Bat parasites (Acari, Anoplura, Cestoda, Diptera, Hemiptera, Nematoda, Siphonaptera, Trematoda) in France (1762-2018): a literature review and contribution to a checklist.

This paper is a bibliographical survey of records of bat parasites in France (including Corsica) between 1762 and 2018. In total, 237 scientific publications were analysed. They show that bats are infected with a large diversity of endoparasites and ectoparasites. A total of 113 parasite taxa were identified from 27 host species; in addition, six bats were not identified to the species-level. The helminth fauna of bats comprises three species of Cestoda, 15 of Trematoda, and 13 of Nematoda. Acari parasites include 53 species (in addition to 22 invalid species). Finally, insect parasites comprise 13 species of Diptera (bat flies), 12 of Siphonaptera (fleas), 3 of Hemiptera (bugs), and 1 Anoplura species. Bat taxa reported with parasites were Barbastella barbastellus, Eptesicus serotinus, Hypsugo savii, Miniopterus schreibersii, Myotis bechsteinii, M. blythii, M. capaccinii, M. dasycneme, M. daubentonii, M. emarginatus, M. myotis, M. mystacinus, M. nattereri, M. punicus, Nyctalus lasiopterus, N. leisleri, N. noctula, Pipistrellus kuhlii, P. nathusii, P. pipistrellus, Plecotus auritus, P. austriacus, Rhinolophus euryale, R. ferrumequinum, R. hipposideros, R. mehelyi, Tadarida teniotis, Eptesicus sp., Myotis sp., Pipistrellus sp., Plecotus sp., Rhinolophus sp. and the species complex Pipistrellus pipistrellus/kuhlii/nathusii. As regards E. nilssonii, Vespertilio murinus (Particoloured Bat), M. alcathoe, M. escalerai, P. macrobullaris and P. pygmaeus, no records were found. These published field data originated from 72 of the 96 departments in metropolitan France. The most commonly cited were Ardèche, Ariège, Bouches-du-Rhône, Haute-Savoie, Maine-et-Loire, Moselle, Meurthe-et-Moselle, Pyrénées-Orientales, Sarthe, Haute-Corse and Corse-du-Sud. © C. Léger, published by EDP Sciences, 2020.

Introduction

Bats (Mammalia: Chiroptera) represent the second-most diverse order of mammals, after rodents. As of 2007, 42 bat species have been reported from Europe (Dietz et al. [84]). According to Arthur & Lemaire [16], 35 species have been unambiguously identified in France. Many aspects of the ecology of bats are under study (e.g. swarming, hunting sites, flight routes, habitat studies, acoustic ecology). One of these aspects is the study of bat parasites, which has a long history in Europe, for instance in French-speaking areas (France, Belgium). Bats are infected with a large diversity of parasites. Around the year 1999, c. 756 taxa were known to be associated with bats worldwide [167]. In Lanza’s book, a wide range of parasitic organisms were presented, belonging to 13 groups: Myconta (two taxa), Acanthocephala (three taxa), Mallophaga (one taxon, accidental exposure), Anoplura (two taxa), Heteroptera (11 taxa), Neobacteria (c. 31 taxa), Protozoa (25 taxa), Cestoda (55 taxa), Digenea (105 taxa), Nematoda (62 taxa), Acari (324 taxa), Diptera (65 taxa) and Siphonaptera (64 taxa). This includes at least ten phyla: Acanthocephala (Spiny-headed worms), Apicomplexa, Arthropoda, Ascomycota (Ascomycete fungi), Euglenozoa, Firmicutes, Nematoda, Platyhelminthes, Protobacteria and Spirochaetes. Similar findings were noted by Stiles & Nolan [235] in their “key catalogue” of bat parasites. In addition to the high diversity of bat parasites, these findings point out the predominant share, in the published records, of metazoan parasites. They also point out the issue of diseases in bats and the issue of bat parasites as disease vectors for their hosts. Indeed, we know that bats are hosts to a large range of infections (transmission linked with their ecology) and they seemingly are able to control these infections so that they are mostly asymptomatic. Some bat parasites (e.g. bat flies) are known to be disease vectors for their hosts [83, 129, 181, 184, 192, 261].

Among the earliest works on bat parasites in France is Étienne-Louis Geoffroy’s Histoire abrégée des insectes [123], published in 1762 (Fig. 1). This book marks the starting point for research on bat parasites in France. The present paper reviews metazoan parasites reported on bats in France between 1762 and 2018, with the exception of acanthocephalans. According to the Host-Parasite Database of the Natural History Museum, London [125], no bat parasites belonging to the Acanthocephala phylum are currently known in France. In addition, hyperparasites are excluded from this paper. Nevertheless, it should be noted that bat parasites have their own parasites, such as Laboulbeniales fungi associated with bat flies or viruses of haemosporidian parasites. Some of these hyperparasites have reports from France, specifically in the department of Gard (Arthrorhynchus eucampsipodae Thaxt., 1901 and A. nycteribiae (Peyr.) Thaxt., 1931) [55, 129, 238]. The purpose of the present paper is twofold: the primary aim is to summarize the large body of published field data; and secondly to inform the reader about the geographical origin of the data and to contribute to a general overview and checklist of bat-parasite associations in France.

Figure 1

Number of studies (n = 237) that include bat parasites observed in France since 1760, by decade.

Methods

Initially, I used the works of nine authors: Anciaux de Faveaux [5-7], Beaucournu [28, 29], Beaucournu and Launay [37], Hůrka [149], Lanza [167], Maa [185] and Szentiványi et al. [237]. The list of all the sources used in these papers offers an essential bibliographical guide. The online catalog of the Library of the Muséum National d’Histoire Naturelle (MNHN) was also used. I checked all available publications on each of the searched terms including a combination of France or the name of administrative departments (n = 111) or the names of former administrative regions of France (n = 22) with one of the generic names of the bat parasites, as mentioned in I parassiti dei pipistrelli (Mammalia, Chiroptera) della fauna italiana [167], Parasite diversity of European Myotis species with special emphasis on Myotis myotis (Microchiroptera, Vespertilionidae) from a typical nursery roost [121], Les puces de France et du basin méditerranéen occidental [37], and Checklist of host associations of European bat flies (Diptera: Nycteribiidae, Streblidae) [237]. For the study area, see Figure 5. I searched Google Scholar, ISI Web of Science, Hyper-Article en Ligne (HAL), Biodiversity Heritage Library (BHL), Gallica, and Archives. The collated sources (n = 237) were then analysed. I then proceeded to index them in terms of their chronology, taxonomy, and geography. The validity of all the taxa found was checked using the comprehensive synonymies provided by the Host-Parasite Database of the Natural History Museum, London [125] and the Inventaire National du Patrimoine Naturel of the Muséum National d’Histoire Naturelle (MNHN) [153]. The taxonomic works of Fain [106-108], Lanza [167], Neumann [197], Da Fonseca [72], Radovsky [212], Roy and Chauve [221, 222], Rudnick [223], Stiles and Nolan [235] and Theodor & Moscona [241] were also used. The bat classification and taxonomy, in this paper, is based on Dietz et al. (2009) and Arthur and Lemaire (2015). Authorities for the host taxa and parasites species are given in Tables 1 and 2. The map in Figure 5 was created using Carto-SI (https://www.carto-si.com/).

Figure 5

Study area and the number of publications that include data on parasites of bats in each French administrative region (department).

Table 1

List of bat species and their associated metazoan parasites in France (including Corsica), based on the published literature. Authors are listed in the bibliography. See also the work titled Les parasites métazoaires des Chiroptères de France (Acari, Anoplura, Cestoda, Diptera, Hemiptera, Nematoda, Siphonaptera, Trematoda) : contribution à un état des lieux bibliographique (1762–2018) et à l’établissement d’une liste nationale (2019). Invalid species are listed in brackets. Records marked with an exclamation mark (!) are invalid. Records marked with a question mark (?) are dubious. They may require further clarification.

Bat host	Number of parasite species reported	Parasite occurrence and citation
Barbastella barbastellus (Schreber, 1774)	10 species (9 recognised species and 1 invalid species)	Acari
		Absence of Ixodes canisuga Johnston, 1849 [28]
		Absence of Pholeoixodes hexagonus (Leach, 1815) [28]
		[Pteroptus vespertilionis L. Duf. [=Dufour] = Pteroptus vespertilionis (Dufour, 1832] [127, 177]
		[Pteroptus vespertilionis] [67]
		Acanthophthirius (Myotimyobia) pantopus (Poppe et Trouessart, 1895) [102, 114, 167, 235, 249]
		Argas vespertilionis (Latreille, 1796) [28]
		Ixodes (Eschatocephalus) vespertilionis C.L. Koch, 1844 [28, 32, 200]
		Spinturnix punctata (Sundevall, 1833) [80, 167]
		See also [128]
		Siphonaptera
		Ischnopsyllus (Hexactenopsylla) hexactenus (Kolenati, 1856) [2, 29, 33, 37, 38, 44, 200]
		Ischnopsyllus (Ischnopsyllus) octactenus (Kolenati, 1856) [200, 228]
		Ischnopsyllus (Ischnopsyllus) simplex Rothschild, 1906 [29, 37]
		Nycteridopsylla longiceps Rothschild, 1908 [29, 33, 37, 167]
		Nycteridopsylla pentactena (Kolenati, 1856) [2, 29, 33, 37, 44, 167, 200]
Chiroptera gen. sp.	30 species	Acari
	(22 recognised species and 8 invalid species)	[Acarus vespertilionis Hermann] [257, 258]
	1 innominate species	[Dermanyssus vespertilionis Dugès, 1834] [94]
		[Leiognathus armatus (=Hirstionyssus arcuatus (C.L. Koch, 1839)?)] [160, 177]
		[Liponyssus setosus (Kolenati)] [72]
		[Nycteribia vespertilionis Meig.] [12]
		[Pteroptus vespertilionis Dufour] [120, 177]
		[Pteroptus vespertilionis] [160]
		Argas (Secretargas) transgariepinus White, 1846 [189]
		Ixodes (Eschatocephalus) vespertilionis C.L. Koch, 1844 [45, 69, 160, 190, 197–199, 248]
		Macronyssus uncinatus (Canestrini, 1885) [160, 177]
		Neotrombicula autumnalis (Shaw, 1790) [62]
		Sasatrombicula (Sasatrombicula) hexasternalae (Vercammen-Grandjean, 1963) [7]
		Spinturnix myoti (Kolenati, 1856) [7, 65?, 167, 223, 246]
		Spinturnix plecotina (C.L. Koch, 1839) [7]
		Spinturnix psi (Kolenati, 1856) [7, 167, 210, 223]
		Diptera
		[Nycteribia vespertilionis Meig.] [12]
		Nycteribia (Nycteribia) pedicularia Latreille, 1805 ?[170]
		Phthiridium biarticulatum Hermann, 1804 ?[173, 204]
		Nycteribia (Nycteribia) latreillii (Leach, 1817) [148]
		Nycteribia (Nycteribia) schmidlii Schiner, 1853
		Penicillidia (Penicillidia) dufourii (Westwood, 1834) [43]
		Hemiptera
		Cimex dissimilis (Horváth, 1910) [207]
		Cimex lectularius Linnaeus, 1758 [28, 35]
		Cimex pipistrelli Jenins, 1839 [28, 236]
		Cimex sp. [28, 177]
		Nematoda
		Molinostrongylus alatus (Ortlepp, 1932)? [167]
		Siphonaptera
		Ischnopsyllus (Hexactenopsylla) hexactenus (Kolenati, 1856) [142]
		Ischnopsyllus (Ischnopsyllus) intermedius (Rothschild, 1898) [44]
		Ischnopsyllus (Ischnopsyllus) octactenus (Kolenati, 1856) [219]
		Nycteridopsylla dictena (Kolenati, 1856) ! see Beaucournu et Launay (1990; p. 314 [paper n°36]) [37, 146, 167, 227]
		Nycteridopsylla pentactena (Kolenati, 1856) [204]
Eptesicus serotinus (Schreber, 1774)	17 species	Acari
	(16 recognised species and 1 invalid species)	Absence of Ixodes canisuga Johnston, 1849 [28]
	1 innominate species	Absence of Pholeoixodes hexagonus (Leach, 1815) [28]
		[Pteroptus vespertilionis (Dufour, 1832)] [190]
		Argas (Carios) vespertilionis (Latreille, 1802) [31, 45, 86]
		Argas (Secretargas) transgariepinus White, 1846 [45, 189]
		Argas vespertilionis (Latreille, 1796) [28, 38]
		Ixodes (Eschatocephalus) vespertilionis C.L. Koch, 1844 [28, 32, 166]
		Notoedres (Notoedres) chiropteralis (Trouessart, 1896) [48, 106, 167, 250]
		Spinturnix psi (Kolenati, 1856) [48]
		Cestoda
		Vampirolepis acuta (Rudolphi, 1819) [158, 159, 177, 196, 255]
		Vampirolepis balsaci (Joyeux & Baer, 1934) [158, 159, 167, 177, 180, 196, 255]
		Vampirolepis sp./Milina sp. [86]
		Diptera
		Basilia (Basilia) mediterranea Hůrka, 1970 [43, 167, 237]
		Basilia (Basilia) nattereri (Kolenati, 1857) [241]
		Nycteribia (Nycteribia) kolenatii Theodor & Moscona, 1954 [147, 241]
		Siphonaptera
		Ischnopsyllus (Ischnopsyllus) intermedius (Rothschild, 1898) [29, 33, 37, 38, 142, 167, 227]
		Nycteridopsylla pentactena (Kolenati, 1856) [29, 33, 37, 164, 167]
		Trematoda
		Lecithodendrium linstowi Dollfus, 1931 [68, 88, 167]
		Plagiorchis vespertilionis (O.F. Müller, 1784) [68, 167]
		Prosthodendrium (Prosthodendrium) longiforme (Bhalerao, 1926) [88, 167]
Eptesicus sp.	1 recognised species	Acari
		Neotrombicula racovitzai Feider, 1970 [6, 118, 167]
Hypsugo savii (Bonaparte, 1837)	4 recognised species	Acari
		Argas (Carios) vespertilionis (Latreille, 1802) [45]
		Argas (Secretargas) transgariepinus White, 1846 [45, 189]
		Spinturnix nobleti Deunff, Volleth, Keller et Aellen, 1990 [81, 167]
		Diptera
		Basilia (Basilia) mediterranea Hůrka, 1970 [41, 167, 237]
Miniopterus schreibersii (Natterer in Kuhl, 1817)	38 species	Acari
	(33 recognised species and 5 invalid species)	[Pteroptus vespertilionis (Dufour, 1832)] [203]
		[Spinturnix vespertilionis] [24]
		[Spinturnix vespertilionis (C.L. Koch)] [70]
		Argas vespertilionis (Latreille, 1796) ? [85]
		Calcarmyobia rhinolophia (Radford, 1940)? There is only one case. According to Lanza (1999), this field data published by Beron (1971) is doubtful: it could be Calcarmyobia dusbabeki Uchikawa, 1985 [50, 51, 167]
		Eyndhovenia euryalis (G. Canestrini, 1885) [76]
		Ichoronyssus diversipilis Vitzthum, 1920 [24]
		Ichoronyssus spinosus (Oudemans) 1902) [24, 70, 72]
		Ixodes (Eschatocephalus) simplex simplex Neumann, 1906 [15, 31, 32, 45, 51, 143, 166, 167, 176] see Beaucournu (1966; p. 498) paper n°30
		Ixodes (Eschatocephalus) vespertilionis C.L. Koch, 1844 [6, 28, 176]! see Beaucournu (1966; p. 498) paper n°30 [200, 242]
		Macronyssus granulosus (Kolenati, 1856) [50, 167]
		Spinturnix myoti (Kolenati, 1856) [76]
		Spinturnix psi (Kolenati, 1856) [50, 63, 76, 139]
		Cestoda
		Milina grisea van Beneden, 1873 [68, 86, 167, 200?]
		Diptera
		[Nycteribia vespertilionis Meig.] [203]
		Nycteribia (Nycteribia) pedicularia Latreille, 1805 ! [85]
		Phthiridium biarticulatum Hermann, 1804 ? [85, 126, 237]
		Nycteribia (Nycteribia) kolenatii Theodor & Moscona, 1954 [200]
		Nycteribia (Nycteribia) latreillii (Leach, 1817) [1, 167]
		Nycteribia (Nycteribia) schmidlii Schiner, 1853 [1, 24, 28, 30, 43, 115, 149, 154, 167, 242]; see also [117, 150]
		Nycteribia (Achrocholidia) vexata Westwood, 1835 [1, 43, 85, 167]
		Penicillidia (Penicillidia) dufourii (Westwood, 1834) [1, 24, 30, 237]
		Penicillidia (Neopenicillidia) conspicua Speiser, 1901 ([1, 28, 30, 115, 154, 167, 242], see also [117, 150, 237])
		Nematoda
		Litomosa ottavianii Lagrange & Bettini, 1948 [19, 167]
		Molinostrongylus panousei Dollfus, 1954 [99, 167]
		Molinostrongylus tipula (Beneden, 1873) [200]
		Riouxgolvania rhinolophi (Bain & Chabaud, 1968) [21, 167]
		Strongylacantha glycirrhyza van Beneden, 1873 [99, 167, 200]
		Trichosomum speciosum van Beneden, 1873 [200]
		See also [174]
		Siphonaptera
		Nycteridopsylla pentactena (Kolenati, 1856) [37, 164, 167]
		Trematoda
		[Paralecithodendrium chilostomum (Mehlis) (=Prosthodendrium (Prosthodendrium) chilostomum (Mehlis, 1831) ?)] [200]
		Lecithodendrium granulosum Looss, 1907 ? [200]
		Lecithodendrium linstowi Dollfus, 1931 [68, 167]
		Mesotretes peregrinus (Braun, 1900) [68, 167, 188]
		Parabascus lepidotus Looss, 1907 [68, 167]
		Plagiorchis vespertilionis (O.F. Müller, 1784) [68, 167, 200, 235]
		Prosthodendrium (Prosthodendrium) chilostomum (Mehlis, 1831) [68, 167, 200, 235]
		Prosthodendrium parvouterus (Bhalerao, 1926) [68, 167]
		See also [174]
Myotis bechsteinii (Leisler in Kuhl, 1817)	4 recognised species	Acari
		Absence of Ixodes canisuga Johnston, 1849 [28]
		Absence of Pholeoixodes hexagonus (Leach, 1815) [28]
		Ixodes (Eschatocephalus) vespertilionis C.L. Koch, 1844 [28, 32, 38]
		Spinturnix bechsteini (Deunff, Walter, Bellido et Volleth, 2004) [82, 130]
		Cestoda
		Vampirolepis balsaci (Joyeux & Baer, 1934) [159, 167, 177, 180, 196, 255]
		Diptera
		Basilia (Basilia) nana Theodor & Moscona, 1954 [4, 28, 30, 38, 237]
Myotis blythii (Tomes, 1857) and M. blythii oxygnathus Monticelli, 1885	9 recognised species	Acari
	1 innominate species	Absence of Ixodes (Eschatocephalus) simplex simplex Neumann, 1906 [32]
		Nycteridocoptes poppei Oudemans, 1898 [113]
		Spinturnix myoti (Kolenati, 1856) [14, 76]
		Diptera
		Nycteribia (Achrocholidia) vexata Westwood, 1835 [30]
		Nycteribia (Nycteribia) latreillii (Leach, 1817) [30, 167]
		Nycteribia (Nycteribia) pedicularia Latreille, 1805 [115, 154, 167]
		Nycteribia (Nycteribia) schmidlii Schiner, 1853 [30]
		Penicillidia (Penicillidia) dufourii (Westwood, 1834) [14, 43, 115, 154, 237]
		Nematoda
		Riouxgolvania nyctali (Bain & Chabaud, 1979) [21, 167]
		Riouxgolvania sp. [86]
		Siphonaptera
		Ischnopsyllus (Hexactenopsylla) hexactenus (Kolenati, 1856) ? [14]
Myotis capaccinii (Bonaparte, 1837)	9 recognised species	Acari
		Spinturnix myoti (Kolenati, 1856) [73]
		Spinturnix psi (Kolenati, 1856) [63, 76, 121, 139]
		Diptera
		Nycteribia (Nycteribia) pedicularia Latreille, 1805 [1, 30, 85]
		Penicillidia (Penicillidia) dufourii (Westwood, 1834) [1, 237]
		Siphonaptera
		Nycteridopsylla longiceps Rothschild, 1908 [37, 164, 167]
		Nycteridopsylla pentactena (Kolenati, 1856) [37, 164, 167]
		Trematoda
-		Lecithodendrium granulosum Looss, 1907 [68]
		Lecithodendrium linstowi Dollfus, 1931 [68, 167]
		Plagiorchis vespertilionis (O.F. Müller, 1784) [68, 167]
Myotis dasycneme (Boie, 1825)	1 recognised species	Acari
		Argas (Carios) vespertilionis (Latreille, 1802) [6, 31, 85]
Myotis daubentonii (Leisler in Kuhl, 1817) and Myotis daubentonii nathalinae	19 species (17 recognised species and 2 invalid species)	Acari
	1 innominate species	Absence of Ixodes canisuga Johnston, 1849 [28]
		Absence of Pholeoixodes hexagonus (Leach, 1815) [28]
		[Pteroptus trouessarti Mégnin (=Haemomyson trouessarti (Mégnin)?)] [248]
		Ixodes (Eschatocephalus) vespertilionis C.L. Koch, 1844 [28, 32]
		Leptotrombidium russicum (Oudemans, 1902) [122, 167, 177, 256] see also [10]
		Neotrombicula autumnalis (Shaw, 1790) [8–10, 122, 167, 177, 256]
		Nycteridocoptes poppei Oudemans, 1898 [113]
		Oudemansidium musca (Oudemans, 1906) [122, 167, 177]
		Spinturnix andegavinus (Deunff, 1977) [76]
		Diptera
		[Nycteribia eparticulata] [248]
		Nycteribia (Nycteribia) pedicularia Latreille, 1805 ! [85]
		Basilia (Basilia) nattereri (Kolenati, 1857) [187]
		Nycteribia (Nycteribia) kolenatii Theodor & Moscona, 1954 [1, 28, 38, 43, 167, 241]
		Nycteribia (Nycteribia) schmidlii Schiner, 1853 [147]
		Nycteribia sp. [247]
		Siphonaptera
		Ischnopsyllus (Ischnopsyllus) simplex Rothschild, 1906 [44]
		Ischnopsyllus (Ischnopsyllus) variabilis (Wagner, 1898) [44]
		Nycteridopsylla longiceps Rothschild, 1908 [29, 37, 167]
		Trematoda
		Allassogonoporus amphoraeformis (Mödlinger, 1930) [89, 167]
		Parabascus duboisi (Hurkova, 1961) [89, 167]
		Plagiorchis vespertilionis (O.F. Müller, 1784) [89, 167]
		Prosthodendrium (Prosthodendrium) hurkovaae Dubois, 1960 [89]
Myotis emarginatus (Geoffroy, 1806)	19 recognised species	Acari
	2 innominate species	Absence of Ixodes canisuga Johnston, 1849 [28]
		Absence of Pholeoixodes hexagonus (Leach, 1815) [28]
		Argas (Secretargas) transgariepinus White, 1846 [45, 189]
		Argas vespertilionis (Latreille, 1796) [28]
		Dermanyssus sp. [86]
		Eyndhovenia euryalis (G. Canestrini, 1885) [63, 121]
		Ixodes (Eschatocephalus) vespertilionis C.L. Koch, 1844 [28, 32, 38, 136, 137, 177]
		Spinturnix emarginatus (Kolenati, 1856) [7, 76, 78]
		Cestoda
		Milina grisea van Beneden, 1873 [68, 86, 167]
		Diptera
		Nycteribia (Achrocholidia) vexata Westwood, 1835 ! [23]
		Nycteribia (Nycteribia) pedicularia Latreille, 1805 ? [23]
		Basilia (Basilia) italica Theodor, 1954 [4, 34]
		Basilia (Basilia) nana Theodor & Moscona, 1954 [43, 237]
		Nycteribia (Nycteribia) latreillii (Leach, 1817) [22]
		Hemiptera
		Cimex lectularius Linnaeus, 1758 ? [253, 254]
		Cimex dissimilis (Horváth, 1910) [254]
		Nematoda
		Litomosa dogieli Bogdanov & Vladimirov, 1956 [19]
		Litomosa filaria (Beneden, 1873) [86, 167]
		Litomosa ottavianii Lagrange & Bettini, 1948 [68, 167]
		Rictularia sp. [86]
		Siphonaptera
		Ischnopsyllus (Ischnopsyllus) simplex Rothschild, 1906 [29, 37, 142]
		Trematoda
		Lecithodendrium linstowi Dollfus, 1931 [68, 167]
		Plagiorchis vespertilionis (O.F. Müller, 1784) [68, 86, 167]
Myotis myotis (Borkhausen, 1797)	38 species	Acari
	(33 recognised species and 5 invalid species)	Absence of Ixodes canisuga Johnston, 1849 [28]
	3 innominate species	Absence of Pholeoixodes hexagonus (Leach, 1815) [28]
		[Dermanyssus murinus (Lucas, 1840)] [101, 104, 167, 183, 195]
		[Liponyssus decussatus (Kolenati) (=Caris decussata Kolenati, 1856 ?)] [23]
		[Pteroptus vespertilionis (Dufour, 1832)] [90, 92]
		[Spinturnix vespertilionis (C.L. Koch)] [70]
		[Spinturnix vespertilionis] [25]
		Ixodes (Eschatocephalus) simplex simplex Neumann, 1906! [85]
		Ichoronyssus sp. ? [25]
		Macronyssus rhinolophi (Oudemans, 1902) ? [152, 212]
		Argas vespertilionis (Latreille, 1796) [28]
		Ixodes (Eschatocephalus) vespertilionis C.L. Koch, 1844 [23, 28, 32, 38, 154, 166]
		Nycteridocoptes poppei Oudemans, 1898 [24, 113]
		Spinturnix myoti (Kolenati, 1856) [14, 76, 78, 177, 226]
		Spinturnix psi (Kolenati, 1856) [76]
		Ichoronyssus spinosus (Oudemans, 1902)/Ichoronyssus scutatus (Kolenati, 1856) [24, 70, 72]
		Ixodes sp. [178]
		Spinturnix sp. [133]
		Steatonyssus sp. [152, 212]
		Cestoda
		Milina grisea van Beneden, 1873 [68, 159, 167]
		Diptera
		Nycteribia (Nycteribia) latreillii (Leach, 1817) [1, 28, 30]
		Nycteribia (Nycteribia) pedicularia Latreille, 1805 [25, 186]
		Nycteribia (Nycteribia) schmidlii Schiner, 1853 [1, 30]
		Nycteribia (Achrocholidia) vexata Westwood, 1835 [1, 23, 25, 28, 30, 38, 43, 167]
		Penicillidia (Neopenicillidia) conspicua Speiser, 1901 [25]
		Penicillidia (Penicillidia) dufourii (Westwood, 1834) [1, 14, 22, 28, 30, 43, 91, 186, 237, 242]
		Hemiptera
		Cimex dissimilis (Horváth, 1910) ? [177]
		Cimex lectularius Linnaeus, 1758 [253, 254]
		Nematoda
		Rictularia plagiostoma (Wedl, 1861)! [233]
		Litomosa dogieli Bogdanov & Vladimirov, 1956 [19]
		Litomosa filaria (Beneden, 1873) [75, 86, 167]
		Molinostrongylus ornatus (Monnig, 1927) [68]
		Molinostrongylus tipula (Beneden, 1873) [57]
		Pterygodermatites (Neopaucipectines) bovieri (Blanchard, 1886) [56, 167, 177, 211, 233, 235, 245]
		Siphonaptera
		[Pulex vespertilionis Dugès, 1832] [95]
		Nycteridopsylla longiceps Rothschild, 1908 ! [228]
		Ischnopsyllus (Ischnopsyllus) intermedius (Rothschild, 1898) [2, 29, 33, 37, 142, 167]
		Ischnopsyllus (Ischnopsyllus) octactenus (Kolenati, 1856) [25, 29, 37, 167]
		Ischnopsyllus (Hexactenopsylla) hexactenus (Kolenati, 1856) [14, 29, 33, 37, 167, 228]
		Ischnopsyllus (Ischnopsyllus) simplex Rothschild, 1906 [29, 37, 85]
		Nycteridopsylla pentactena (Kolenati, 1856) [29, 37, 167]
		Trematoda
		Lecithodendrium linstowi Dollfus, 1931 [68, 167]
		Plagiorchis vespertilionis (O.F. Müller, 1784) [86]
		Prosthodendrium (Prosthodendrium) chilostomum (Mehlis, 1831) [57, 61, 68, 167]
		See also [217, 220]
Myotis mystacinus (Leisler in Kuhl, 1817)	14 species	Acari
	(11 recognised species and 3 invalid species)	Absence of Ixodes canisuga Johnston, 1849 [28]
		Absence of Pholeoixodes hexagonus (Leach, 1815) [28]
		[Gamassus dermanyssoïdes (Mégnin)] [248]
		[Pteroptus trouessarti Mégnin (=Haemomyson trouessarti (Mégnin)?)] [248]
		[Pteroptus vespertilionis (Dufour, 1832)] [248]
		Ixodes (Eschatocephalus) simplex simplex Neumann, 1906! [85]
		Argas (Carios) vespertilionis (Latreille, 1802) [45]
		Argas vespertilionis (Latreille, 1796) [28]
		Ixodes (Eschatocephalus) vespertilionis C.L. Koch, 1844 [28, 32, 38, 85]
		Spinturnix mystacinus (Kolenati, 1857) [76, 78]
		Diptera
		Basilia (Basilia) italica Theodor, 1954 [34, 35, 43, 237]
		Basilia (Basilia) nana Theodor & Moscona, 1954 [241]
		Nycteribia (Nycteribia) kolenatii Theodor & Moscona, 1954 [147, 241]
		Siphonaptera
		Ischnopsyllus (Ischnopsyllus) simplex Rothschild, 1906 [2, 29, 33, 37, 44, 85?, 142, 156]
		Nycteridopsylla longiceps Rothschild, 1908 [29, 37, 38, 167]
		Nycteridopsylla pentactena (Kolenati, 1856) [29, 37, 167]
Myotis nattereri (Kuhl, 1817)	8 recognised species	Acari
		Absence of Ixodes canisuga Johnston, 1849 [28]
		Absence of Pholeoixodes hexagonus (Leach, 1815) [28]
		Ixodes (Eschatocephalus) vespertilionis C.L. Koch, 1844 [28, 32]
		Spinturnix myoti (Kolenati, 1856) [63, 121]
		Diptera
		Basilia (Basilia) nattereri (Kolenati, 1857) [4, 28, 30, 43, 187]
		Basilia (Basilia) nana Theodor & Moscona, 1954 [4, 30, 237]
		Nycteribia (Nycteribia) kolenatii Theodor & Moscona, 1954 [28, 147]
		Siphonaptera
		Ischnopsyllus (Ischnopsyllus) simplex Rothschild, 1906 [29, 33, 37, 44, 85, 142]
		Nycteridopsylla longiceps Rothschild, 1908 [29, 37, 167]
		Nycteridopsylla pentactena (Kolenati, 1856) [29, 37, 167]
Myotis punicus Felten, Spitzenberger & Storch, 1977	1 recognised species	Spinturnix myoti (Kolenati, 1856) [64]
Myotis sp.	4 recognised species	Acari
		Argas vespertilionis (Latreille, 1796) [171, 242]
		Spinturnix myoti (Kolenati, 1856) ? [14]
		Hemiptera
		Cimex dissimilis (Horváth, 1910) [253]
		Siphonaptera
		Ischnopsyllus (Hexactenopsylla) hexactenus (Kolenati, 1856) [44]
Nyctalus lasiopterus (Schreber, 1780)	1 recognised species	Acari
		Pteracarus pipistrellius maximis Uchikawa, 1989 [251]
Nyctalus leisleri (Kuhl, 1817)	2 recognised species	Acari
		Argas (Carios) vespertilionis (Latreille, 1802) [45]
		Siphonaptera
		Ischnopsyllus (Ischnopsyllus) intermedius (Rothschild, 1898) [44]
Nyctalus noctula (Schreber, 1774)	14 species (8 recognised species and 6 invalid species)	Acari
		[Acarus vespertilionis Hermann, 1804 - la Mite de la chauve-souris] [135]
		[Caris vespertilionis = Argas vespertilionis (Latreille, 1796) ?] ? [168, 171, 201]
		[Dermanyssus coriaceus Gervais (=Hirstionyssus arcuatus (Koch, 1839)?)] [11, 13, 103, 212, 221, 222, 258, 259]
		[Haemomyson trouessarti (Mégnin) (=Leiognathus arcuatus=Hirstionyssus arcuatus (Koch, 1839)? See [244]]
		[Pteroptus vespertilionis (Dufour, 1832)] [131, 177]
		[Pteroptus vespertilionis] [131]
		Hirstionyssus arcuatus (Koch, 1839) [11, 12, 103, 124, 127, 167, 177, 212, 221, 258, 259]
		Spinturnix helvetiae (Deunff, Aellen & Keller, 1986) [recorded at Col de Bretolet, at the border between France and Switzerland] [79]
		Spinturnix acuminatus (C.L. Koch, 1836) [recorded at Col de Bretolet, at the border between France and Switzerland] [79]
		Diptera
		Nycteribia (Nycteribia) pedicularia Latreille, 1805![85]
		Siphonaptera
		Ischnopsyllus (Ischnopsyllus) variabilis (Wagner, 1898) ? [37]
		Ischnopsyllus (Ischnopsyllus) elongatus (Curtis, 1832) [37, 42, 44, 167, 204, 228!, 231!]
		Ischnopsyllus (Ischnopsyllus) intermedius (Rothschild, 1898) [3, 44]
		Nycteridopsylla eusarca Dampf, 1908 [37, 42]
Pipistrellus kuhlii (Kuhl, 1817)	9 recognised species	Acari
		Absence of Ixodes canisuga Johnston, 1849 [28]
		Absence of Ixodes (Eschatocephalus) simplex simplex Neumann, 1906 [32]
		Absence of Pholeoixodes hexagonus (Leach, 1815) [28]
		Argas (Carios) vespertilionis (Latreille, 1802) [45]
		Argas vespertilionis (Latreille, 1796) [28, 166]
		Ixodes (Eschatocephalus) vespertilionis C.L. Koch, 1844 [28]
		Siphonaptera
		Ischnopsyllus (Hexactenopsylla) hexactenus (Kolenati, 1856) [29]
		Ischnopsyllus (Ischnopsyllus) octactenus (Kolenati, 1856) [29, 33, 37, 167]
		Ischnopsyllus (Ischnopsyllus) variabilis (Wagner, 1898) [37, 44, 142]
		Nycteridopsylla longiceps Rothschild, 1908 [37, 44, 156]
		Trematoda
		Lecithodendrium linstowi Dollfus, 1931 [86]
		Pycnoporus heteroporus (Dujardin, 1845) [86]
Pipistrellus nathusii (Keyserling & Blasius, 1839)	8 recognised species	Acari
		Absence of Ixodes canisuga Johnston, 1849 [28]
		Absence of Ixodes (Eschatocephalus) simplex simplex Neumann, 1906 [32]
		Absence of Pholeoixodes hexagonus (Leach, 1815) [28]
		Acanthophthirius (Acanthophthirius) poppei (Trouessart, 1895) [52, 102, 114, 167, 249]
		Argas (Carios) vespertilionis (Latreille, 1802) [45]
		Argas vespertilionis (Latreille, 1796) [28]
		Ixodes (Eschatocephalus) vespertilionis C.L. Koch, 1844 [28]
		Siphonaptera
		Ischnopsyllus (Ischnopsyllus) octactenus (Kolenati, 1856) [29, 33, 37, 167]
		Ischnopsyllus (Ischnopsyllus) variabilis (Wagner, 1898) [2, 29, 33, 37, 44]
		Nycteridopsylla longiceps Rothschild, 1908 [37, 44, 167]
		Nycteridopsylla pentactena (Kolenati, 1856) [44]
Pipistrellus pipistrellus (Schreber, 1774)	23 species	Acari
	(17 recognised species and 6 invalid species and one species complex)	Absence of Ixodes canisuga Johnston, 1849 [28]
		Absence of Ixodes (Eschatocephalus) simplex simplex Neumann, 1906 [32]
		Absence of Pholeoixodes hexagonus (Leach, 1815) [28]
		Absence of [Pteroptus vespertilionis] [67]
		[Argas caris] [191]
		[Caris vespertilionis = Argas vespertilionis (Latreille, 1796) ?] ? [258]
		[Gamasus pteroptoides (Mégnin)] [190]
		[Haemomyson trouessarti (Mégnin) (=Leiognathus arcuatus=Hirstionyssus arcuatus (Koch, 1839)? See [221]]
		[Laelaps (Iphis) agilis Koch, Berlese = Gamasus pteroptoides (Mégnin)? = Laelaps agilis C.L. Koch, 1836 ?] ([244], see also [190])
		[Pteroptus vespertilionis Hermann] [215]
		Argas (Carios) vespertilionis (Latreille, 1802) [31, 45, 177, 239]
		Argas vespertilionis (Latreille, 1796) [18, 28, 38, 67, 166]
		Ixodes (Eschatocephalus) vespertilionis C.L. Koch, 1844 [28, 86, 199]
		See also [201]
		Diptera
		Nycteribia (Nycteribia) pedicularia Latreille, 1805 ? [214, 215]
		Basilia (Basilia) mediterranea Hůrka, 1970 [40, 43, 167]
		Nycteribia (Nycteribia) kolenatii Theodor & Moscona, 1954 [241]
		Nycteribia (Nycteribia) schmidlii Schiner, 1853 [147]
		Nycteribia sp. - Complex of species Nycteribia kolenatii/latreillii/pedicularia [177, 214, 215]
		Siphonaptera
		Ischnopsyllus (Ischnopsyllus) octactenus (Kolenati, 1856) [2, 29, 33, 36, 37, 44, 142, 164, 167, 177, 227]
		Ischnopsyllus (Ischnopsyllus) variabilis (Wagner, 1898) [33, 37, 85]
		Nycteridopsylla ancyluris ancyluris Jordan, 1942 [33, 37, 44, 142, 157, 164]
		Nycteridopsylla longiceps Rothschild, 1908 [2, 29, 33, 37, 85, 167]
		Nycteridopsylla pentactena (Kolenati, 1856) [29, 37, 38, 167]
		Trematoda
		Lecithodendrium linstowi Dollfus, 1931 [68, 167]
		Parabascus semisquamosus (Braun, 1900) [60, 68, 167]
		Plagiorchis vespertilionis (O.F. Müller, 1784) [68, 86, 167]
		Pycnoporus heteroporus (Dujardin, 1845) [57, 61, 68, 97, 163, 167, 235]
Pipistrellus sp.	3 recognised species	Acari
Nota bene: according to Lord et al., for studies published before the 1990’s, Pipistrellus pipistrellus and Pipistrellus sp. specimens should be treated as potentially also including P. pygmaeus.		Argas (Carios) vespertilionis (Latreille, 1802) [45]
		Diptera
		Basilia (Basilia) mediterranea Hůrka, 1970 [41]
		Penicillidia (Penicillidia) dufourii (Westwood, 1834) [53]
Plecotus auritus (Linné, 1758)	16 recognised species	Acari
	(15 recognised species and 1 invalid species)	Absence of Ixodes canisuga Johnston, 1849 [28]
	1 innominate species	Absence of Ixodes (Eschatocephalus) simplex simplex Neumann, 1906 [32]
		Absence of Pholeoixodes hexagonus (Leach, 1815) [28]
		[Spinturnix vespertilionis] [70]
		Argas (Carios) vespertilionis (Latreille, 1802) [45]
		Argas vespertilionis (Latreille, 1796) [28]
		Ixodes (Eschatocephalus) vespertilionis C.L. Koch, 1844 [28, 199]
		Spinturnix plecotina (C.L. Koch, 1839) [63, 76, 167]
		See also [201]
		Cestoda
		Vampirolepis sp./Milina sp. [86]
		Diptera
		Basilia (Basilia) nana Theodor & Moscona, 1954 [28, 237]
		Basilia (Basilia) nattereri (Kolenati, 1857) [28, 187]
		Phthiridium biarticulatum Hermann, 1804 [85, 86, 237]
		Nematoda
		Litomosa filaria (Beneden, 1873) [19, 167]
		Seuratum mucronatum (Rudolphi, 1809) [54, 86, 194]
		Siphonaptera
		Ischnopsyllus (Ischnopsyllus) simplex Rothschild, 1906 ? [29, 37]
		Ischnopsyllus (Ischnopsyllus) variabilis ! (Wagner, 1898) [85]
		Ischnopsyllus (Hexactenopsylla) hexactenus (Kolenati, 1856) [2, 33, 37, 44, 142, 177, 227, 228]
		Nycteridopsylla longiceps Rothschild, 1908 [33, 37, 85, 167, 227]
		Nycteridopsylla pentactena (Kolenati, 1856) [29, 33, 37, 167]
		Trematoda
		Plagiorchis vespertilionis (O.F. Müller, 1784) [86, 167]
		Prosthodendrium sp. [86]
Plecotus austriacus Fischer, 1829	7 recognised species	Acari
		Absence of Ixodes (Eschatocephalus) simplex simplex Neumann, 1906 [32]
		Argas (Carios) vespertilionis (Latreille, 1802) [31]
		Argas (Secretargas) transgariepinus White, 1846 [45, 189]
		Spinturnix plecotina (C.L. Koch, 1839) [63, 76, 167]
		See also [128]
		Siphonaptera
		Ischnopsyllus (Hexactenopsylla) hexactenus (Kolenati, 1856) [33, 37, 44]
		Nycteridopsylla pentactena (Kolenati, 1856) [33, 37, 167]
		Trematoda
		Lecithodendrium linstowi Dollfus, 1931 [68, 167]
		Parabascus lepidotus Looss, 1907 [68, 167]
Plecotus sp.	1 recognised species	Nematoda
		Seuratum mucronatum (Rudolphi, 1809) [97]
Rhinolophus euryale Blasius, 1853	21 recognised species	Acari
		Absence of Ixodes canisuga Johnston, 1849 [28]
		Absence of Pholeoixodes hexagonus (Leach, 1815) [28]
		Eyndhovenia euryalis (G. Canestrini, 1885) [50, 63, 70, 76, 78, 105, 139, 167, 235]
		Ichoronyssus scutatus (Kolenati, 1856) [50, 65, 167, 205, 212, 235, 246]
		Ixodes (Eschatocephalus) vespertilionis C.L. Koch, 1844 [28, 32, 85, 166]
		Macronyssus granulosus (Kolenati, 1856) [51, 167]
		Neomyobia slovenica Dusbábek, 1969 [50–52, 167]
		Neotrombicula vandeli Kolebinova & Vercammen-Grandjean, 1971 [7, 162, 167]
		Paraperiglischrus rhinolophinus (C.L. Koch, 1841) [50, 76, 138, 167, 235]
		Riedlinia (Riedlinia) petarberoni (Kolebinova & Vercammen-Grandjean, 1970) [7, 51, 161, 167]
		Sasatrombicula (Sasatrombicula) hexasternalae (Vercammen-Grandjean, 1963) [51, 162, 167]
		Spinturnix myoti (Kolenati, 1856) [76]
		Spinturnix psi (Kolenati, 1856) [76]
		Diptera
		Nycteribia (Nycteribia) pedicularia Latreille, 1805 ! [85]
		Nycteribia (Achrocholidia) vexata Westwood, 1835! [85]
		Phthiridium biarticulatum Hermann, 1804 [28, 30, 38, 115, 167, 242]
		Hemiptera
		Cimex lectularius Linnaeus, 1758? [253, 254]
		Nematoda
		Litomosa ottavianii Lagrange & Bettini, 1948 [68, 167]
		Riouxgolvania rhinolophi (Bain & Chabaud, 1968) [20, 167]
		Siphonaptera
		Rhinolophopsylla unipectinata unipectinata (Taschenberg, 1880) [2, 29, 33, 142, 167]
		Trematoda
		Lecithodendrium linstowi Dollfus, 1931 [68, 167]
		Parabascus lepidotus Looss, 1907 [68, 167]
		Plagiorchis vespertilionis (O.F. Müller, 1784) [68, 167]
Rhinolophus ferrumequinum (Schreber, 1774)	50 species	Acari
	(44 recognised species and 6 invalid species)	Absence of Ixodes canisuga Johnston, 1849 [28]
	2 innominate species	Absence of Pholeoixodes hexagonus (Leach, 1815) [28]
		[Haemogamasus setosus (Kolenati)] [5]
		[Liponyssus arcuatus Koch (=Hirstionyssus arcuatus (C.L. Koch, 1839) (pro parte) et Steatonyssus murinus (Lucas, 1840) (pro parte)] [200]
		[Pteroptus vespertilionis Hermann] ([215], see also [177])
		[Pteroptus vespertilionis (Dufour, 1832)] [200]
		Macronyssus rhinolophi (Oudemans, 1902) ? [152, 212]
		Ixodes (Eschatocephalus) simplex simplex Neumann, 1906! [85]
		Macronyssus ellipticus (Kolenati, 1856) ! [152, 177, 212]
		According to Radovsky [212], this observation published by Husson & Daum [152] is dubious.
		Macronyssus longimanus (Kolenati) ! [152, 177, 212]: according to Radovsky [212], this observation published by Husson & Daum [152] is dubious.
		Alabidocarpus diceratops Lawrence, 1952 [5, 110, 167]
		Alabidocarpus megalonyx (Trouessart, 1895) [52, 111, 167, 208, 218, 249]
		Alabidocarpus minor (Rollinat & Trouessart, 1897) [52, 111, 167, 208, 218, 249]
		Argas (Carios) vespertilionis (Latreille, 1802) [45]
		Eyndhovenia euryalis (G. Canestrini, 1885) [50, 63, 76, 78, 167]
		Eyndhovenia euryalis oudemansi (Eyndhoven, 1941) [18, 105, 252]
		Ixodes (Eschatocephalus) vespertilionis C.L. Koch, 1844 [28, 32, 38, 45, 51, 58, 154, 167, 177, 199, 200, 214, 215, 218, 229, 239]
		Labidocarpus (Labidocarpus) rollinati Trouessart, 1895 [167, 218, 232]
		Neomyobia rollinati (Poppe, 1908) [50, 52, 167]
		Notoedres (Notoedres) chiropteralis (Trouessart, 1896) [106, 109, 167, 218, 250]
		Nycteridocoptes eyndhoveni Fain, 1959 [49, 107]
		Paraperiglischrus rhinolophinus (C.L. Koch, 1841) [50, 78, 167, 235]
		Psorergates rhinolophi Fain, 1959 [108, 167]
		Sasatrombicula (Sasatrombicula) hexasternalae (Vercammen-Grandjean, 1963) [51, 162, 167]
		Spinturnix psi (Kolenati, 1856) [139, 223]
		Steatonyssus spinosus Willmann, 1936 [50, 51, 167]
		See also [201]
		Cestoda
		Hymenolepis sp. [86]
		Milina grisea van Beneden, 1873 [68, 71, 159, 196, 200, 243]
		Diptera
		[Nycteribia vespertilionis Latreille] [73]
		Nycteribia (Achrocholidia) vexata Westwood, 1835 ! [85]
		Penicillidia (Penicillidia) dufourii (Westwood, 1834) ? [200]
		Brachytarsina flavipennis Macquart, 1851 [26, 40, 43, 117, 155, 237]
		Nycteribia (Nycteribia) schmidlii Schiner, 1853 [1]
		Nycteribia sp. [73, 74]
		Phthiridium biarticulatum Hermann, 1804 [1, 28, 30, 43, 85, 115, 126, 135, 151, 152, 154, 156, 175, 177, 237, 242]
		Hemiptera
		Cimex dissimilis (Horváth, 1910) [254]
		Cimex lectularius Linnaeus, 1758 [253, 254]
		Nematoda
		Litomosa ottavianii Lagrange & Bettini, 1948 [19, 167]
		Strongylacantha glycirrhyza van Beneden, 1873 [68, 167, 200, 243]
		Trichosomum speciosum van Beneden, 1873 [200]
		See also [174]
		Siphonaptera
		Ischnopsyllus (Ischnopsyllus) simplex Rothschild, 1906 ?[152, 177]
		Ischnopsyllus (Ischnopsyllus) intermedius (Rothschild, 1898) [29]
		Ischnopsyllus (Hexactenopsylla) hexactenus (Kolenati, 1856) [152]
		Nycteridopsylla pentactena (Kolenati, 1856) [29, 37, 152, 164, 167, 177]
		Rhinolophopsylla unipectinata unipectinata (Taschenberg, 1880) [2, 33, 38, 44, 142, 152, 156, 164, 167, 177, 227, 242]
		Trematoda
		[Paralecithodendrium chilostomum (Mehlis) (=Prosthodendrium (Prosthodendrium) chilostomum (Mehlis, 1831)?)] [200]
		Lecithodendrium granulosum Looss, 1907 ? [200]
		Lecithodendrium linstowi Dollfus, 1931 [68, 167, 243]
		Lecithodendrium moedlingeri (Pande, 1935) [243]
		Mesotretes peregrinus (Braun, 1900) [68, 71, 88, 167, 188, 243]
		Parabascus lepidotus Looss, 1907 [68, 167]
		Plagiorchis vespertilionis (O.F. Müller, 1784) [68, 71, 86–88, 167, 200, 243]
		Prosthodendrium (Prosthodendrium) carolinum Hurková, 1959 [167, 243]
		Prosthodendrium (Prosthodendrium) chilostomum (Mehlis, 1831) [200, 243]
		Prosthodendrium (Prosthodendrium) longiforme (Bhalerao, 1926) [71, 167, 243]
		See also [174]
Rhinolophus hipposideros (Bechstein, 1800)	21 species (19 recognised species and 2 invalid species)	Acari
	2 innominate species	Absence of Ixodes canisuga Johnston, 1849 [28]
		Absence of Pholeoixodes hexagonus (Leach, 1815) [28]
		[Pteroptus vespertilionis (Dufour, 1832)] [200]
		Ixodes sp. [178]
		Ixodes (Eschatocephalus) vespertilionis C.L. Koch, 1844 [23, 28, 32, 152, 154, 199, 200]
		Labidocarpus (Labidocarpus) rollinati Trouessart, 1895 ([167, 218] see also [52, 208])
		Leptotrombidium russicum (Oudemans, 1902) [161]
		Macronyssus rhinolophi (Oudemans, 1902) [50, 152, 167, 212]
		Neomyobia chiropteralis chiropteralis (Michael, 1884) [50, 52, 100, 114, 167]
		Paraperiglischrus rhinolophinus (C.L. Koch, 1841) [50, 138, 167, 223, 235]
		Sasatrombicula (Sasatrombicula) bureschi Kolebinova & Beron, 1965 [51, 52, 162, 167]
		Sasatrombicula (Sasatrombicula) hexasternalae (Vercammen-Grandjean, 1963) [51, 162, 167]
		Anoplura
		Polyplax serrata (Burmeister, 1839) ! [17, 177, 216]
		Cestoda
		Milina grisea van Beneden, 1873 ? [200]
		Diptera
		Nycteribia (Achrocholidia) vexata Westwood, 1835! [85]
		Nycteribia (Nycteribia) pedicularia Latreille, 1805 ? [200]
		Phthiridium biarticulatum Hermann, 1804 [115, 126?, 135?, 167, 237]
		Nematoda
		Trichosomum speciosum van Beneden, 1873 [200]
		See also [174]
		Siphonaptera
		Rhinolophopsylla unipectinata unipectinata (Taschenberg, 1880) [2, 86, 142, 167, 200, 228]
		Trematoda
		Lecithodendrium granulosum Looss, 1907? [200]
		[Paralecithodendrium chilostomum (Mehlis) (=Prosthodendrium (Prosthodendrium) chilostomum (Mehlis, 1831)?)] [200]
		See also [174]
		Lecithodendrium sp. [86]
		Mesotretes peregrinus (Braun, 1900) [68, 167, 188]
		Plagiorchis vespertilionis (O.F. Müller, 1784) [68, 86, 167, 200]
		Prosthodendrium (Prosthodendrium) chilostomum (Mehlis, 1831) [86, 167, 200, 235]
Rhinolophus mehelyi Matschie, 1901	1 recognised species	Diptera
		Nycteribia (Nycteribia) schmidlii Schiner, 1853 [147]
Rhinolophus sp.	5 species(4 recognised species and 1 invalid species)	Acari
	1 innominate species	Eyndhovenia euryalis (G. Canestrini, 1885) [51, 76]
		Ixodes (Eschatocephalus) vespertilionis C.L. Koch, 1844 [5, 58, 126, 197]
		[Liponyssus setosus (Kolenati)] [72]
		Neomyobia sp. [126]
		Nematoda
		Litomosa ottavianii Lagrange & Bettini, 1948 [19, 167]
		Siphonaptera
		Rhinolophopsylla unipectinata unipectinata (Taschenberg, 1880) [44]
Tadarida teniotis (Rafinesque, 1814)	1 recognised species	Siphonaptera
		Araeopsylla gestroi (Rothschild, 1906) [3, 37, 44, 85]

Table 2

List of bat parasites (Acari, Anoplura, Cestoda, Diptera, Hemiptera, Nematoda, Siphonaptera, Trematoda) and their hosts in France (including Corsica), based on the published literature, with reported synonyms. Authors are listed in the bibliography. See also the work entitled Les parasites métazoaires des Chiroptères de France (Acari, Anoplura, Cestoda, Diptera, Hemiptera, Nematoda, Siphonaptera, Trematoda) : contribution à un état des lieux bibliographique (1762-2018) et à l’établissement d’une liste nationale (2019). Invalid species are listed in brackets. Records marked with an exclamation mark (!) are invalid. Records marked with a question mark (?) are dubious. They may require further clarification.

Parasite species	Number of reported hosts	Bat species and citation
Acari
Acari, part 1/3: generally recognised taxa (n = 53), with their reported synonyms, and records without identification to species level (n = 6)
Acanthophthirius (Myotimyobia) pantopus (Poppe et Trouessart, 1895)	1	Barbastella barbastellus (Schreber, 1774) [102, 114, 167, 235, 249]
Acanthophthirius (Acanthophthirius) poppei (Trouessart, 1895)	1	Pipistrellus nathusii (Keyserling & Blasius, 1839) [52, 102, 114, 167, 249]
Alabidocarpus diceratops Lawrence, 1952	1	Rhinolophus ferrumequinum (Schreber, 1774) [5, 110, 167]
Alabidocarpus megalonyx (Trouessart, 1895)	1	Rhinolophus ferrumequinum (Schreber, 1774) [52, 111, 167, 208, 218, 249]
Alabidocarpus minor (Rollinat & Trouessart, 1897)	1	Rhinolophus ferrumequinum (Schreber, 1774) [52, 111, 167, 208, 218, 249]
Argas pipistrellae (Audouin, 1832) = Argas vespertilionis (Latreille, 1796)
			Argas transgariepinus White, 1846 = Argas (Secretargas) transgariepinus White, 1846
Argas vespertilionis (Latreille, 1796)	11	Barbastella barbastellus (Schreber, 1774) [28]
		Eptesicus serotinus (Schreber, 1774) [28, 38]
		Miniopterus schreibersii (Natterer in Kuhl, 1817)? [85]
		Myotis emarginatus (Geoffroy, 1806) [28]
		Myotis myotis (Borkhausen, 1797) [28]
		Myotis mystacinus (Leisler in Kuhl, 1817) [28]
		Myotis sp. [171?, 242]
		Pipistrellus kuhlii (Kuhl, 1817) [28, 166]
		Pipistrellus nathusii (Keyserling & Blasius, 1839) [28]
		Pipistrellus pipistrellus (Schreber, 1774) [18, 28, 38, 67, 166]
		Plecotus auritus (Linné, 1758) [28]
Argas (Carios) vespertilionis (Latreille, 1802)	12	Eptesicus serotinus (Schreber, 1774) [31, 45, 86]
		Hypsugo savii (Bonaparte, 1837) [45]
		Myotis dasycneme (Boie, 1825) [31]
		Myotis mystacinus (Leisler in Kuhl, 1817) [45]
		Nyctalus leisleri (Kuhl, 1817) [45]
		Pipistrellus kuhlii (Kuhl, 1817) [45]
		Pipistrellus nathusii (Keyserling & Blasius, 1839) [45]
		Pipistrellus pipistrellus (Schreber, 1774) [31, 45, 177, 239]
		Pipistrellus sp. [45]
		Plecotus auritus (Linné, 1758) [45]
		Plecotus austriacus Fischer, 1829 [31]
		Rhinolophus ferrumequinum (Schreber, 1774) [45]
		See also [5, 167–169]
Argas (Secretargas) transgariepinus White, 1846	5	Chiroptera gen. sp. [189]
		Eptesicus serotinus (Schreber, 1774) [45, 189]
		Hypsugo savii (Bonaparte, 1837) [45, 189]
		Myotis emarginatus (Geoffroy, 1806) [45, 189]
		Plecotus austriacus Fischer, 1829 [45, 189]
		See also [31, 167]
Chiroptella muscae (Oudemans, 1906) = Oudemansidium musca (Oudemans, 1906)
			Calcarmyobia dusbabeki Uchikawa, 1985?	1?	Miniopterus schreibersii (Natterer in Kuhl, 1817) [50, 51, 167]
There is only one case. According to Lanza (1999) [publication n°167], this field data published by Beron (1971) [publication n°50] is doubtful. It could be Calcarmyobia dusbabeki Uchikawa, 1985 / Calcarmyobia rhinolophia (Radford, 1940).
Calcarmyobia rhinolophia (Radford, 1940)!	1	Miniopterus schreibersii (Natterer in Kuhl, 1817) [50, 51, 167]
		There is only one case. According to Lanza (1999) [publication n°167], this field data published by Beron (1971) [publication n°50] is doubtful. It could be Calcarmyobia dusbabeki Uchikawa, 1985 / Calcarmyobia rhinolophia (Radford, 1940).
Dermanyssus coriaceus = Hirstionyssus arcuatus (Koch, 1839). See [11, 13, 221, 222]
Dermanyssus sp.	1	Myotis emarginatus (Geoffroy, 1806) [86]
Dermanyssus pipistrellae (Gervais, 1841) = Hirstionyssus arcuatus (Koch, 1839)
Diplostaspis daubentonii Kolenati, 1857 = Spinturnix andegavinus (Deunff, 1977) according to Deunff (1977) [76]
Diplostaspis stellata Kolenati, 1859 = Spinturnix andegavinus (Deunff, 1977) according to Deunff (1977) [76]
Eschatocephalus flavipes (Koch) = Ixodes (Eschatocephalus) vespertilionis C.L. Koch, 1844
			Eschatocephalus vespertilionis C.L. Koch = Ixodes (Eschatocephalus) vespertilionis C.L. Koch, 1844
Eschatocephalus vespertilionis (Koch 1844) = Ixodes (Eschatocephalus) vespertilionis C.L. Koch, 1844
			Eyndhovenia euryalis (G. Canestrini, 1885)	5	Miniopterus schreibersii (Natterer in Kuhl, 1817) [76]
Myotis emarginatus (Geoffroy, 1806) [63, 121]
Rhinolophus euryale Blasius, 1853 [50, 63, 70, 76, 78, 105, 139, 167, 235]
Rhinolophus ferrumequinum (Schreber, 1774) [50, 63, 76, 78, 167]
Rhinolophus sp. [51, 76]
Eyndhovenia euryalis oudemansi (Eyndhoven, 1941)	1	Rhinolophus ferrumequinum (Schreber, 1774) [18?, 105, 252]
Haemalastor vespertilionis = Ixodes (Eschatocephalus) vespertilionis C.L. Koch, 1844
Hirstionyssus arcuatus (Koch, 1839)	1	Nyctalus noctula (Schreber, 1774) [11, 12, 103, 124, 127, 167, 177, 212, 221, 258, 259]
Ichoronyssus diversipilis Vitzthum, 1920	1	Miniopterus schreibersii (Natterer in Kuhl, 1817) [24]
Ichoronyssus scutatus (Kolenati, 1856)	1	Rhinolophus euryale Blasius, 1853 [50, 65, 167, 205, 212, 235, 246]
Ichoronyssus spinosus (Oudemans) 1902)	1	Miniopterus schreibersii (Natterer in Kuhl, 1817) [24, 70, 72]
Ichoronyssus spinosus (Oudemans, 1902) / Ichoronyssus scutatus (Kolenati, 1856)	1	Myotis myotis (Borkhausen, 1797)? [24, 70, 72]
Ichoronyssus sp.	1	Myotis myotis (Borkhausen, 1797)? [25]
Ixodes chiropterum Babos et Janisch, 1958 = Ixodes (Eschatocephalus) simplex simplex Neumann, 1906
Ixodes gracilipes = Ixodes (Eschatocephalus) vespertilionis C.L. Koch, 1844 ?
Ixodes hexagonus Leach, 1815 = Pholeoixodes hexagonus (Leach, 1815)
Ixodes longipes (Lucas, 1872) = Ixodes (Eschatocephalus) vespertilionis C.L. Koch, 1844
Ixodes (Pomerantzevella) simplex Neumann 1906 = Ixodes (Eschatocephalus) simplex simplex Neumann, 1906
Ixodes pospelovae Emtchuck, 1955 = Ixodes (Eschatocephalus) simplex simplex Neumann, 1906
Ixodes siculifer Mégnin, 1880 = Ixodes (Eschatocephalus) vespertilionis C.L. Koch, 1844
Ixodes reduvius Geer = Ixodes (Ixodes) ricinus (Linnaeus, 1758)
Ixodes (Eschatocephalus) vespertilionis C.L. Koch, 1844	18	Barbastella barbastellus (Schreber, 1774) [28, 32, 200]
		Chiroptera gen. sp. [45, 69, 160, 177, 197–199, 248]
		Eptesicus serotinus (Schreber, 1774) [28, 32, 166]
		Miniopterus schreibersii (Natterer in Kuhl, 1817) ([6, 28, 176!] see Beaucournu, 1966, p. 498; paper n°30 [200, 242])
		Myotis bechsteinii (Leisler in Kuhl, 1817) [28, 32, 38]
		Myotis daubentonii (Leisler in Kuhl, 1817) [28, 32]
		Myotis emarginatus (Geoffroy, 1806) [28, 32, 38, 136, 137, 177]
		Myotis myotis (Borkhausen, 1797) [23, 28, 32, 38, 154, 166]
		Myotis mystacinus (Leisler in Kuhl, 1817) [28, 32, 38, 85]
		Myotis nattereri (Kuhl, 1817) [28, 32]
		Pipistrellus kuhlii (Kuhl, 1817) [28]
		Pipistrellus nathusii (Keyserling & Blasius, 1839) [28]
		Pipistrellus pipistrellus (Schreber, 1774) [28, 86, 199]
		Plecotus auritus (Linné, 1758) [28, 199]
		Rhinolophus euryale Blasius, 1853 [28, 32, 85, 166]
		Rhinolophus ferrumequinum (Schreber, 1774) [28, 32, 38, 45, 51, 58, 154, 177, 199, 200, 214, 215, 218, 229, 239]
		Rhinolophus hipposideros (Bechstein, 1800) [23, 28, 32, 152, 154, 199, 200]
		Rhinolophus sp. [5, 58, 126, 197]
		See also [14, 31, 59, 143, 167, 201, 213]
Ixodes (Eschatocephalus) simplex simplex Neumann, 1906	4	Absence in Myotis blythii (Tomes, 1857) and M. blythii oxygnathus Monticelli, 1885 [32]
		Absence in Pipistrellus kuhlii (Kuhl, 1817) [32]
		Absence in Pipistrellus nathusii (Keyserling & Blasius, 1839) [32]
		Absence in Pipistrellus pipistrellus (Schreber, 1774) [32]
		Absence in Plecotus auritus (Linné, 1758) [32]
		Absence in Plecotus austriacus Fischer, 1829 [32]
		Rhinolophus ferrumequinum (Schreber, 1774) ! [85]
		Myotis mystacinus (Leisler in Kuhl, 1817) ! [85]
		Myotis myotis (Borkhausen, 1797) ! [85]
		Miniopterus schreibersii (Natterer in Kuhl, 1817) ([15, 31, 32, 45, 51, 143?, 166, 167, 176] see Beaucournu [1966; p. 498] paper n°30)
		See also [213]
Ixodes sp.	2	Myotis myotis (Borkhausen, 1797) [178]
		Rhinolophus hipposideros (Bechstein, 1800) [178]
Labidocarpus megalonyx (Trouessart, 1895) = Alabidocarpus megalonyx (Trouessart, 1895)
			Labidocarpus minor (Rollinat & Trouessart, 1897) = Alabidocarpus minor (Rollinat & Trouessart, 1897)
Labidocarpus (Labidocarpus) rollinati Trouessart, 1895	2	Rhinolophus ferrumequinum (Schreber, 1774) [167, 218, 232]
		Rhinolophus hipposideros (Bechstein, 1800) [167, 218] see also [52, 191]
Leiognathus arcuatus = Hirstionyssus arcuatus (Koch, 1839)
			Leiognathus uncinatus = Macronyssus uncinatus (Canestrini, 1885)
Leptotrombidium (Leptotrombidium) russicum (Oudemans, 1902) = Leptotrombidium russicum (Oudemans, 1902)
			Leptotrombidium russicum (Oudemans, 1902)	2	Myotis daubentonii (Leisler in Kuhl, 1817) [122, 167, 177, 256] see also [10]
Rhinolophus hipposideros (Bechstein, 1800) [161]
Leptus autumnalis = Neotrombicula autumnalis (Shaw, 1790)
			Liponyssus ellipticus = Macronyssus ellipticus (Kolenati, 1856)
Liponyssus euryale Canestrini = Macronyssus rhinolophi (Oudemans, 1902)
			Liponyssus longimanus = Macronyssus longimanus (Kolenati)
Liponyssus spinosus Oudemans, 1902 = Ichoronyssus scutatus (Kolenati, 1856)
Macronyssus ellipticus (Kolenati, 1856) !	1	Rhinolophus ferrumequinum (Schreber, 1774) ! [152, 177, 212]
According to Radovsky [212], this observation published by Husson & Daum [152] is dubious.
Macronyssus granulosus (Kolenati, 1856)	2	Miniopterus schreibersii (Natterer in Kuhl, 1817) [50, 167]
		Rhinolophus euryale Blasius, 1853 [51, 167]
Macronyssus longimanus (Kolenati) !	1	Rhinolophus ferrumequinum (Schreber, 1774) ! [152, 177, 212]
According to Radovsky [212], this observation published by Husson & Daum [152] is dubious.
Macronyssus uncinatus (Canestrini, 1885)	1	Chiroptera gen. sp. [160, 177]
Macronyssus rhinolophi (Oudemans, 1902)	3	Myotis myotis (Borkhausen, 1797)? [152?, 212?]
		Rhinolophus ferrumequinum (Schreber, 1774)? [152?, 212?]
		Rhinolophus hipposideros (Bechstein, 1800) [50, 152?, 167, 212?]
Macronyssus spinosus (Oudemans, 1902) = Ichoronyssus scutatus (Kolenati, 1856)
			Myobia pantopus = Acanthophthirius (Myotimyobia) pantopus (Poppe et Trouessart, 1895)
Myobia poppei = Acanthophthirius (Acanthophthirius) poppei (Trouessart, 1895)
			Neomyobia chiropteralis chiropteralis (Michael, 1884)	1	Rhinolophus hipposideros (Bechstein, 1800) [50, 52, 100, 167], see also [114]
See also [112]
Neomyobia rollinati (Poppe, 1908)	1	Rhinolophus ferrumequinum (Schreber, 1774) [50, 52, 167]
Neomyobia slovenica Dusbábek, 1969	1	Rhinolophus euryale Blasius, 1853 [50–52, 167]
Neomyobia sp.	1	Rhinolophus sp. [126]
Neotrombicula autumnalis (Shaw, 1790)	2	Chiroptera gen. sp. [62]
		Myotis daubentonii (Leisler in Kuhl, 1817) [8–10, 122, 167, 177, 256]
Neotrombicula racovitzai Feider, 1970	1	Eptesicus sp. [5, 118, 167]
Neotrombicula vandeli Kolebinova & Vercammen-Grandjean, 1971	1	Rhinolophus euryale Blasius, 1853 [7, 162, 167]
Notoedres (Notoedres) chiropteralis (Trouessart, 1896)	2	Rhinolophus ferrumequinum (Schreber, 1774) [106, 109, 167, 218, 250]
		Eptesicus serotinus (Schreber, 1774) [48, 106, 167, 250]
Nycteridocoptes eyndhoveni Fain, 1959	1	Rhinolophus ferrumequinum (Schreber, 1774) [49, 107]
Nycteridocoptes poppei Oudemans, 1898	3	Myotis blythii (Tomes, 1857) and M. blythii oxygnathus Monticelli, 1885 [113]
		Myotis daubentonii (Leisler in Kuhl, 1817) [113]
		Myotis myotis (Borkhausen, 1797) [24]
Oudemansidium musca (Oudemans, 1906)	1	Myotis daubentonii (Leisler in Kuhl, 1817) [122, 167, 177]
Paraperiglischrus rhinolophinus (C.L. Koch) = Paraperiglischrus rhinolophinus (C.L. Koch, 1841)
			Paraperiglischrus rhinolophinus Koch, 1841 = Paraperiglischrus rhinolophinus (C.L. Koch, 1841)
Paraperiglischrus rhinolophinus (C.L. Koch, 1841)	3	Rhinolophus euryale Blasius, 1853 [50, 76, 138, 167, 235]
		Rhinolophus ferrumequinum (Schreber, 1774) [50, 78, 167, 235]
		Rhinolophus hipposideros (Bechstein, 1800) [50, 138, 167, 223?, 235]
Periglischrus interruptus (Kolenati, 1856) = Paraperiglischrus rhinolophinus (C.L. Koch, 1841)
Pteracarus pipistrellius maximis Uchikawa, 1989	1	Nyctalus lasiopterus (Schreber, 1780) [251]
Prosopodectes chiropteralis (Trouessart, 1896) = Notoedres (Notoedres) chiropteralis (Trouessart, 1896)
Psorergates rhinolophi Fain, 1959	1	Rhinolophus ferrumequinum (Schreber, 1774) [108, 167]
Riedlinia (Riedlinia) petarberoni (Kolebinova & Vercammen-Grandjean, 1970)	1	Rhinolophus euryale Blasius, 1853 [7, 51, 161, 167]
Sarconissus vespertilionis = Ixodes (Eschatocephalus) vespertilionis C.L. Koch, 1844
			Sarcoptes chiropteralis Trouessart, 1896 = Notoedres (Notoedres) chiropteralis (Trouessart, 1896)
Sasatrombicula (Sasatrombicula) bureschi Kolebinova & Beron, 1965	1	Rhinolophus hipposideros (Bechstein, 1800) [51, 52, 162, 167]
Sasatrombicula (Sasatrombicula) hexasternalae (Vercammen-Grandjean, 1963)	4	Chiroptera gen. sp. [7]
		Rhinolophus euryale Blasius, 1853 [51, 162, 167]
		Rhinolophus ferrumequinum (Schreber, 1774) [51, 162, 167]
		Rhinolophus hipposideros (Bechstein, 1800) [51, 162, 167]
Spinturnix acuminatus (C.L. Koch, 1836)	1	Nyctalus noctula (Schreber, 1774) [recorded at Col de Bretolet, at the border between France and Switzerland] [79]
Spinturnix andegavinus (Deunff, 1977)	1	Myotis daubentonii (Leisler in Kuhl, 1817) [76], see also [167]
This species is treated as Spinturnix andegavina Deunff, 1977 by some authors [167, 209, 225]. According to Sachanowicz et al. (publication n°223; p. 49), this species “is a member of the myoti species group, which is actually recognised as also containing”: S. dasycnemi (Kolenati, 1856), S. myoti (Kolenati, 1856), S. bechsteini Deunff et al. 2004, S. emarginata (Kolenati, 1856), and S. mystacina (Kolenati, 1857)”. According to Lanza [167], this taxon is actually a synonym of Spinturnix daubentonii (Kolenati, 1857).
Spinturnix bechsteini (Deunff, Walter, Bellido et Volleth, 2004)	1	Myotis bechsteinii (Leisler in Kuhl, 1817) [82, 130]
Spinturnix daubentonii (Kolenati, 1857). This species is treated as a synonym of Spinturnix andegavinus (Deunff, 1977) by some authors [167, 209, 225].
			Spinturnix helvetiae (Deunff, Aellen & Keller, 1986).	1	Nyctalus noctula (Schreber, 1774) [recorded at Col de Bretolet, at the border between France and Switzerland] [79]
In the opinion of Uchikawa et al. (1994), this taxon is actually a subspecies of S. acuminata (C.L. Koch, 1836). See the article entitled “Contribution to the taxonomy of the genus Spinturnix (Acari: Spinturnicidae), with the erection of a new genus, Emballonuria” (Folia Parasitologica, 41 (4), p. 295).
Spinturnix emarginatus (Kolenati, 1856)	1	Myotis emarginatus (Geoffroy, 1806) [7, 76, 78]
		See also [77]
Spinturnix euryalis (G. Canestrini) = Eyndhovenia euryalis (G. Canestrini, 1885)
Spinturnix murinus (Walckenaer, 1847) = Spinturnix myoti (Kolenati, 1856)
			Spinturnix myoti (Kolenati, 1856)	8	Chiroptera gen. sp. [7, 65?, 167, 246]
Miniopterus schreibersii (Natterer in Kuhl, 1817) [76]
Myotis blythii (Tomes, 1857) and M. blythii oxygnathus Monticelli, 1885 [14, 76]
Myotis capaccinii (Bonaparte, 1837) [76]
Myotis myotis (Borkhausen, 1797) [14?, 76, 78, 177, 226]
Myotis nattereri (Kuhl, 1817) [63, 121]
Myotis punicus Felten, Spitzenberger & Storch, 1977 [64]
Rhinolophus euryale Blasius, 1853 [76]
See also [77]
Spinturnix mystacinus (Kolenati, 1857)	1	Myotis mystacinus (Leisler in Kuhl, 1817) [76, 78]
		See also [77]
Spinturnix nobleti Deunff, Volleth, Keller et Aellen, 1990	1	Hypsugo savii (Bonaparte, 1837) [81, 167]
Spinturnix oudemansi van Eyndhoven, 1941 = Eyndhovenia euryalis (G. Canestrini, 1885)
See also Eyndhovenia euryalis oudemansi (Eyndhoven, 1941).
Spinturnix plecotina (C.L. Koch, 1839)	3	Chiroptera gen. sp. [7]
		Plecotus auritus (Linné, 1758) [63, 76, 167]
		Plecotus austriacus Fischer, 1829 [63, 76, 167]
Spinturnix plecotinus Koch, 1839 = Spinturnix plecotina (C.L. Koch, 1839)
			Spinturnix punctata (Sundevall, 1833)	1	Barbastella barbastellus (Schreber, 1774) [80, 167]
Spinturnix psi (Kolenati, 1856)	7	Chiroptera gen. sp. [7, 167, 210?, 223]
		Eptesicus serotinus (Schreber, 1774) [48]
		Miniopterus schreibersii (Natterer in Kuhl, 1817) [50, 63, 76, 139]
		Myotis capaccinii (Bonaparte, 1837) [63, 76, 121, 139]
		Myotis myotis (Borkhausen, 1797) [76]
		Rhinolophus euryale Blasius, 1853 [76]
		Rhinolophus ferrumequinum (Schreber, 1774) [139, 223?]
		See also [210]
Spinturnix sp.	1	Myotis myotis (Borkhausen, 1797) [133]
Steatonyssus sp.	1	Myotis myotis (Borkhausen, 1797) [152, 212]
Steatonyssus spinosus Willmann, 1936	1	Rhinolophus ferrumequinum (Schreber, 1774) [50, 51, 167]
Thrombicula autumnalis Shaw = Neotrombicula autumnalis (Shaw, 1790)
			Thrombicula russica Oudemans = Leptotrombidium russicum (Oudemans, 1902)
Thrombicula russicum = Leptotrombidium russicum (Oudemans, 1902)
			Trombicula muscae = Oudemansidium musca (Oudemans, 1906)
Trombicula russicum = Leptotrombidium russicum (Oudemans, 1902)

Results and discussion

Based on published data, eight groups of bat parasites reported from France have been identified (Fig. 2). The majority of the analysed papers (94%) were published between 1762 and 1999 (Fig. 1). All host-parasite associations are listed in Tables 1 and 2. What follows is an overview of all bat parasites, arranged by higher taxonomic group.

Figure 2

Overview of the 113 generally recognised parasite taxa that are mentioned in the analysed papers (n = 237) per host taxonomic group. Invalid species (n = 22 Acari and 3 Diptera) recorded in the literature, records reported from France without identification to species level (n = 6 Acari; 1 Cestoda; 2 Diptera; 1 Hemiptera; 2 Nematoda and 2 Trematoda) and species only noted as absent (n = 3 Acari and 1 Diptera) are not included here.

Phylum Arthropoda Latreille, 1829

Subphylum Chelicerata Heymons, 1901

Subclass Acari Leach, 1817

Most of the studied papers (n = 112) deal with 75 species (53 generally recognised species and 22 invalid species) of Acari of four groups: Ixodida, Mesostigmata, Trombidiformes (suborder Prostigmata), and Sarcoptiformes (Fig. 3). A total of 53 recognised species (including two subspecies) of mites and ticks were reported to be parasites of bats in France prior to 2018. Of these, two species have only been collected from the border between Switzerland and the French department of Haute-Savoie (Col de Bretolet): Spinturnix helvetiae and S. acuminatus.

Figure 3

Number of recognised species of Acari (n = 53), per host order (n = 4) and genus (n = 23). Invalid species (n = 22 Acari) recorded in the literature (n = 237 papers) and species only noted as absent (n = 3 Acari) are not included here.

These recognised species found in the literature (n = 53) reported from France belong to 23 genera: Acanthophthirius, Alabidocarpus, Argas, Calcarmyobia, Eyndhovenia, Hirstionyssus, Ichoronyssus, Ixodes, Labidocarpus, Leptotrombidium, Macronyssus, Neomyobia, Neotrombicula, Notoedres, Nycteridocoptes, Oudemansidium, Paraperiglischrus, Pteracarus, Psorergates, Riedlinia, Sasatrombicula, Steatonyssus and Spinturnix. Among this group, the most diverse genera reported in France are Spinturnix and Macronyssus (Fig. 3): these genera account for 30% of all documented Acari infections involving valid species.

Six more records reported from France without identification to species level were found. These comprise Dermanyssus sp. [86], Ichoronyssus sp. [25], Ixodes sp. [178], Neomyobia sp. [126], Steatonyssus sp. [152, 212] and Spinturnix sp. [133]. Three other species are noted as absent from bats in western France, namely Ixodes canisuga, I. ricinus, and Pholeoixodes hexagonus [28]. Finally, 22 invalid taxa reported from France were found in the analysed papers. Examples for this category are Dermanyssus murinus (Lucas, 1840) [101, 104, 167, 183, 195], D. vespertilionis Dugès, 1834 [94], Pteroptus vespertilionis, and Spinturnix vespertilionis (C.L. Koch)] [70].

A large host group comprising the following taxa was identified with Acari infections: Barbastella barbastellus, Eptesicus serotinus, Eptesicus sp., Hypsugo savii, Miniopterus schreibersii, Myotis bechsteinii, M. blythii, M. blythii oxygnathus, M. capaccinii, M. dasycneme, M. daubentonii, M. emarginatus, M. myotis, M. mystacinus, M. nattereri, M. punicus, Myotis sp., Nyctalus lasiopterus, N. leisleri, N. noctula, Pipistrellus kuhlii, P. nathusii, P. pipistrellus, Pipistrellus sp., Plecotus auritus, P. austriacus, Rhinolophus euryale, R. ferrumequinum, R. hipposideros and Rhinolophus sp. (see Tables 1 and 2). The oldest works dealing with Acari parasitising bats in France are Geoffroy’s Histoire abrégée des insectes [123] and Latreille’s Précis des caractères génériques des insectes, disposés dans un ordre naturel [168], dated 1762 and 1797. Geoffroy mentioned the tick (found on an unidentified bat) as Acarus fuscus ovatus, pedibulus pallidis, vespertilionis, a taxon treated as Caris vespertilionis by Lamarck in the Histoire naturelle des animaux sans vertèbres (1839). The book of Latreille contains an observation of Carios on “la Chauve-Souris noctule” (=Nyctalus noctula or Nyctalus sp.). This taxon is very likely Argas (Carios) vespertilionis (Latreille, 1802). Descriptions of some species and subspecies are based on type material from France (e.g. Spinturnix nobleti [81], S. bechsteini [82], Pteracarus pipistrellius maximis [251], Myobia poppei (= Acanthophthirius (Acanthophthirius) poppei) [249] and Spinturnix andegavinus [76]). The majority of published data on parasites of chiropteran populations in France deal with Arachnida and similar findings were noted by Lanza [167], Krištofík & Danko [165] and Frank et al. [121] in Italy, Slovakia and other European countries.

Subphylum Hexapoda Latreille, 1825

Suborder Anoplura Leach, 1815 (order Phthiraptera Haeckel, 1896)

Only one species has been reported as a bat parasite in France, i.e. Polyplax serrata. The scientist Paul Rémy (1894–1962) published in 1948 the only record; this was from R. hipposideros (Borkhausen, 1797) in north-eastern France at Trémont-sur-Saulx (Meuse area [177, 216]). The mentioned locality in the original paper, “Frémont-sur-Saulx” [216], contains a typographical error. These field data, dated 1925–1926, and published in the journal La Feuille des Naturalistes, are surprising and dubious [Beaucournu, in litt.]. In fact, this species is more likely to be an ectoparasite of mammals of the Rodentia order (e.g. Apodemus, Clethrionomys and Mus genera) and Eulipotyphla (Crocidura leucodon) [98, 234]. It should be noted, however, that Polyplax sp. was also reported on Rhinolophus mehelyi by Gadžiev et al. in 1990 in Eastern Europe. These are the only data on Polyplax sp. in Lanza’s analysis [167] and hence they may be unreliable. Rémy’s observation is not mentioned in the works of Durden & Musser [98], Ferris [119] and Hopkins [140].

Order Diptera Linnaeus, 1758

According to Szentiványi et al., 17 species of bat flies are currently known in Europe [237]. Thirteen species of bat flies have been reported from France and two more records without identification to species level have been found. These are Basilia (Basilia) italica, B. (Basilia) mediterranea, B. (Basilia) nana, 1954, B. (Basilia) nattereri, Brachytarsina flavipennis, Nycteribia (Achrocholidia) vexata, N. (Nycteribia) kolenatii, N. (Nycteribia) latreillii, N. (Nycteribia) pedicularia, N. (Nycteribia) schmidlii, Penicillidia (Neopenicillidia) conspicua, P. (Penicillidia) dufourii, Phthiridium biarticulatum, Nycteribia sp. and Nycteribia kolenatii/latreillii/pedicularia [1, 4, 12, 22–30, 34, 35, 38–41, 43, 45, 47, 53, 73, 74, 86, 91, 93, 115–117, 126, 135, 146–152, 154, 156, 167, 170, 172, 173, 175, 177, 179, 186, 187, 200, 203, 204, 206, 214, 215, 237, 240–242, 247, 248, 260]. Three invalid taxa reported from France were found in the analysed papers: Nycteribia eparticulata, N. vespertilionis Meig., and N. vespertilionis Latreille. Penicillidia (Penicillidia) monoceros is noted as absent in western France [28]. Another species, B. (Basilia) daganiae could be distributed in France [28, 34]. As far as hosts of the bat flies (Diptera: Nycteribiidae and Streblidae) are concerned, E. serotinus, H. savii, M. schreibersii, M. bechsteinii, M. blythii, M. capaccinii, M. daubentonii, M. emarginatus, M. myotis, M. mystacinus, M. nattereri, M. species, P. pipistrellus, Pipistrellus sp., P. auritus, R. euryale, R. ferrumequinum, R. hipposideros, and R. mehelyi have been recorded in the literature. Jean-Frédéric Hermann’s record of Phthiridium vespertilionis, dated 1804, is the oldest French record of a dipteran as a bat parasite [135]. The bibliographical survey of the published data (n = 111 papers) written by Szentiványi et al. has shown that ten bat fly species are known to be associated with bats in Albania, Romania, and Italy [237]. Europe’s most species-rich communities have been reported in Spain (11 species), Switzerland (11 species), Hungary (11 species) and France (13 species). As such, France has the most diverse community reported in the literature.

Order Hemiptera Linnaeus, 1758

The literature provides well-documented cases for Cimex dissimilis, C. lectularius, and C. pipistrelli on M. myotis, M. emarginatus, R. euryale, and R. ferrumequinum [28, 167, 177, 207, 236, 253, 254]. In addition to these species, records reported from France without identification to species level have been found (Cimex sp. on M. myotis, M. emarginatus, R. euryale, and R. ferrumequinum) [28, 177]. The first published data in the analysed literature are dated 1961 [28] when Beaucournu published his observations on Cimex sp. in the department of Maine-et-Loire. True bugs (Hemiptera: Cimicidae) using bats as hosts have not been well studied in France. Studies on bat guano deposits could provide data on the species distribution. For instance, this method provided new records of Cimex sp. and Cimex dissimilis (Horváth, 1910) in a roosting colony of M. myotis in June 2017 and June 2018 in north-eastern France [177] (about C. dissimilis in roosting colony of M. myotis, see also [121] and [224]).

Order Siphonaptera Latreille, 1825

Twelve species are known to be associated with bats in France. They belong to the family Ischnopsyllidae Wahlgren, 1907. The species are Araeopsylla gestroi, Ischnopsyllus (Hexactenopsylla) hexactenus, I. (Ischnopsyllus) elongatus, I. (Ischnopsyllus) intermedius, I. (Ischnopsyllus) octactenus, I. (Ischnopsyllus) simplex, I. (Ischnopsyllus) variabilis, Nycteridopsylla ancyluris ancyluris, N. eusarca, N. longiceps, N. pentactena, and Rhinolophopsylla unipectinata unipectinata. According to Beaucournu and Launay [37], the published records of Nycteridopsylla dictena are dubious [146, 227]. Bat fleas have been observed on 20 hosts in France [3, 14, 23, 29, 33, 36–38, 42–44, 46, 85, 86, 90, 95, 141, 142, 152, 156, 157, 167, 177, 200, 204, 219, 227, 228, 231, 242, 257]. These hosts are E. serotinus, M. schreibersii, M. blythii, M. capaccinii, M. daubentonii, M. emarginatus, M. myotis, M. mystacinus, M. nattereri, Myotis sp., N. leisleri, N. noctula, P. kuhlii, P. nathusii, P. pipistrellus, P. auritus, P. austriacus, R. euryale, R. ferrumequinum, R. hipposideros, Rhinolophus sp., and Tadarida teniotis. Studies on bat fleas have a long history in France and the book of Walckenaer on insects, entitled Faune parisienne, insectes ou Histoire abrégée des insectes des environs de Paris [257], may be among the earliest such works. According to the Inventaire National du Patrimoine Naturel, 91 autochthonous (indigenous) species of the Siphonaptera order have been reported in France. Species of the Ischnopsyllidae found in France represent almost 13% of fleas in the country [153].

Phylum Nematoda Diesing, 1861

In France, the helminth fauna of bats is varied, with more than 30 species. Nematodes recorded in bats in France are divided into three orders: Muspiceida Bain & Chabaud, 1959, Rhabditida Chitwood, 1933, and Strongylida Molin, 1861; and six families: Onchocercidae Leiper, 1911 (three species), Molineidae Skrjabin & Schulz, 1937 (four species), Rictulariidae Railliet, 1916 (one species), Muspiceidae Bain & Chabaud, 1959 (two species), Seuratidae Hall, 1916 (one species), and Strongylacanthidae (Yorke & Maplestone, 1926, subfamily) Chabaud, 1960 (one species). Thirteen species and two nematodes identified to genus-level have been reported: Litomosa dogieli, L. filaria, L. ottavianii, Molinostrongylus alatus, M. ornatus, M. panousei, M. tipula, Pterygodermatites (Neopaucipectines) bovieri, Riouxgolvania nyctali, R. rhinolophi, Seuratum mucronatum, Strongylacantha glycirrhyza, Rictularia sp., Riouxgolvania sp., and Trichosomum speciosum. These parasites were documented in the following bat species: M. schreibersii, M. blythii, M. emarginatus, M. myotis, Plecotus sp., Rhinolophus sp. and three species of the genus Rhinolophus (R. euryale, R. ferrumequinum, and R. hipposideros) [19–21, 54, 56, 57, 68, 75, 86, 97, 99, 167, 177, 194, 200, 211, 233, 235, 243, 245]. The original description of the rare nematode P. (Neopaucipectines) bovieri is based on material from M. myotis in France. To my knowledge, it is the first published observation in the country (dated September 1885) [56, 66, 230]. The original descriptions of Riouxgolvania nyctali and R. rhinolophi are based on material from M. blythii and R. euryale in the Netherlands and the French departments of Ariège and Pyrénées-Orientales [20, 21]. A re-description of Seuratum mucronatum was based on material from Plecotus auritus (dated 1950) in the French department of Indre-et-Loire [54]. As a comparison, according to Horvat et al. [144, 145], two Nematode species (associated with M. myotis and R. ferrumequinum) are known in Serbia to be associated with bats, whilst in Croatia, these authors noted three species.

Phylum Platyhelminthes Minot, 1876

Class Trematoda

Records of 15 recognised species of trematodes from bats have been found from over 17 published papers. They belong to the order Plagiorchiida La Rue, 1957 and are divided into three families: Lecithodendriidae Lühe, 1901 (13 species), Mesotretidae Poche, 1926 (one species), and Plagiorchiidae Lühe, 1901 (one species). These species are Allassogonoporus amphoraeformis, Lecithodendrium granulosum, L. linstowi, L. moedlingeri, Mesotretes peregrinus, Parabascus duboisi, P. lepidotus, P. semisquamosus, Plagiorchis vespertilionis, Prosthodendrium (Prosthodendrium) carolinum, P. (Prosthodendrium) hurkovaae, P. (Prosthodendrium) chilostomum, P. (Prosthodendrium) longiforme, P. parvouterus, and Pycnoporus heteroporus [1, 57, 60, 61, 68, 71, 86–89, 97, 163, 167, 177, 188, 200, 202, 243]. Two trematodes identified to genus-level have been reported (Lecithodendrium sp. and Prosthodendrium sp.) [86]. These parasites were documented in the following bat species: Eptesicus serotinus [68, 88, 167], Miniopterus schreibersii [68, 167, 188, 200, 235], Myotis daubentonii [89, 167], M. capaccinii [68, 167], M. emarginatus [68, 86, 167], M. myotis [57, 61, 68, 167], Pipistrellus kuhlii [86], P. pipistrellus [57, 60, 61, 68, 86, 97, 163, 167, 235], Plecotus auritus [86, 167], P. austriacus [68, 167], Rhinolophus euryale [68, 167], R. ferrumequinum [68, 86–88, 167, 188, 200, 243], and R. hipposideros [68, 86, 167, 188, 200, 235].

As a comparison, in the United Kingdom, Lord et al. [181, 182] noted four trematode species and one trematode identified to genus-level in bats. In Serbia, Horvat et al. [144, 145] noted a total of seven trematode taxa associated with bats, and only one species in Croatia. According to the Inventaire National du Patrimoine Naturel, 28 species of the family Lecithodendriidae and 27 species of the family Plagiorchiidae have been reported in France. Species of the Lecithodendriidae found in bats represent almost 46% of the family [153]. The observations on Mesotretes peregrinus (Braun, 1900), published by Combes & Clerc, Dubois and Matskási [68, 88, 188], are of particular interest since this is the only species of the family Mesotretidae reported in Europe [125, 153]. As regards Plagiorchis vespertilionis (O.F. Müller, 1784) [68, 86–89, 200, 235], it is worth noting that this species is the only member of the family Plagiorchiidae reported from bats in France. According to Lanza [167], 18 taxa of the Plagiorchiidae family have been reported worldwide. The Histoire naturelle des helminthes ou vers intestinaux (1845) by Félix Dujardin (1801–1860) appears to be the earliest French source mentioning bat-associated trematodes [93]. The first documented reports from France, P. (Prosthodendrium) chilostomum and P. heteroporus, were published in the Notices helminthologiques (deuxième série) by Raphaël Blanchard (1857–1919) [57].

Class Cestoda

Three recognised and one innominate species of cestodes have been reported in bats in France, which makes it the richest community reported in Europe. These species are Milina grisea, Vampirolepis acuta, and V. balsacii [68, 86, 158, 159, 167, 177, 180, 196, 200, 243, 255, 262]. They belong to one family: Hymenolepididae Ariola, 1899 (Cyclophyllidea van Beneden in Braun, 1900 order). In addition to these species, one cestode identified to genus-level has been reported (Vampirolepis sp.) [86]. The description of H. balsacii is based on material from Myotis bechsteinii and Eptesicus serotinus from north-eastern France collected by the naturalist Henri Heim de Balsac (1899–1979) at a place called Buré d’Orval in Allondrelle-la-Malmaison [158, 159]. Some authors have noted that the observation was made at “Buré” ([158] see also [142]). Joyeux & Baer’s paper on cestodes, entitled Sur quelques Cestodes de France, is the first work on the bat cestodes in France (published in 1934 in the journal Archives du Muséum national d’Histoire naturelle). Bat cestodes are not well studied, especially in France, where the most recent data from field research is almost 50 years old [68, 86, 158, 159, 167, 177, 180, 196, 200, 243, 255, 262]. France has the most diverse community reported in the literature. According to Frank et al. [121], three species are known to be associated with bats in Poland and Hungary. These authors also reported one species in Germany and two species in Austria. It is worth noting that, according to the Inventaire National du Patrimoine Naturel, 75 species of the family Hymenolepididae have been reported in France and species of the family found in bats represent 4% of the family [153].

Hosts and geographical distribution of bat-parasite associations

Over a 256-year period, the 113 recognised taxa of bat parasites from France were collected from 27 bats species and six other bats that were not identified to species-level (five genera and the Pipistrellus species complex) (Figs. 2 and 4). The taxa are B. barbastellus, E. serotinus, H. savii, M. schreibersii, M. bechsteinii, M. blythii, M. capaccinii, M. dasycneme, M. daubentonii, M. emarginatus, M. myotis, M. mystacinus, M. nattereri, M. punicus, N. lasiopterus, N. leisleri, N. noctula, P. kuhlii, P. nathusii, P. pipistrellus, P. auritus, P. austriacus, R. euryale, R. ferrumequinum, R. hipposideros, R. mehelyi, T. teniotis, Eptesicus sp., Myotis sp., Pipistrellus sp., Plecotus sp., Rhinolophus sp., and the species complex Pipistrellus pipistrellus/kuhlii/nathusii. These species represent almost 79% of the bat fauna of France (including Corsica). The most commonly reported hosts, which are mentioned in more than 29 papers, are E. serotinus, R. euryale, R. hipposideros, M. schreibersii, P. pipistrellus, M. myotis, and R. ferrumequinum. However, the most cited species is the Greater horseshoe bat (R. ferrumequinum); 30% of the analysed publications deal with this species. Some bat species have no records of associated metazoan parasites in the analysed publications (n = 237), because the ecology of the host is poorly studied (E. nilssonii, Vespertilio murinus (Particoloured Bat), M. alcathoe, M. escalerai and P. macrobullaris). In addition to this, since P. pygmaeus was identified in the 1990’s, we cannot rule out that some of the records of Pipistrellus sp. or P. pipistrellus may refer to P. pygmaeus [41, 45, 53, 58, 142].

Figure 4

Histogram showing the number of studies (n = 237) per host taxon (n = 34; species: 27; complex: 1; genera: 6) during the period 1762–2018.

Published field data originated from 72 French departments (Fig. 5). One of them is mentioned as a non-prospected area (department of Vosges). Indeed, there is no publication about bat parasites in this department. Associations with specified geographical locations were most commonly from Ardèche (11 papers), Ariège (13 papers), Bouches-du-Rhône (15 papers), Haute-Savoie (12 papers), Maine-et-Loire (19 papers), Moselle (11 papers), Meurthe-et-Moselle (23 papers), Pyrénées-Orientales (24 papers), Sarthe (12 papers), Haute-Corse, and Corse-du-Sud (23 papers). Importantly, these distribution patterns are influenced more by biased sampling efforts than by actual geographical and ecological patterns. This distribution map only helps to point to well-studied areas. The relative prominence of the departments Ardèche, Ariège, Pyrénées-Orientales, Haute-Corse and Corse-du-Sud compared to other departments is most likely a result of the attention given to karstic areas by European biospeleology (for instance [14, 23–25, 48–53, 69, 70, 115, 118, 120, 126, 131–133, 136, 137, 151, 152, 154, 178, 186, 214–216, 246]). This is closely linked to the success and prevalence of M. schreibersii, R. euryale, and R. ferrumequinum compared to other bat species. This prevalence could be the result of their ability to roost in the summer in limestone areas and underground sites. The focus on E. serotinus, R. ferrumequinum, R. hipposideros, P. pipistrellus, and M. myotis is the result of their ability to exploit anthropogenic environments (i.e. farmland, urban areas). As a consequence of this, these species have more contact with human populations (about P. pipistrellus, see [182]) and were the first to be studied in France (for the 1762–1844 period see [11, 13, 18, 90–92, 94–97, 124, 183, 220, 258, 260]).

Conflict of interest

The author declares no conflict of interest.