abundant inland and on two sites on the east coast. C. kibatiensis and C. grahamii were less abundant than the other three species and limited to two foci. Spatial distribution analysis of the different species showed that C. bolitinos, C. enderleini and C. imicola were collected at low altitudes, while the other two species were found at higher altitude. A morphological identification key for adult females and males is given, as well as cytochrome oxydase subunit I sequences. Phylogenetic reconstructions showed a clear divergence between C. bolitinos from Reunion Island and mainland Africa. This monograph will help to identify the Culicoides species in the poorly known entomological fauna of the south-western Indian Ocean region.

Culicoides Latreille (Diptera: Ceratopogonidae) taxonomy: current challenges and future directions.

Culicoides Latreille biting midges (Diptera: Ceratopogonidae) cause a significant biting nuisance to humans, livestock and equines, and are the biological vectors of a range of internationally important pathogens of both veterinary and medical importance. Despite their economic significance, the delimitation and identification of species and evolutionary relationships between species within this genus remains at best problematic. To date no phylogenetic study has attempted to validate the subgeneric classification of the genus and the monophyly of many of the subgenera remains doubtful. Many informal species groupings are also known to exist but few are adequately described, further complicating accurate identification. Recent contributions to Culicoides taxonomy at the species level have revealed a high correlation between morphological and molecular analyses although molecular analyses are revealing the existence of cryptic species. This review considers the methods for studying the systematics of Culicoides using both morphological and genetic techniques, with a view to understanding the factors limiting our current understanding of Culicoides biology and hence arbovirus epidemiology. In addition, we examine the global status of Culicoides identification, highlighting areas that are poorly addressed, including the potential implementation of emerging technologies.

Host preferences of Palaearctic Culicoides biting midges: implications for transmission of orbiviruses.

Feeding success depends on host availability, host defensive reactions and host preferences. Host choice is a critical determinant of the intensity at which pathogens are transmitted. The aim of the current study was to describe host preferences of Palaearctic Culicoides species (Diptera: Ceratopogonidae) Latreille using traps baited with the five different host species of poultry, horse, cattle, sheep and goat. Collections were carried out nightly in July and August 2009 in western France with three replicates of a 5 × 5 randomized Latin square (five sites, five hosts). Moreover, an ultraviolet (UV) light/suction trap was operated during host-baited collections to correlate Culicoides biting rates and UV light/suction trap catches. A total of 660 Culicoides belonging to 12 species, but comprised mainly of Culicoides scoticus Downes and Kettle, Culicoides dewulfi Goetghebuer and Culicoides obsoletus Meigen, were collected on animal baits. Abundance was highest for the horse, which accounted for 95% of all Culicoides caught, representing 10 species. The horse, the largest bait, was the most attractive host, even when abundance data were corrected by weight, body surface or Kleiber's scaling factor. Culicoides obsoletus was the only dominant species attracted by birds. Both C. scoticus and C. dewulfi were collected mainly from the upper body of the horse. Finally, the quantification of host preferences allows for discussion of implications for the transmission of Culicoides-borne pathogens such as bluetongue virus.

A geographical information system-based multicriteria evaluation to map areas at risk for Rift Valley fever vector-borne transmission in Italy.

Rift Valley fever (RVF) is a severe mosquito-borne disease that is caused by a Phlebovirus (Bunyaviridae) and affects domestic ruminants and humans. Recently, its distribution widened, threatening Europe. The probability of the introduction and large-scale spread of Rift Valley fever virus (RVFV) in Europe is low, but localized RVF outbreaks may occur in areas where populations of ruminants and potential vectors are present. In this study, we assumed the introduction of the virus into Italy and focused on the risk of vector-borne transmission of RVFV to three main European potential hosts (cattle, sheep and goats). Five main potential mosquito vectors belonging to the Culex and Aedes genera that are present in Italy were identified in a literature review. We first modelled the geographical distribution of these five species based on expert knowledge and using land cover as a proxy of mosquito presence. The mosquito distribution maps were compared with field mosquito collections from Italy to validate the model. Next, the risk of RVFV transmission was modelled using a multicriteria evaluation (MCE) approach, integrating expert knowledge and the results of a literature review on host sensitivity and vector competence, feeding behaviour and abundance. A sensitivity analysis was performed to assess the robustness of the results with respect to expert choices. The resulting maps include (i) five maps of the vector distribution, (ii) a map of suitable areas for vector-borne transmission of RVFV and (iii) a map of the risk of RVFV vector-borne transmission to sensitive hosts given a viral introduction. Good agreement was found between the modelled presence probability and the observed presence or absence of each vector species. The resulting RVF risk map highlighted strong spatial heterogeneity and could be used to target surveillance. In conclusion, the geographical information system (GIS)-based MCE served as a valuable framework and a flexible tool for mapping the areas at risk of a pathogen that is currently absent from a region.

A review of trends in the distribution of vector-borne diseases: is international trade contributing to their spread?

It is difficult to determine the part that international trade has played in the expansion of vector-borne diseases, because of the multitude of factors that affect the transformation of habitats and the interfaces between vectors and hosts. The introduction of pathogens through trade in live animals or products of animal origin, as well as the arrival of arthropod vectors, is probably quite frequent but the establishment of an efficient transmission system that develops into a disease outbreak remains the exception. In this paper, based on well-documented examples, the authors review the ecological and epidemiological characteristics of vector-borne diseases that may have been affected in their spread and change of distribution by international trade. In addition, they provide a detailed analysis of the risks associated with specific trade routes and recent expansions of vector populations. Finally, the authors highlight the importance, as well as the challenges, of preventive surveillance and regulation. The need for improved monitoring of vector populations and a readiness to face unpredictable epidemiological events are also emphasised, since this will require rapid reaction, not least in the regulatory context.

Laboratory and field-based tests of deltamethrin insecticides against adult Culicoides biting midges.

Bluetongue virus (BTV) is an economically important arbovirus of ruminants transmitted by Culicoides biting midges. Vector control using residual spraying or application to livestock is recommended by many authorities to reduce BTV transmission; however, the impact of these measures in terms of both inflicting mortality on Culicoides and subsequently upon BTV transmission is unclear. This study consisted of a standardized World Health Organization laboratory assay to determine the susceptibility of European Culicoides species to deltamethrin and a field trial based upon allowing individuals of a laboratory strain of Culicoides nubeculosus Meigen to feed upon sheep treated with Butox 7.5 pour-on (a deltamethrin-based topical formulation). Susceptibility in the laboratory trial was higher in colony C. nubeculosus (24-h LC90 = 0.00106%), than in field populations of Culicoides obsoletus Meigen (24-h LC90 = 0.00203%) or Culicoides imicola Kieffer (24-h LC90 = 0.00773%). In the field, the pour-on formulation was tested with a total of 816 C. nubeculosus specimens fed upon on the thigh of treated sheep. The study revealed a maximum mortality rate of 49% at 4 d postapplication, and duration of lethal effect was predicted to be as short as 10 d, despite testing being carried out with a highly susceptible strain. The reasons for this low efficacy are discussed with reference both to the potential for lack of spread of the active ingredient on the host and feeding patterns of the major potential vector species on the sheep host. Practical implications for vector control strategies during BTV incursions are also detailed.

Mosquito density forecast from flooding: population dynamics model for Aedes caspius (Pallas).

Insect population dynamics depend strongly on environmental factors. For floodwater mosquitoes, meteorological conditions are crucial in the rhythm of mosquito abundances. Indeed, rainfall triggers the egg hatching after flooding breeding sites, and temperature controls the duration of the aquatic immature development up to adult emergence. According to this, we have developed a simple mechanistic and tractable model that describes the population dynamics of floodwater mosquitoes as a function only of the most accessible meteorological variables, rainfall and temperature. The model involves three parameters: development duration tdev of the immature aquatic stages, the adult emergence rate function f(t) (characterized by the emergence time scale tau and shaping the profile of adult population abundance), and the depletion rate, alpha, of adult disappearance. The developed model was subsequently applied to fit experimental field data of the dynamics of Aedes caspius (Pallas), the main pest mosquito in southern France. First, it was found that the emergence rate function of adult mosquitoes very well reproduce experimental data of the dynamics of immature development for all sampled temperatures. The estimated values of tdev and tau both exhibit Arrhenius behaviour as a function of temperature. Second, using the meteorological records of rainfall and temperature as inputs, the model correctly fit data from a two-site CO2 trapping survey conducted in 2004 and 2005. The estimated depletion rates (summation of the mortality and the emigration rates) were found to be a concave quadratic function of temperature with a maximum of 0.5 per days at about 22 degrees C.

Population sizes and dispersal pattern of tsetse flies: rolling on the river?

The West African trypanosomoses are mostly transmitted by riverine species of tsetse fly. In this study, we estimate the dispersal and population size of tsetse populations located along the Mouhoun river in Burkina Faso where tsetse habitats are experiencing increasing fragmentation caused by human encroachment. Dispersal estimated through direct (mark and recapture) and indirect (genetic isolation by distance) methods appeared consistent with one another. In these fragmented landscapes, tsetse flies displayed localized, small subpopulations with relatively short effective dispersal. We discuss how such information is crucial for designing optimal strategies for eliminating this threat. To estimate ecological parameters of wild animal populations, the genetic measures are both a cost- and time-effective alternative to mark-release-recapture. They can be applied to other vector-borne diseases of medical and/or economic importance.

Horse-, bird-, and human-seeking behavior and seasonal abundance of mosquitoes in a West Nile virus focus of southern France.

After 35 yr of disease absence, West Nile virus (family Flaviviridae, genus Flavivirus, WNV) circulation has been regularly detected in the Camargue region (southern France) since 2000. WNV was isolated from Culex modestus Ficalbi, which was considered the main vector in southern France after horse outbreaks in the 1960s. Recent WNV transmissions outside of the Cx. modestus distribution suggested the existence of other vectors. To study potential WNV vectors, horse- and bird-baited traps and human landing collections of mosquitoes were carried out weekly from May to October 2004 at two Camargue sites: one site in a wet area and the other site in a dry area, both chosen for their past history of WNV transmission. At the wet site, the most abundant species in bird-baited traps were Culex pipiens L. and Cx. modestus; both species also were found in lower proportions on horses and humans. The most abundant species in horse-baited traps and human landing collections were Aedes caspius (Pallas), Aedes vexans (Meigen), and Anopheles hyrcanus (Pallas) sensu lato; some of these species were occasionally collected with avian blood at the end of the summer. Anopheles maculipennis Meigen sensu lato was an abundant horse feeder, but it was rarely collected landing on human bait and never contained avian blood. At the dry site, Cx. pipiens was the most abundant species in bird- and horse-baited traps. The seasonal and circadian dynamics of these species are analyzed, and their potential in WNV transmission in Camargue discussed.

Modelling and determination of the transmission contact rate for contagious bovine pleuropneumonia.

Contagious bovine pleuropneumonia (CBPP) is a cattle respiratory disease that represents one of the major threats to cattle health and production in sub-Saharan Africa. The transmission contact rate of CBPP plays a key role in the spreading dynamics of the disease. We have developed an approach based on the combination of a SEIR model describing the spread of CBPP with the dynamic of seroconversion to determine the transmission contact rate for CBPP. This method has been subsequently applied to serological diagnostic data obtained from an experimental vaccine trial. As a result, we find that the transmission contact rates for subclinical, clinical and chronic infective states are respectively, 0.084/N, 0.45 and 0.14/N per animal per day, where N is the herd population size, and the basic reproductive number corresponding to this trial (N=28) is R0=27.