Abstract
Following the first record of Phlebotomine sand flies in Luxembourg in 2023, we performed during the summer 2024 the first extensive field survey investigating the sand fly fauna diversity and distribution in the Grand Duchy. Overall, 181 samples (insect batches/site/trapping method/date) were collected from 165 sites across the country (44 localities in 28 communes), mainly by running light traps but occasionally sticky traps and conducting resting catches. Only Phlebotomus mascittii was found, in a total of 15 samples collected at 10 localities, revealing 23 specimens (12 females and 11 males). More trapping effort might be implemented in the country to confirm the absence of Ph. perniciosus. Together with the first observations of 2023 (at two localities), the occurrence of Ph. mascittii has increased to 12 localities in total, which are widely distributed over south Luxembourg, but with an overall low density. This low density suggests the risk for Leishmania transmission in Luxembourg, if introduced, to be very low. The two northernmost localities where sand flies were detected are in the commune of Rosport-Mompach. Our data do not suggest northwards spread of sand flies, but they complete the distribution data north to the Alps, demonstrating they are more widespread than previously known. Further assessing the risk for sand fly-borne diseases in Luxembourg would request the acquisition of information on Leishmania infections in dogs for imported cases and, if existing, suspected locally acquired cases. Additional entomological field surveys may be implemented to further determine the local distribution of sand flies, the period of activity of adults, the abundance, or the presence of pathogens in insects when their circulation is suspected.
1 Introduction
Phlebotomine sand flies are vectors of Leishmania spp. and phleboviruses in the Mediterranean region where they are common, while they are rare north to the Alps. They are non-invasive insects, but there is evidence that they are expanding northwards and to higher altitudes as a result of climate change (Medlock et al., 2014). This was evidenced in particular for Phlebotomus (Larroussius) perniciosus Newstead, 1911 and Ph. (Larroussius) neglectus Tonnoir, 1921 in Italy (Maroli).
Phlebotomus (Transphlebotomus) mascittii Grassi, 1908, a suspected vector of Leishmania infantum (Nicolle, 1908) shows the widest and northernmost distribution in Europe (Kniha et al., 2023). The species has been sporadically reported in northern France and south-eastern Belgium (Depaquit et al., 2005; Schaffner, 2023) as well as in Germany, where it has been detected in the federal state of Rhineland-Palatinate, at the same latitudes as southern Luxembourg and southern Belgium (Oerther et al., 2020) but also more northern near Cochem (Nauke et al., 2008), Mörz and Lahnstein (Krüger et al., 2025). Also, both Ph. mascittii and Ph. perniciosus, the latter being the main L. infantum vector in Western Europe, were reported in the federal state of Baden-Württemberg, southern Germany (Nauke et al., 2004). Moreover, autochthonous L. infantum infections have been reported in the Städteregion Aachen (Germany), bordering Belgium and the Netherlands (Nauke et al., 2008). Similarly, in France, autochthonous L. infantum infections have been reported as far north as Indre-et-Loire, Loiret and Jura (Chamaillé et al., 2010; Kasbari et al., 2012). Similarly to other insects, climate change might favour some sand fly species, in particular Ph. mascittii, to spread more northern and to develop larger populations where they are already established (Medlock et al., 2014).
In 2023, a capacity building session together with a field study was organised by the European network for medical and veterinary entomology VectorNet, to promote sand fly survey in northern France and the Benelux countries (Risueño et al., 2024). During that session, four locations were surveyed in Luxembourg (Bech-Kleinmacher, Luxembourg-Grund, Machtum and Remich) and sand flies were detected in the Moselle valley, at both Bech-Kleinmacher and Remich locations (Figures 1, 3). In addition, sand flies were detected in bordering France (Conz-les-Bains) and Germany (Nittel) (Figure 1). These findings, obtained within a survey limited in time and space, encouraged performing additional investigations but with a stronger sampling effort to obtain a better image of the sand fly situation in the area.
Preliminary selection of localities to be investigated for the possible occurrence of Phlebotomine sand flies in Luxembourg, 2024. (A) With probability of presence model output (the more red, the more suitable habitat for Ph. mascittii); (B) Satellite image background (© Google Earth). Red and yellow pins: localities respectively with or without sand flies detected in 2023 in Luxembourg and neighbouring regions); White pins: suggested localities for the cross-sectional survey in 2024.
Citation: Journal of the European Mosquito Control Association 2025; 10.52004/2054930x-20251019
Subsequently, to better assess the risk to public and veterinary health related to the occurrence of sand flies in Luxembourg, we performed a field study during the summer 2024 with the objective to evaluate the distribution range of Phlebotomine sand flies in the country. This investigation was also performed to assess sand fly species diversity, i.e. if Ph. mascittii is the only species present or if congeners such as Ph. perniciosus are co-occurring.
2 Materials and methods
Study area
The Grand Duchy of Luxembourg is located between 49.45N and 50.18N latitude and 5.78E and 6.53E longitude covering an area of 2,586 km2. Besides the administrative divisions, Luxembourg is divided into two biogeographical regions (Oesling and Guttland) due to distinct geological, topographical and climatic conditions. The Oesling in the north of the country covers about 32% of the country’s area and belongs to the Ardennes and Eifel Mountain range. It is a plateau consisting of slate rocks and quartzites with an average altitude of about 450 m a.s.l. which is cut by deep and steep valleys. It has the coolest and wettest climate of the country, with an annual mean temperature of 7.5 °C on the plateau and 9.0 °C in the valleys and a mean annual precipitation between 800 and 950 mm for the 1991-2020 normals (STATEC, 2022). South of the Oesling, the Guttland covers about 68% of the country’s area and belongs to the area of the Lorraine stratified plain. It is an undulating hilly landscape (average altitude of about 300 m a.s.l.) which was formed by a succession of hard sandstone and shell limestone strata and soft strata such as Keuper. The climate of the Guttland is warmer and in many areas drier than that of the Oesling, with a mean annual temperature between 9.5 and 10.7 °C and an annual precipitation varying between 750 and 1000 mm for the 1991-2020 normals (STATEC, 2022).
Sampling strategy
This study was a cross-sectional field survey based on the use of light traps (overnight), interception sticky traps (over several nights) and ad hoc resting catches at various locations that look suitable for sand flies. In a first step, trapping locations were selected based on a probability of presence model output for Ph. mascittii, i.e. at localities with theoretical high habitat suitability (Figure 1A) (based on Alten et al., 2016). In a second step, natural suitable sites (e.g. cliffs) or buildings (e.g. old barns) were identified at these localities on satellite images (Google Earth™) and on street panorama views (Google Street View™). Finally, on the field, the best trapping sites were selected at these localities but also at additional ones according to environmental features considered suitable for sand flies, i.e. old barns preferably with beaten earth floor and with poultry or dogs nearby, cellars with beaten earth floor, stone walls, cliff bases, rocks or caves (Feliciangeli et al., 2004; Risueño et al., 2017), and to their accessibility. The initial plan was to investigate at least 16 localities (Figure 1B) during one period of eight days in July 2024 (16–23/07/24). Though, a second field trip could be organised in August 2024 to perform additional trapping at the same and at new localities, at the occasion of the completion of a VectorNet capacity building session on mosquitoes and sand fly collection and identification (20–30/08/24).
Sand fly sampling and identification
Sand flies were mainly collected by light traps (LT), run overnight. Two LT models were used, i.e. CDC-like traps, assembled by the entomology team of the laboratory of Parasitology-Mycology, University of Reims Champagne-Ardenne, powered by four HR6/AA 1.2 V / >2000 mAh Ni-MH rechargeable batteries (Figure 2B) and Laika traps (LaikaLab, Castiglione-del-Lago, Italy) powered by a 12 V / 7.2 Ah lead battery (Figure 2A). The Laika traps were modified by removing the UV LED, to avoid catching non-target insects such as moths, and when the white light LED was defective, we added a battery-operated warm yellow LED light candle at the trap entrance (Figure 2A). All traps were used with a net ending into a plastic vial containing 80 ml of water and two droplets of dish soap, in which the insects were sinking. Interception sticky traps (ST), consisting in 80 g white 20×20 cm paper sheets impregnated with castor oil were occasionally placed in stone wall holes for 4 to 5 consecutive nights (Figure 2C). In addition, a few resting catches (RC) were performed with a hand-made mechanical aspirator in holes of stone walls, in basements or around human (Figure 2D). The LTs were hung indoor or outdoor old barns and cellars, or in proximity of poultry houses, cliffs, rocks or stone walls, at 0.5 to 1 m height, while the STs were fixed with wood sticks in drain holes or crevices of stone walls, according to standard procedures (Alten et al., 2015).
Sampling tools used to investigate for sand fly presence in Luxembourg, 2024. A- Laika trap with its 12V battery and the vial for collection in liquid, and with a battery-powered candle (top left vignette); B- CDC-like trap, with the vial for collection in liquid concealed by the black tissue; C- A sticky trap placed in a hole of a stone wall; D- A specimen of Phlebotomus mascittii female (centre) in between two mosquitoes, caught by aspirating in holes of a stone wall (Clemency, site SFLU-031A, 23/07/2024).
Citation: Journal of the European Mosquito Control Association 2025; 10.52004/2054930x-20251019
Photo credits: Francis Schaffner.Sand flies were extracted from LT sample liquid under a stereomicroscope and placed in 70% ethanol before further processing, as were other Psychodidae and Culicidae (data not shown here). ST paper sheets were inspected using a LED lighted head visor magnifier (Carson Pro® CP-60, Carson Optical Inc., Rijan, Netherlands) and sand fly specimens were gently removed from with a tiny forceps after application of a droplet of ethanol and placed in 70% ethanol. Insect catches obtained by RC were left in a freezer for 30 min. before sand flies were sorted and placed in 70% ethanol. Later, specimens were dissected (head and genitalia) and mounted on glass slides in CMCP-10 mounting media (Polysciences Europe GmbH, Eppelheim, Germany) for morphological identification which was performed based on available identification keys (Dantas-Torres et al., 2014; Fauran et al., 1998; Gatt et al., 2009).
Statistics
Statistics were performed on catches and trapping methods, using a data set that includes in addition to our data from Luxembourg also data from Belgium and France, all collected in summer 2024 with the same methods and material. This allowed to increase the number of data and thus strengthen the analysis. However, the RC and ST trap catches were excluded since they provided too little data, i.e. these methods were only used six times, resulting in limited sample size. The effect of trap type and place (indoor, outdoor or at indoor/outdoor interface) on the collected sand flies was investigated with a generalised linear model (GLM) with negative binomial distribution with log link and dispersion estimated (Schaffner et al., 2023). Data were weighted for sampling effort by the number of collection LT/nights. Effects and their two-way interactions were fitted in the GLM, and nonsignificant factors were removed. Models were compared and the best model selected using the corrected Akaike’s information criterion (AIC). Differences between trapping methods and trap placement were further tested when significant in the GLM using post-hoc pairwise comparisons with Least Square Differences (LSD) correction. P-values below 0.05 were considered significant. All statistical analyses were performed using SPSS® statistics software (Version 29.0.0, IBM Corp., Armonk, NY, USA).
3 Results and discussion
Sampling effort
Sand fly sampling was performed in both biogeographical regions of Luxembourg, but a single locality only was investigated in Oesling (Figure 3). This was due to the much lower theoretical habitat suitability of the Oesling compared to the Guttland (Figure 1A). Overall, we sampled at 44 localities, distributed over 28 communes out of 100 and 10 cantons out of 12 that form the Grand Duchy. The sampling effort consisted in 163 LT/nights, with from 1 to 13 LT/nights per locality (median = 4 LT/nights per locality), 16 RC (12 in stone walls holes and cellars, 1 in cliff holes, 1 in castle basement, 2 around human at night), and 2 ST sessions, with a total of 29 paper sheets run for 4 or 5 consecutive nights (Table 1).
Sampling effort of our sand fly cross-sectional survey in Luxembourg, summer 2024.1
Citation: Journal of the European Mosquito Control Association 2025; 10.52004/2054930x-20251019
Sampling results
Sand flies were detected in 15 samples collected at 10 localities, revealing 23 specimens (Table 2). All specimens, including 12 females and 11 males, belong to Ph. mascittii and are stored in the microscopic slide collection of the National Museum of Natural History Luxembourg (collection numbers MNHNL57206 to MNHNL57222). Specimens were caught in small quantities, i.e. 1 or 2 specimens per trap night only, with an average density at positive sites of 0.14 specimens per LT/night, which is similar to the density observed for the same species in Germany (0.17; Oerther et al., 2020) but lower than in southern Switzerland (0.54; Schaffner et al., 2024). With ST, the average density was 1.15 specimens per m2, which is higher than the density observed in north-eastern France and in southern Switzerland for that species (0.65 and 0.66, respectively; Schaffner, 2023; Schaffner et al., 2024). Though, these densities are much lower than the densities observed in the leishmaniasis endemic regions for other sand fly species, such as in the French Mediterranean region, where the density of Ph. (Larroussius) ariasi Tonnoir, 1921 reached up to 50.53 (Prudhomme et al., 2015). The low density revealed for Luxembourg suggests the risk for Leishmania transmission, if introduced, to be very low.
Characteristics of the localities where sand flies (SF) were detected in Luxembourg, summer 2024, and numbers of specimens collected.1
Citation: Journal of the European Mosquito Control Association 2025; 10.52004/2054930x-20251019
The two sites where sand flies were detected in 2023, at Bech-Kleinmacher (SFLU-004) and Remich (SFLU-002), were investigated again in 2024 but no sand fly was caught (3 and 4 LT/nights, respectively). However, sand flies were detected at another site in Remich (SFLU-011). The third locality, which was investigated in 2023 without detecting any sand fly in the 12 LT/nights performed, Luxembourg-Grund (SFLU-001), revealed one sand fly specimen in 2024 caught by STs. The fourth locality where traps were run in 2023 was not investigated in 2024.
Samples were collected within two periods of 8 days in July and 10 days in August 2024, allowing the collections of 98 and 83 samples, of which 8 and 7 (8.2 and 8.4%) revealed positive, respectively. Six localities were investigated during both periods, and two revealed sand fly presence in both periods, namely Remich (SFLU-011) and Born (SFLU-017).
LTs and RCs were positive at 7.4 and 6.3%, respectively, while the two performed ST sessions revealed positive. This suggests a trapping over several consecutive days to be more efficient in sand fly detection and hence, it can be suggested to adjust the sampling strategy of a follow-up field study by running more ST sessions. The type of LT used did not significantly influence the number of Ph. mascittii caught (GLM, df = 310, P = 0.632). The CDC-like trap (72 trap/nights) and the Laika traps (91 trap/nights) showed almost similar attractivity, with 6.9 and 7.7% of the trap/nights, respectively, that revealed positive. The Laika traps run with the original LED light (51 trap/nights) or with the LED candle (40 trap/nights) were positive at 5.9 and 10.0%, respectively. Hence, the Laika traps with LED were run in July while the Laika traps with candle were run in August, but the overall results obtained during both periods showing to be similar, the trapping period may not influence the trap efficiency comparison. The CDC-like LT revealed more handy to use, thanks to the small and light battery block, compared to the large and heavy 12V battery of the Laika LT. However, the use of rechargeable batteries of >2,000 mAh is recommended to ensure the power source for 20 h, and the use of fast chargers is essential to allow traps to be reset the following afternoon. The trap design of the CDC-like LT is also more functional than the one of the Laika LT since the fan speed and size of the latter leads to the destruction of all insects larger than 5 mm. The yellow candlelight shows to efficiently attract sand flies despite the low light flux. As a result, we could recommend further using the CDC-like trap but maybe by using a reduced LED light flux, to bring down the insect by-catch and the power consumption.
LTs were run indoor (e.g. barn, cellar; 10 trap/nights; 6.1%), outdoor (126 trap/nights; 77.3%), or at the indoor/outdoor interface (e.g. open barn, ruin with destroyed roof, cellar or barn entrance; 27 trap/nights; 16.6%) and found positive at 30.0, 4.8 and 11.4%, respectively. Placement of the trap indoors or outdoors did influence the trap catches of Ph. mascittii (GLM, df = 312, P<0.001). Traps that were placed outdoor caught a lower number of sand flies compared to indoor or indoor/outdoor (GLM, df = 312, P <0.011). Hence further studies aiming at detecting sand flies at additional locations may better perform trapping indoor or at open barns/cellars or their entrance.
Sampling localities were located on an altitude between 148 and 411 m a.s.l., while sand flies were detected in the range 148 to 359 m a.s.l.. Positive localities are distributed from east to west of Luxembourg, but mainly in the southernmost quarter of the country (Figure 3). The two northernmost localities where sand flies were detected are in the commune of Rosport- Mompach, localities of Born (10 specimens; 6.514539E, 49.760627N, altitude 150 m a.s.l.) and Osweiler (1 specimen only; 6.440222E, 49.775282N, altitude 359 m a.s.l.).
Location of sampling sites and results of our cross-sectional sand fly survey in Luxembourg province of Belgium, 2024, together with record of 2023. White disks: no sand fly detected in 2024; Black disks: sand fly detected in 2024; Black triangles: sand flies detected in 2023; White triangle: no sand flies detected in 2023 (Risueño et al., 2024).
Citation: Journal of the European Mosquito Control Association 2025; 10.52004/2054930x-20251019
The 181 samples, all methods, were collected from a total of 165 sites that can be grouped into 6 types: (1) chicken run or goat fen, (2) cliffs or rocks, in limestone or sandstone, (3) forest, (4) house or farm courtyard, (5) unused house or barn, ruin, diverse shelters, or castle ruin, and (6) stone walls (Figures 4 and 5). Sand flies were mainly detected at unused houses or barns (n=6; 12.2% of the investigated sites in that category), but also in/at stone walls (n=4; 14.3%), at cliffs or rocks (n=3; 4.2%). The barns had a beaten earth (n=4) or cobblestones floor (n=1) or both (n=1). The stone walls revealed sand flies in inspected holes in nature (n=1) or in rural (n=2) or urban environment (n=1), with grass and trees in front of the wall (n=2), or asphalt only (n=2). The cliffs all consisted in limestone (n=3), shaded by trees or bushes (n=2) or open (n=1). The catch of a sand fly in a forest is surprising since this environment is not considered favourable to sand flies. However, it happened along a gravel road made with limestone substrate, of which the margin could provide suitable microhabitat. A second trapping performed the following day remained negative, and a trap contamination is unlikely considering the little number of sand flies caught during our survey.
Types of sites investigated for sand fly presence in Luxembourg, 2024, and results, for all trapping methods.
Citation: Journal of the European Mosquito Control Association 2025; 10.52004/2054930x-20251019
Examples of sites investigated for sand fly presence in Luxembourg, 2024, where Phlebotomus mascittii was detected (A–E) or not (F). (A) Open barn with beaten earth floor in Garnich (site SFLU-029A), on 22 July 2024; (B) Entrance of a cellar with beaten earth floor in Garnich (SFLU-029E), on 22 July 2024 ; (C) Stable with cobblestone floor in Filsdorf (SFLU-042B), on 25 August 2024; (D) Stone wall in nature/agriculture environment in Gostingen (SFLU-013C), on 25 August 2024; (E) Stone wall open onto a street in rural environment in Born (SFLU-017D), where sticky traps where used, on 20 August 2024; (F) Intermediary level of limestone cliffs, south to Hettermillen (SFLU-012A) in the Moselle valley, on 17 July 2024.
Citation: Journal of the European Mosquito Control Association 2025; 10.52004/2054930x-20251019
Photo credits: Francis Schaffner.With regards to the site environments, our findings are in line with records from neighbouring countries, where Ph. mascittii was recently found in house ruins (Belgium, 2024 and France, 2023; Risueño et al., 2024; Schaffner et al., 2025), near cliffs or rocks (Germany and France, 2023; Oerther et al., 2020; Risueño et al., 2024), and at old barns (France, 2023 and Germany, 2015-2018; Oerther et al., 2020; Risueño et al., 2024).
5 Conclusions
This study suggests the phlebotomine fauna of Luxembourg to be restricted to Ph. mascittii. However, more trapping effort might be implemented in the country to confirm the absence of Ph. perniciosus. Together with the first observations in 2023 (Risueño et al., 2024), the occurrence of Ph. mascittii is shown at 12 localities in total, which are widely distributed over south Luxembourg, but with an apparent low density. Our data do not suggest northwards spread of sand flies. They complete the first field information collected about sand fly occurrence in Luxembourg in 2023 and the distribution data of sand flies north to the Alps, demonstrating they are more widespread than previously known (Prudhomme et al., 2024). Additional investigations in northern France, Belgium, Southern Netherlands and Germany are necessary to understand the current distribution of sand flies north of the Alps and later on to assess the possible northward spread or the building up of larger populations as a consequence of climate change.
The vector role of Ph. mascittii for Leishmania parasites is not yet evidenced but suspected because of the report of autochthonous cases from its distribution area without evidence of presence of another or a better vector species (Nauke et al., 2008). Further assessing the risk for sand fly-borne diseases in Luxembourg would request the acquisition of information on Leishmania infections in dogs for imported cases and, if existing, suspected locally acquired cases. The entomological data gathered by our studies will support the planning of further studies, e.g. a further cross-sectional investigation at additional suitable sites or a longitudinal survey to assess the period of activity of adults, the abundance, or to screen for pathogens in insects in the leishmaniasis cases environment.
Supplementary material
Supplementary material can be found online at https://doi.org/10.6084/m9.figshare.28254140
Table S1. Detailed field data of Phlebotomine sand fly investigation in Luxembourg, 2024.
Acknowledgements
We acknowledge the project ‘European network for medical and veterinary entomology’ (VectorNet3) and its contracting agencies EFSA, the European Food Safety Authority, and ECDC, the European Centre for Disease Prevention and Control, for supporting the capacity building field training that was linked to this study (Framework contract number EFSA/2023/OP/0009 (OC/EFSA/BIOHAW/2023/05) and specific contract VECTORNET3 EFSA SC 01), with the valuable contribution of Cedric Marsboom, Vit Dvorak, and all trainees. We are also grateful to Jérôme Depaquit (University of Reims Champagne-Ardennes, Faculty of Pharmacy) for kindly lending trapping material, William Wint (Environmental Research Group Oxford) for providing the Ph. mascittii habitat suitability model output, Paul Braun (digital curator at MNHNL) for producing the distribution map (Figure 3), and Niels Verhulst (University of Zürich, Institute of Parasitology) for the statistical analysis. We thank all private ground and building owners who allowed us to access their propriety to place traps.
Conflict of interest
Francis Schaffner is editor-in-chief of the Journal of the European Mosquito Control Association; he had no influence in the review process and decision making on this manuscript. The other co-author declares no conflict of interest.
Data availability
The detailed data that supports the findings of this study are available in the supplementary material of this article.
References
Alten, B., Ozbel, Y., Ergunay, K., Kasap, O.E., Cull, B., Antoniou, M., Velo, E., Prudhomme, J., Molina, R., Bañuls, A.L., Schaffner, F., Hendrickx, G., Van Bortel, W. and Medlock J.M., 2015. Sampling strategies for phlebotomine sand flies (Diptera: Psychodidae) in Europe. Bulletin of Entomological Research 105: 664–678. https://doi.org/10.1017/s0007485315000127
Alten, B., Versteirt, V., Van Bortel, W., Zeller, H., Wint, W. and Alexander, N.S., 2016. VBORNET gap analysis: Sand fly vector distribution models utilised to identify areas of potential species distribution in areas lacking records. Journal of Open Health Data 4(1): p.e5. https://doi.org/10.5334/ohd.26
Chamaillé, L., Tran, A., Meunier, A., Bourdoiseau, G., Ready, P. and Dedet J.-P., 2010. Environmental risk mapping of canine leishmaniasis in France. Parasites and Vectors 3: 31. https://doi.org/10.1186/1756-3305-3-31
Dantas-Torres, F., Tarallo, V.D. and Otranto, D., 2014. Morphological keys for the identification of Italian phlebotomine sand flies (Diptera: Psychodidae: Phlebotominae). Parasites and Vectors 7: 479. https://doi.org/10.1186/s13071-014-0479-5
Depaquit, J., Naucke, T.J., Schmitt, C., Ferté, H. and Léger, N., 2005. A molecular analysis of the subgenus Transphlebotomus Artemiev, 1984 (Phlebotomus, Diptera, Psychodidae) inferred from ND4 mtDNA with new northern records of Phlebotomus mascittii Grassi, 1908. Parasitology Research 95: 113–116. https://doi.org/10.1007/s00436-004-1254-x
Fauran, P., Izri, A., Delaunay, P. and Marty, P., 1998. Les phlébotomes (Diptera, Phlebotominae) des Alpes-Maritimes et de Monaco. [The phlebotomine sand flies (Diptera, Phlebotominae) of the Alpes-Maritimes and Monaco]. Riviéra Scientifique 1998: 41–48.
Feliciangeli, M.D., 2004. Natural breeding places of phlebotomine sandflies. Medical and Veterinary Entomology 18: 71–80. https://doi.org/10.1111/j.0269-283x.2004.0487.x
Gatt, P., Williams, J. and Mifsud, D., 2009. New distributional data on sandflies from rubble walls in the Maltese Islands with an illustrated key to the Maltese species (Diptera: Phlebotominae). Bulletin of the Entomological Society of Malta 2: 95–110.
Kasbari, M., Ravel, C., Harold, N., Pesson, B., Schaffner, F. and Depaquit, J., 2012. Possibility of leishmaniasis transmission in Jura, France. Emerging Infectious Diseases 18: 1030. https://doi.org/10.3201/eid1806.120158
Kniha, E., Dvořák, V., Koblmüller, S., Prudhomme, J., Ivović, V., Hoxha, I., Oerther, S., Heitmann, A., Lühken, R., Bañuls, A.L., Sereno, D., Michelutti, A., Toniolo, F., Alarcón-Elbal, P.M., Bravo-Barriga, D., González, M.A., Lucientes, J., Colella, V., Otranto, D., Bezerra-Santos, M.A., Kunz, G., Obwaller, A.G., Depaquit, J., Alić, A., Kasap, O.E., Alten, B., Omeragic, J., Volf, P., Walochnik, J., Sebestyén, V. and Trájer, A.J., 2023. Reconstructing the post-glacial spread of the sand fly Phlebotomus mascittii Grassi, 1908 (Diptera: Psychodidae) in Europe. Communications Biology 6: 1244. https://doi.org/10.1038/s42003-023-05616-1
Krüger, A., Crecelius, A., Fischer, D. and Hagen R.M., 2025. New records of Phlebotomus (Transphlebotomus) mascittii Grassi, 1908 in northern Rhineland-Palatinate (Diptera: Psychodidae, Phlebotominae). Contributions to Entomology: In press.
Maroli, M., Rossi, L., Baldelli, R., Capelli, G., Ferroglio, E., Genchi, C., Gramiccia, M., Mortarino, M., Pietrobelli, M. and Gradoni, L., 2008. The northward spread of leishmaniasis in Italy: evidence from retrospective and ongoing studies on the canine reservoir and phlebotomine vectors. Tropical Medicine and International Health 13: 256–264. https://doi.org/10.1111/j.1365-3156.2007.01998.x
Medlock, J.M., Hansford, K.M., Van Bortel, W., Zeller, H. and Alten, B., 2014. A summary of the evidence for the change in European distribution of phlebotomine sand flies (Diptera: Psychodidae) of public health importance. Journal of Vector Ecology 39: 72–77. https://doi.org/10.1111/j.1948-7134.2014.12072.x
Naucke, T.J. and Schmitt, C., 2004. Is leishmaniasis becoming endemic in Germany? International Journal of Medical Microbiology 293 (Suppl. 37): 179–181. https://doi.org/10.1016/s1433-1128(04)80036-6
Naucke, T.J., Menn, B., Massberg, D. and Lorentz, S., 2008. Sandflies and leishmaniasis in Germany. Parasitology Research 103 (Suppl. 1): S65–S68. https://doi.org/10.1007/s00436-008-1052-y
Oerther, S., Jöst, H., Heitmann, A., Lühken, R., Krüger, A., Steinhausen, I., Brinker, C., Lorentz, S., Marx, M., Schmidt-Chanasit, J., Naucke, T. and Becker, N., 2020. Phlebotomine sand flies in Southwest Germany: an update with records in new locations. Parasites and Vectors 13: 173. https://doi.org/10.1186/s13071-020-04058-6
Prudhomme, J., Rahola, N., Toty, C., Cassan, C., Roiz, D., Vergnes, B., Thierry, M., Rioux, J.-A., Alten, B., Sereno, D. and Bañuls, A.-L., 2015. Ecology and spatiotemporal dynamics of sandflies in the Mediterranean Languedoc region (Roquedur area, Gard, France). Parasites and Vectors 8: 642. https://doi.org/10.1186/s13071-015-1250-2
Prudhomme, J., Depaquit, J. and Robert-Gangneux F., 2024. Phlebotomine sand fly distribution and abundance in France: A systematic review. Parasite 31: 45. https://doi.org/10.1051/parasite/2024045
Risueño, J., Bersihand, S., Bender. C., Cornen, T., De Boer, K., Ibáñez-Justicia A., Rey, D., Rozier, Y., Schneider, A., Stroo, A. Vanslembrouck, A., Van Bortel, W., Weigand, A., Zambianchi, D., Pérez Cutillas, P., Oerther, S., Braks, M., Wint, W.G.R., Berriatua, E. and Schaffner, F., 2024. A survey of Phlebotomine sand flies across their northern distribution range limit in Western Europe. Journal of the European Mosquito Control Association, 42(2): 105–115. https://doi.org/10.52004/2054930x-20241008
Risueño, J., Muñoz, C., Pérez-Cutillas, P., Goyena, E., Gonzálvez, M., Ortuño, M., Bernal, L.J., Ortiz, J., Alten, B. and Berriatua, E., 2017. Understanding Phlebotomus perniciosus abundance in south-east Spain: assessing the role of environmental and anthropic factors. Parasites and Vectors 10: 189. https://doi.org/10.1186/s13071-017-2135-3
Schaffner, F., 2023. Occurrence of Phlebotomus mascittii (Diptera: Psychodidae: Phlebotominae) in the upper Rhine Valley of Alsace, France. Annales de la Société Entomologique de France 59(4): 278–284. https://doi.org/10.1080/00379271.2023.2228741
Schaffner, F., Schneider, A., Debbaut, R. and Van Bortel, W., 2025. Confirmation of the occurrence of Phlebotomine sand flies (Diptera: Psychodidae, Phlebotominae) in Belgium, Luxembourg Province. Journal of the European Mosquito Control Association, in press. https://doi.org/10.52004/2054930x-20251021
Schaffner, F., Silaghi, C., Verhulst, N.O., Depaquit, J. and Mathis, A., 2024. The Phlebotomine sand fly fauna of Switzerland revisited. Medical and Veterinary Entomology 38: 13–22. https://doi.org/10.1111/mve.12690
STATEC, 2022. The statistics portal. https://statistiques.public.lu
