Modelling the roles of climate, landscape and biotic factors in the distribution and maximum trap catch of Culicoides vectors across Europe. S. Withenshaw1, K. Searle1, A. Butler2, A. Allepuz3, J. Barber4, S. Carpenter4, E. Denison4, K. Labuschagne5, P. Mellor4, M. Miranda6, S. Napp3, N. Pages3, C. Sanders4, F. Stubbins7, B. Purse1 1. Centre for Ecology and Hydrology 2. Biomathematics and Statistics Scotland 3. Centre de Recerca en Sanitat Animal 4. The Pirbright Institute 5. PVVD, ARC-Onderstepoort Veterinary Institute 6. Laboratory of Zoology, University of the Balaeric Island 7. School of Agricultural, Forest and Environmental Sciences, Clemson University Table 1: Covariates investigated in models of Culicoides female maximum annual trap abundance. All covariates were extracted for the location of each trap site for model fitting, and the centroids of 1km grid squares across Europe for predictions. Background Culicoides biting midges (Diptera: Ceratopogonidae) transmit bluetongue virus (BTV), which has caused significant disruption to ruminant production systems in northern Europe over the past decade. The presence and abundance of female Culicoides vectors are important determinants of the risk of BTV infection within a region, therefore understanding the drivers of these spatial patterns will aid the prediction and Figure 1: Culicoides spp. mitigation of future BTV incursions. Here we present preliminary results of a novel Bayesian modelling framework for determining the climate, landscape and biotic factors driving the maximum annual trap catch of three Culicoides species groups sampled across the UK and Spain. The resulting models were used to make predictions of this variable across west and central Europe for past years. Methods 1. Data • Midges sampled weekly at sites in UK and Spain (2005-2010). • 404 site-year combinations (Fig 2). • Light-suction traps. • Species identified morphologically and counted (C. imicola, C. obsoletus complex, C. pulicaris complex). • Maximum size of weekly female catch per species per year at each site was determined. Covariate Culicoides species C. imicola Precipitation Summed precipitation in mm (ppt) per year Land cover Covariates included in the 20 best models 0.58 Temperature (20) Sheep density (16) Summed NDVI (15) Elevation (12) Cattle density (8) Roe deer density (5) Red deer density (4) Number of dry days (dd) per year Coefficient of variation in summed precipitation (cvppt) per year Categorical variable indicating the type of land cover present MODIS 8-day 1km Land Surface Temp ENSEMBLES daily 0.25° precipitation MODIS annual 500m land cover (Crop, Crop/Natural Vegetation Mosaic, Deciduous Broadleaf/Mixed Forest, Evergreen Needleleaf Forest, Grassland, Open Shrubland, Savanna, Woody Savanna, Urban) NDVI Summed values for the Normalised Difference Vegetation Index per year Elevation Metres above sea level MODIS 8-day 250m Vegetation Indices WORLDCLIM 1km elevation Deer density Modelled density per km2 of roe deer, red deer and total [1] [2] deer (roe + red) Livestock density Modelled density per km2 of cattle, sheep, goats and total (cattle+sheep+goats) [3] Results and Discussion • The covariates included in models of maximum trap catch differed between the Culicoides spp. groups considered here (Table 2). • The model predicts the distribution of C. imicola to be restricted to southern Europe (Fig 3a), in line with previous prediction maps (Fig 3d) and sampled presence (Fig 3e)[4,5]. However, there is under-prediction of presence in Spain and over-prediction in Italy[6] (Fig 3f) and Bulgaria[7]. • The model predicts higher abundance of C. obsoletus complex and C. pulicaris complex in northern Europe compared to the south. However, the southern range limits for both of these groups are known to stretch down to north Africa[5] (Fig 3e). The prediction of zero abundance in the south Mediterranean region is therefore likely to be an artefact of the sampling range (Fig 2). • The predicted distribution of the C. pulicaris complex in Portugal looks more similar to the known distribution of a closely-related species, C. punctatus, which penetrates less far south[8]. This may indicate complications related to morphological identification of some midge specimens. • Models will be further refined, including a comparison of performance using an alternative source of temperature data (ENSEMBLES). The maximum catch of further Culicoides species will also be modelled and predicted (C. newsteadii and C. impunctatus). • Models of the timing and size of seasonal peaks of Culicoides abundance are also in development using the same data set and will be used to predict the seasonal dynamics of midges across the same geographical range. C. obsoletus complex C. imicola a b Key to midge abundance No prediction d C. pulicaris complex 0-10 c 0 0 00 00 ,0 0 000 , 0 0, 000 0 000 ,00 0, 000 ,00 0, 0 00 ,00 0, 00 ,00 0, 00 ,00 -1 -1 1 -1 -1 -1 -1 -1 -1 0 10 0 - 0 0 0 0 0 0 10 00 ,00 ,00 ,00 ,00 ,00 , 1 0 0 0 10 0 10 1,00 ,00 ,00 0 0 1 10 10 – 102 <VALUE> Mean outof-sample R2 Source Temperature Summed degree days >12°C per year 2. Model fitting • Poisson GLMM fit in INLA to identify the best-fitting models using covariates listed in Table 1. • Full subset model selection • All models also included year as a fixed effect and site-year as a random effect • Out-of-sample model fit (R2) was used to identify the best model, using a subset of data not used to fit the model. This process was iterated 20 times, resulting Figure 2: Location of midge trap sites. in 20 best models per species (Table 2). 3. Predictions • Predictions of the maximum annual abundance per species were made for each 1km pixel across Europe (Fig 3). • Predictions from each of the top 20 models (based on out of sample prediction performance) were averaged for each species, weighted by their relative support in the data. • Predictions were only made for pixels that were environmentally similar to the training data set, defined as those with a Malhalanobis Distance (MD) falling within the 0-90% cumulative distribution of MDs from the training data set. Table 2: Covariates included in the 20 best models of maximum annual Culicoides female trap abundance. Mean R2 of the 20 best models for each species is given. Number of times a predictor variable was included in the best model for a species is in brackets. Description 102 – 103 C. obsoletus complex 0.58 Elevation (20) Precipitation (dd) (20) Summed NDVI (20) Roe deer density (12) Precipitation (cvppt) (10) Red deer density (10) Sheep density (8) Cattle density (4) e 104 – 105 f 105 – 106 106 – 107 107 – 108 108 – 109 0 C. pulicaris complex 0.65 103 – 104 Temperature (20) Precipitation (cvppt) (11) Precipitation (dd) (11) Cattle density (8) Sheep density (6) Elevation (4) Precipitation (ppt) (2) Roe deer density (2) Red deer density (1) Figure 3: (a-c) Model averaged predictions of the mean maximum catch of three Culicoides species females in 2007 according to the 20 models with the best fit to the data. Predictions were made for 1km2 pixels across Europe. No predictions were made if a pixel was too environmentally dissimilar from the training data set according to Malhalanobis Distances (d) Previously modelled predictions of C. imicola presence taken from [4] (e) Previously identified range-limits of Culicoides spp. groups taken from [5], (f) Previously identified distribution of C. imicola and C. obsoletus complex in Italy taken from [6]. References: [1] Wint et al. (2014) Open Health Data 2: e1 [2] Alexander et al. (2014) Open Health Data 2: e2 [3] Robinson et al. (2014) PlosOne 9: e96084 [4] Purse et al. (2007). Journal of Applied Ecology 44: 1231-1242 [5] Purse et al. (2005) Nature Reviews Microbiology 3: 171-182 [6] Conte et al. (2007). Veterinary Parasitology 150: 333-344 [7] Purse et al. (2006). Medical and Veterinary Entomology 20: 335-344 [8] Capela et al. (2003). Medical and Veterinary Entomology 17: 165-177. WHY: develop models to predict the distribution and abundance of Culicoides spp. across Europe? BECAUSE: they are vectors of arboviruses that infect livestock and knowing their distribution and population dynamics will aid control efforts.
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