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Accepted by D. Whitmore: 19 Oct. 2016; published: 6 Dec. 2016
ZOOTAXA
ISSN 1175-5326 (print edition)
ISSN
1175-5334
(online edition)
Copyright © 2016 Magnolia Press
Zootaxa 4205 (2): 101
–
126
http://www.mapress.com/j/zt/
Article
101
http://doi.org/10.11646/zootaxa.4205.2.1
http://zoobank.org/urn:lsid:zoobank.org:pub:7FC9F0FE-4FB7-483B-ADF1-ECA34841E40B
Checklist of host associations of European bat flies
(Diptera: Nycteribiidae, Streblidae)
TAMARA SZENTIVÁNYI
1,2,3,7
, PÉTER ESTÓK
4,5
& MIHÁLY FÖLDVÁRI
6
1
University of Debrecen, Department of Evolutionary Zoology and Human Biology, Egyetem tér 1., H-4032, Debrecen, Hungary
2
University of Lausanne, Department of Ecology and Evolution, CH-1015, Lausanne, Switzerland
3
Museum of Zoology, Palais de Rumine, Place de la Riponne 6, CH-1014, Lausanne, Switzerland
4
Eszterházy Károly University, Department of Zoology, Eszterházy tér 1., H-3300, Eger, Hungary
5
Bükk Mammalogical Society, Maklári u. 77/A, H-3300, Eger, Hungary
6
University of Debrecen, MTA–DE "Lendület" Behavioural Ecology Research Group, Department of Evolutionary Zoology and
Human Biology, Egyetem tér 1., H-4032, Debrecen, Hungary
7
Corresponding author. E-mail: tamaraszentivanyi@gmail.com
Abstract
Bat flies are obligate blood-sucking ectoparasites of bats. They are divided into two families: Nycteribiidae and Streblidae.
Europe has 17 species of bat flies and 45 species of bats (Mammalia: Chiroptera). This checklist is based on both published
records and our own field data and provides updated information on all associations between bat flies and their hosts in
Europe. The host-parasite association between Basilia italica Theodor and Plecotus auritus (Linnaeus) is reported for the
first time. Moreover, our records of B. italica on Myotis alcathoe Helversen & Heller, B. nana Theodor & Moscona on
Plecotus auritus, Nycteribia kolenatii Theodor & Moscona on M. bechsteinii (Kuhl) and Penicillidia dufourii (Westwood)
on M. daubentonii (Kuhl) represent new host associations for Hungary.
Key words: ectoparasites, host associations, parasite specificity, Pupipara
Introduction
Bats (Mammalia: Chiroptera) possess a diverse ectoparasitic fauna including Diptera, Heteroptera, Siphonaptera
and Acari (e.g., Hutson 1972; Marshall 1982). Dipteran parasites are represented by two families: Nycteribiidae
and Streblidae. To the best of our knowledge, 16 nycteribiids and one streblid have been reported for Europe (Pape
et al. 2015).
Nycteribiids and streblids are obligate blood-sucking ectoparasites of bats, and they belong together with
Glossinidae (tsetse) and Hippoboscidae (louse flies) to the calyptrate superfamily Hippoboscoidea. Louse flies and
bat flies form the Pupipara, the species of which reproduce by obligate pseudo-placental unilarviparity (Meier et al.
1999). The female deposits a mature third instar larva (usually on the wall of the host roost), which immediately
begins to pupate (Hill 1963). Nycteribiids have their wings totally reduced, while most streblids have fully
developed wings at least during a certain part of their adult life. Bat flies do not leave their hosts except for the
laying of larvae (both families) or when their hosts are disturbed (only streblids). There are three subfamilies within
the family Nycteribiidae: Archinycteribiinae, Cyclopodiinae and Nycteribiinae (Maa 1965). The family Streblidae
is divided into five subfamilies: the Old World Nycteriboscinae and Ascodipterinae and the New World
Trichobiinae, Nycterophiliinae and Streblinae (Dick & Patterson 2006). The only European streblid species is
Brachytarsina flavipennis Macquart of the Nycteriboscinae subfamily (as Brachytarsininae in Dittmar et al. 2006;
Vasconcelos et al. 2015).
The number of European bat fly species is probably underestimated and is likely to increase as a result of more
research (Pape et al. 2015). This is expected especially on the least known and rare hosts such as, e.g., Nyctalus
azoreum (Thomas), N. lasiopterus (Schreber) and Pipistrellus maderensis (Dobson) (all Chiroptera:
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Vespertilionidae). No recent review of known host associations of bat flies is available, the latest dating back to the
1960s (e.g., Hůrka 1964; Theodor 1967). Our study aims to fill this gap by providing a review of the existing
literature, complemented with unpublished data from fieldwork in Hungary. Moreover, since bat flies could be
vectors of several zoonotic agents such as Polychromophilus Dionisi (Haemosporida: Plasmodiidae) (Gardner &
Molyneux 1988) or BartonellaStrong et al. (Rhizobiales: Bartonellaceae) species (Morse et al. 2012), we urge that
future studies address the ecology and disease transmitting potential of these flies.
Material and methods
We collected literature data from the area of European territory defined by Pape et al. (2015) as “the western
Palaearctic, including the European mainland, Great Britain, the Macaronesian islands, Cyprus, Faroe Islands,
Iceland, Svalbard, Franz Josef Land and Novaya Zemlya, but excluding Turkey, the Caucasus, western
Kazakhstan, the Arabian Peninsula and North Africa” (Fig. 1), and supplemented this with data obtained from our
own fieldwork in Hungary (Fig. 2). Forty-five (45) bat species are reported from Europe (UNEP/Eurobats 2015).
Distributions of bat flies are given based on the Catalogue of Palaearctic Diptera ( Hůrka & Soós 1986a, 1986b), as
well as on more recent works (Tab. 1). Our review lists only bat fly species recorded from Europe, but a few extra-
limital records have been included (indicated in square brackets) for species associated with hosts occurring also in
Europe. In most cases, bat flies have numerous documented occurrences on different bat species, but usually they
have one or two main hosts: we indicated the main host-bat fly associations by putting the names in bold (Tabs 1–
2). We determined host specificity based on the frequency of host use by bat flies. Our own material was collected
in Hungary in 1998–1999 and 2012–2014 (Fig. 2), with full collecting data provided in Table 1. Bats were caught
using mist nets deployed at drinking, foraging or swarming sites. The flies were collected from the bats using
forceps and stored in 70% ethanol. Bat flies were identified using the works of Theodor & Moscona (1954) and
Theodor (1967), whereas their classification and nomenclature follow Hůrka & Soós (1986a, 1986b). For bat
classification, we followed Dietz & Von Helversen (2004). The collected bat fly specimens are deposited at
Eszterházy Károly College (Eger, Hungary). The map in Figure 1 was created with QGIS 2.8.2-Wien.
Results and discussion
All host-parasite associations are presented in Tables 1–2, and the collecting sites surveyed in 1998–1999 and
2012–2014 are summarized in Figure 2.
Many bat species have no records of associated bat flies, generally because the parasitology of the host has
been poorly studied (e.g., Eptesicus bottae (Peters),Myotis escalerai Cabrera, Plecotus kolombatovici Dulic [all
Chiroptera: Vespertilionidae]). During our survey we recorded 24 host associations of bat flies in Hungary,
including that of Basilia italica Theodor (Diptera: Nycteribiidae) on Plecotus auritus (Linnaeus) (Chiroptera:
Vespertilionidae), an association here reported for the first time.
We also provide the second documented cases for Basilia italica on Barbastella barbastellus (Schreber)
(Chiroptera: Vespertilionidae) and B. italica on Myotis alcathoe Helversen & Heller (Chiroptera: Vespertilionidae)
(Czuppon & Molnár 2001; Danko et al. 2010) and the third case for Basilia nattereri (Kolenati) (Diptera:
Nycteribiidae) on Myotis bechsteinii (Kuhl) (Chiroptera: Vespertilionidae) (Dudich 1925; Soós 1955).
In addition, our records of Basilia italica on Myotis alcathoe, B. nana Theodor & Moscona (Diptera:
Nycteribiidae) on Plecotus auritus, Nycteribia kolenatii Theodor & Moscona (Diptera: Nycteribiidae) on M.
bechsteinii and Penicillidia dufourii (Westwood) (Diptera: Nycteribiidae) on M. daubentonii (Kuhl) (Chiroptera:
Vespertilionidae) represent new host associations for Hungary.
We consider Falcoz’s (1926) records of Nycteribia pedicularia Latreille (Diptera: Nycteribiidae) as doubtful
since this species was not well defined and poorly distinguished from N. latreillii (Leach) and N. kolenatii at that
time (Theodor & Moscona 1954). We also indicate some additional early records as doubtful because of the
unclarified taxonomic status of the hosts (e.g., “Vespertilio murinus” in Westwood [1835] and Kolenati [1857]).
Likewise, Dudich’s (1925) records of Basilia nattereri are considered doubtful, as B. nana and B. nattereri were
first distinguished by Theodor (1954).
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EUROPEAN BAT FLY HOST ASSOCIATIONS
Our paper provides a complete checklist of bat flies and their bat hosts from Europe and also provides essential
knowledge for proposed future studies that aim towards a better understanding of the nature of the remarkable bat-
bat fly host-parasite system.
FIGURE 1. Study area (after Pape et al. 2015).
FIGURE 2. Map illustrating sites sampled for bat flies in Hungary in 1998–1999 and 2012–2014.
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TABLE 1. List of bat flies (Nycteribiidae and Streblidae) of Europe and their hosts, based on the literature and
previously unpublished data. Main hosts are in bold font. Records from outside Europe are listed in square brackets. Host
associations marked with an asterisk (*) are newly recorded for Hungary. Countries of bat fly species distributions are
listed in alphabetical order. For authors of bat species, see Table 2.
Bat fly species Host species, references and new data Distribution of bat fly
species
Nycteribiidae Samouelle
Subfamily Nycteribiinae Samouelle
Basilia daganiae Theodor &
Moscona
Pipistrellus kuhlii—Bequaert (1953), Theodor & Moscona (1954),
Theodor (1956, 1967), Kock (1974)
Cyprus
[Pipistrellus pipistrellus—Benda et al. (2010; Jordan)]
Basilia italica Theodor Barbastella barbastellus—Czuppon & Molnár (2001). Present
study: 1 ♂, Hungary, [Heves county], Felsőtárkány, 22.VIII.2013, P.
Estók leg.
Albania, France,
Hungary, Italy, Poland,
Slovakia, Switzerland
Eptesicus serotinus—Czuppon & Molnár (2001)
Myotis alcathoe*—Danko et al. (2010). Present study: 1 ♀,
Hungary, [Heves county], Noszvaj, 04.VI.2012, P. Estók leg.; 1 ♀
and 1 ♂, Hungary, [Heves county], Felsőtárkány, 24.VII.2012, P.
Estók leg.; 1 ♀, Hungary, [Heves county], Egerbakta, 21.VIII.2012,
P. Estók leg.; 1 ♀, Hungary, [Borsod-Abaúj-Zemplén county],
Miskolc, 29.VIII.2012, P. Estók leg.; 1 ♀, Hungary, [Heves county],
Egerbocs, 10.VIII.2014, P. Estók leg.; 2 ♀, Hungary, [Heves
county], Felsőtárkány, 23.VIII.2014, P. Estók leg.
Myotis brandtii—Hůrka (1964, as “M. mystacinus”), Hůrka (1980,
1984), Czuppon (2001), Czuppon & Molnár (2001). Present study:
1 ♂, Hungary, [Heves/Nógrád county] Mátra Mts., 05.VI.1999, P.
Paulovics leg.; 1 ♀ and 1 ♂, Hungary, [Heves county],
Felsőtárkány, 24.VII.2012, P. Estók leg.; 2 ♂, Hungary, [Heves
county], Noszvaj, 17.VIII.2012, P. Estók leg.
Myotis emarginatus—Theodor (1954), Hůrka (1980)
Myotis myotis—Theodor (1954), Hutson (1984)
Myotis mystacinus—Aellen (1963), Nowosad (1990), Beaucournu
& Noblet (1996), Czuppon (2001), Czuppon & Molnár (2001),
Krištofík & Danko (2012), Szentiványi et al. (2016)
Myotis nattereri—Czuppon (2001), Czuppon & Molnár (2001)
Plecotus auritus*—Present study: 1 ♂, Hungary, [Heves county],
Nagyvisnyó, 10.VIII.2012, P. Estók leg.
Basilia mediterranea Hůrka Eptesicus serotinus—Kock & Quetglas (2003, as “Eptesicus
serotinus boscai”)
France (incl. Corsica),
Spain (incl. Mallorca)
Hypsugo savii—Beaucournu & Noblet (1994)
Pipistrellus pygmaeus—Bauer (1957, as “P. pipistrellus
mediterraneus”)
Pipistrellus pipistrellus—Beaucournu & Noblet (1985, 1994)
......continued on the next page
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EUROPEAN BAT FLY HOST ASSOCIATIONS
TABLE 1. (Continued)
Bat fly species Host species, references and new data Distribution of bat fly
species
Basilia mongolensis nudior
Hůrka
Myotis alcathoe—Heddergott (2009) Austria, Bulgaria,
Croatia, Germany,
Greece (incl. Crete)
Myotis aurascens—Ševčík et al. (2013)
Plecotus austriacus—Hůrka (1972)
ex. Myotis mystacinus—Hůrka (1984), Kock (1984)
Basilia nana Theodor &
Moscona
Barbastella barbastellus—Grulich & Povolný (1955) Albania, Austria,
Bulgaria, Czech
Republic, France,
Germany, Great Britain,
Hungary, Poland,
Romania, Slovakia,
Spain, Sweden,
Switzerland, The
Netherlands
Miniopterus schreibersii—Aellen (1963)
Myotis bechsteinii—Theodor (1954), Theodor & Moscona (1954),
Grulich & Povolný (1955), Aellen (1963), Hůrka (1964), Kock
(1973), Hůrka (1984), Hutson (1984), Nowosad (1987, 1990),
Beaucournu & Noblet (1996), Kock (1999) Rupp (1999), Czuppon
(2001), Czuppon & Molnár (2001), Rupp et al. (2004), Reckardt &
Kerth (2006), Scheffler (2010), Scheffler & Hiller (2010), Baagøe
(2011b), Krištofík & Danko (2012), Szentiványi et al. (2016).
Present study: 3 ♀ and 2 ♂, Hungary, [Veszprém county], Eplény,
21.VII.1999, P. Paulovics leg.; 1 ♀ and 3 ♂, Hungary, [Veszprém
county], Bakonybél, 25.VIII.1999, P. Paulovics leg.; 4 ♀ and 1 ♂,
Hungary, [Veszprém county], Bakonybél, 25.VIII.1999, P.
Paulovics leg.; 2 ♀, Hungary, [Veszprém county], Bakony Mts.,
27.VIII.1999, P. Paulovics leg.; 2 ♀, Hungary, [Heves county],
Noszvaj, 18.VIII.2012, P. Estók leg.; 3 ♀, Hungary, [Borsod-Abaúj-
Zemplén county], Miskolc, 29.VIII.2012, P. Estók leg.; 1 ♀,
Hungary, [Heves county], Noszvaj, 05.IX.2012, P. Estók leg.; 2 ♂,
Hungary, [Borsod-Abaúj-Zemplén county], Miskolc, 06.IX.2012, P.
Estók leg.; 1 ♀, Hungary, [Borsod-Abaúj-Zemplén county],
Miskolc, 10.IX.2012, P. Estók leg.; 3 ♀ and 4 ♂, Hungary, [Borsod-
Abaúj-Zemplén county], Miskolc, 14.IX.2012, P. Estók leg.; 1 ♂,
Hungary, [Borsod-Abaúj-Zemplén county], Miskolc, 17.IX.2012, P.
Estók leg.; 3 ♀ and 4 ♂, Hungary, [Borsod-Abaúj-Zemplén county],
Miskolc, 27.IX.2012, P. Estók leg.; 1 ♀, Hungary, [Borsod-Abaúj-
Zemplén county], Miskolc, 04.IX.2013, P. Estók leg.; 1 ♀, Hungary,
[Borsod-Abaúj-Zemplén county], Varbó, 10.IX.2013, P. Estók leg.;
2 ♀ and 1 ♂, Hungary, [Borsod-Abaúj-Zemplén county], Miskolc,
27.IX.2013, P. Estók leg.; 2 ♂, Hungary, [Borsod-Abaúj-Zemplén
county], Miskolc, 29.IX.2013, P. Estók leg.
Myotis blythii—Hůrka (1964, 1980), Czuppon & Molnár (2001)
Myotis dasycneme—Theodor (1954), Theodor & Moscona (1954),
Theodor (1967), Hutson (1984), Nowosad (1987)
Myotis daubentonii—Haitlinger (1978), Czuppon (2001), Czuppon
& Molnár (2001). Present study: 1 ♀, Hungary, [Heves county],
Felsőtárkány, 18.V.2013, P. Estók leg.
Myotis emarginatus—Theodor & Moscona (1954), Theodor (1967),
Beaucournu & Noblet (1996)
Myotis myotis—Theodor & Moscona (1954), Grulich & Povolný
(1955), Aellen (1963), Hůrka (1964, 1980), Krištofík (1982), Hůrka
(1984), Hutson (1984), Czuppon (2001), Czuppon & Molnár
(2001), Güttinger et al. (2011)
Myotis mystacinus—Theodor & Moscona (1954), Theodor (1954,
1967), Kock (1973)
......continued on the next page
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TABLE 1. (Continued)
Bat fly species Host species, references and new data Distribution of bat fly
species
Myotis nattereri—Theodor & Moscona (1954), Grulich & Povolný
(1955), Hůrka (1980, 1984), Hutson (1984), [Amr & Qumsiyeh
(1993; Jordan)], Imaz et al. (1999), Czuppon (2001), Czuppon &
Molnár (2001), Topál (2011). Present study: 1 ♂, Hungary,
[Veszprém county], Bakony Mts., 27.VIII.1999, P. Paulovics leg.; 1
♂, Hungary, [Heves county], Felsőtárkány, 24.VII.2012, P. Estók
leg.
Pipistrellus pipistrellus—Hutson (1984)
Plecotus auritus*—Theodor (1954), Theodor & Moscona (1954),
Beaucournu (1961), Walter (2004). Present study: 1 ♂, Hungary,
[Veszprém county], Bakonybél, 25.VIII.1999, P. Paulovics leg.
Rhinolophus ferrumequinum—Hutson (1984)
Rhinolophus hipposideros—Theodor & Moscona (1954)
Basilia nattereri (Kolenati) Eptesicus serotinus—Kolenati (1857) (doubtful record), Theodor
(1954), Theodor & Moscona (1954), Hůrka (1980), Hutson (1984),
Baagøe (2011a)
Czech Republic, France,
Hungary, Romania,
Slovakia, Spain,
Switzerland, Ukraine
Myotis bechsteinii—Dudich (1925) (doubtful record), Soós (1955).
Present study: 1 ♂, Hungary, [Borsod-Abaúj-Zemplén county],
Miskolc, 21.IX.2014, P. Estók leg.
Myotis daubentonii—Soós (1955), Czuppon & Molnár (2001),
Krištofík & Danko (2012)
Myotis emarginatus—Dudich (1925)
Myotis myotis—Soós (1955), Hůrka (1980)
Myotis mystacinus—Aellen (1963), Hůrka (1980)
Myotis nattereri—Soós (1955), Aellen (1963), Hutson (1984),
Masson (1989), Imaz et al. (1999), Czuppon (2001), Czuppon &
Molnár (2001), Topál (2011), Krištofík & Danko (2012). Present
study: 1 ♂, Hungary, [Borsod-Abaúj-Zemplén county],
Bükkszentkereszt, 05.IX.2013, P. Estók leg.
Plecotus auritus—Balcells (1968b), Hůrka (1980)
Rhinolophus hipposideros—Hutson (1984)
Nycteribia kolenatii Theodor
& Moscona
Barbastella barbastellus—Haitlinger (1978), Nowosad (1987),
Scheffler (2011a)
Austria, Czech
Republic, Denmark,
Eptesicus serotinus—Theodor (1954), Theodor & Moscona (1954),
Hůrka (1964), Hutson (1984), Baagøe (2011a) Estonia, Finland,
France, Germany,
Myotis bechsteinii*—Aellen (1963), Hůrka (1964, 1984), Hutson
(1984), Nowosad (1987), Scheffler (2010), Baagøe (2011b). Present
study: 1 ♂, Hungary, [Borsod-Abaúj-Zemplén county], Miskolc,
27.IX.2012, P. Estók leg.
Great Britain,
Hungary, Ireland, Italy,
Macedonia, Norway,
Poland, Portugal,
Myotis brandtii—Rupp (1999), Rupp et al. (2004), Walter (2004)
Myotis dasycneme—Theodor (1954), Theodor & Moscona (1954),
Hůrka (1964), Hutson (1984), [Orlova (2011; Russia)]
......continued on the next page
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EUROPEAN BAT FLY HOST ASSOCIATIONS
TABLE 1. (Continued)
Bat fly species Host species, references and new data Distribution of bat fly
species
Myotis daubentonii—Theodor (1954), Theodor & Moscona (1954),
Grulich & Povolný (1955), Aellen (1963), Hůrka (1964)
Beaucournu (1972), Kock (1973), Nowosad (1974), Hůrka (1984),
Hutson (1984), Müller & Ohlendorf (1984), Nowosad (1987),
Walter (1987), Nowosad (1990), Haitlinger & Ruprecht (1992),
Dietz & Walter (1995), Beaucournu & Noblet (1996), Kock (1999),
Rupp (1999), Czuppon (2001), Heddergott & Claußen (2004), Rupp
et al. (2004), Scheffler & Ressler (2005, 2007), Scheffler (2008,
2009), Lindecke & Scheffler (2011), [Orlova (2011; Russia)],
Krištofík & Danko (2012). Present study: 15 ♀ and 23 ♂, Hungary,
[Baranya county], Abaliget, 04.IX.1998, P. Paulovics leg.; 1 ♀ and
1 ♂, Hungary, [Veszprém county], Kislőd, 19.VIII.1998, P.
Paulovics leg.; 4 ♀ and 9 ♂, Hungary, [Heves county], Vízfő,
05.IX.1998, P. Paulovics leg.; 20 ♀ and 30 ♂, Hungary, [Fejér
county], Alba, 28.VIII.1999, P. Paulovics leg.; 31 ♀ and 40 ♂,
Hungary, [Veszprém county], Bakony Mts., 1999, P. Paulovics leg.;
2 ♀ and 1 ♂, Hungary, [Veszprém county], Bakony Mts.,
27.VIII.1999, P. Paulovics leg.; 8 ♀ and 11 ♂, Hungary, [Veszprém
county], Eplény, 21.VII.1999, P. Paulovics leg.; 1 ♀ and 2 ♂,
Hungary, [Hajdú-Bihar county], Debrecen, 20.V.1999, P. Paulovics
leg.; 285 ♀ and 186 ♂, Hungary, [Tolna county], Gemenc, 1999, P.
Paulovics leg.; 2 ♀ and 6 ♂, Hungary, [Veszprém county], Kislőd,
16.VI.1999, P. Paulovics leg.; 2 ♀ and 1 ♂, Hungary, [Heves/
Nógrád county], Mátra Mts., 05.VI.1999, P. Paulovics leg.; 11 ♀
and 8 ♂, Hungary, [Veszprém county], Szentgál, 02.IV.1999, P.
Paulovics leg.; 1 ♀, Hungary, [Heves county], Noszvaj, 04.VI.2012,
P. Estók leg.; 1 ♀ and 1 ♂, Hungary, [Borsod-Abaúj-Zemplén
county], Miskolc, 10.VII.2012, P. Estók leg.; 1 ♂, Hungary, [Heves
county], Felsőtárkány, 17.VII.2012, P. Estók leg.; 1 ♂, Hungary,
[Borsod-Abaúj-Zemplén county], Miskolc, 23.VII.2012, P. Estók
leg.; 1 ♀, Hungary, [Heves county], Noszvaj, 16.VIII.2012, P. Estók
leg.; 1 ♀ and 2 ♂, Hungary, [Heves county], Noszvaj, 18.VIII.2012,
P. Estók leg.; 1 ♀ and 2 ♂, Hungary, [Borsod-Abaúj-Zemplén
county], Miskolc, 29.VIII.2012, P. Estók leg.; 1 ♀ and 3 ♂,
Hungary, [Borsod-Abaúj-Zemplén county], Miskolc, 06.IX.2012, P.
Estók leg.; 1 ♀, Hungary, [Borsod-Abaúj-Zemplén county],
Miskolc, 10.IX.2012, P. Estók leg.; 1 ♂, Hungary, [Borsod-Abaúj-
Zemplén county], Miskolc, 14.IX.2012, P. Estók leg.; 1 ♀, Hungary,
[Borsod-Abaúj-Zemplén county], Miskolc, 17.IX.2012, P. Estók
leg.; 1 ♂, Hungary, [Borsod-Abaúj-Zemplén county], Miskolc,
27.IX.2012, P. Estók leg.; 1 ♀, Hungary, [Heves county],
Felsőtárkány, 18.V.2013, P. Estók leg.; 1 ♀, Hungary, [Heves
county], Felsőtárkány, 15.VII.2013, P. Estók leg.; 1 ♀ and 1 ♂,
Hungary, [Borsod-Abaúj-Zemplén county], Miskolc, 04.IX.2013, P.
Estók leg.; 1 ♂, Hungary, [Borsod-Abaúj-Zemplén county],
Bükkszentkereszt, 05.IX.2013, P. Estók leg.; 1 ♀, Hungary,
[Borsod-Abaúj-Zemplén county], Háromhuta, 01.VIII.2014, P.
Estók leg.; 1 ♀, Hungary, [Heves county], Felsőtárkány,
08.IX.2014, P. Estók leg.
Romania, Slovakia,
Slovenia, Sweden,
Switzerland, The
Netherlands
Myotis emarginatus—Aellen (1963)
Myotis myotis—Dudich (1925) (doubtful record), Hůrka (1964),
Nowosad (1974), Hutson (1984), Nowosad (1987), Czuppon
(2001), Heddergott & Claußen (2004), Rupp et al. (2004), Güttinger
et al. (2011), Scheffler (2012)
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TABLE 1. (Continued)
Bat fly species Host species, references and new data Distribution of bat fly
species
Myotis mystacinus—Theodor (1954), Theodor & Moscona (1954),
Aellen (1963), Hůrka (1964), Walter (1987)
Myotis nattereri—Theodor (1954), Theodor & Moscona (1954),
Hůrka (1964), Kock (1973), Nowosad (1974), Walter & Benk
(1982), Hutson (1984), Nowosad (1987), Czuppon (2001), Scheffler
& Ressler (2005, 2007), Topál (2011)
Nyctalus leisleri—Walter (2004)
Nyctalus noctula—Walter (1987, 2004)
Pipistrellus pipistrellus—Theodor & Moscona (1954), Hutson
(1984), Walter (1987, 2004)
Plecotus auritus—Hůrka (1964), Nowosad (1974, 1987)
Rhinolophus hipposideros—Hůrka (1964), Hutson (1984)
Vespertilio murinus—Theodor & Moscona (1954) (doubtful
record), Hutson (1984)
Nycteribia latreillii (Leach)
Syn.: N. biscutata Gil Collado
Listropodia latreillii africana
Karaman
Eptesicus serotinus—Gil Collado (1934), Theodor (1954), Hutson
(1984), Baagøe (2011a)
Albania, Austria,
Belgium, Bosnia and
Herzegovina, Bulgaria,
Croatia, Czech
Republic, European
Turkey, France,
Germany, Greece,
Hungary, Italy (incl.
Sardinia and Sicily),
Macedonia, Poland,
Portugal, Romania,
Slovakia, Spain,
Switzerland, The
Netherlands, ex-
Yug o s la v ia
Miniopterus schreibersii—Grulich & Povolný (1955), Hůrka (1964,
1980), Krištofík (1982)
Myotis blythii—Hůrka (1962, 1964), Balcells (1968a, 1968b),
Hůrka (1984), Nowosad (1990), Beaucournu & Noblet (1996),
Czuppon (2001), Scheffler (2011b), Ševčík et al. (2013). Present
study: 2 ♀, Hungary, [Baranya county], Szársomlyó 1998, P.
Paulovics leg.
Myotis capaccinii—Theodor & Moscona (1954), Theodor (1967)
Myotis daubentonii—Colyer & Hammond (1951), Walter (2004)
Myotis emarginatus—Krištofík (1982)
Myotis myotis—Theodor (1954), Theodor & Moscona (1954),
Grulich & Povolný (1955), Aellen (1963), Hůrka (1964), Kock
(1973), Krištofík (1982), Hůrka (1984), Hutson (1984), Nowosad
(1990), Beaucournu & Noblet (1996), Rupp (1999), Czuppon
(2001), Rupp et al. (2004), Scheffler (2009), Uhrin et al. (2010),
Güttinger et al. (2011), Scheffler (2011b), Krištofík & Danko
(2012), Scheffler et al. (2013). Present study: 1 ♀, Hungary,
[Borsod-Abaúj-Zemplén county], Miskolc, 11.IX.2012, P. Estók
leg.
Myotis nattereri—Beaucournu (1961)
Nyctalus leisleri—Kaňuch et al. (2005)
Plecotus auritus—Theodor & Moscona (1954)
Rhinolophus euryale—Theodor & Moscona (1954)
Rhinolophus ferrumequinum—Theodor (1954, 1967), Hutson
(1984)
Rhinolophus hipposideros—Theodor & Moscona (1954), Theodor
(1967)
Vespertilio murinus—Westwood (1835) (doubtful record), Theodor
& Moscona (1954) (doubtful record), Theodor (1967)
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EUROPEAN BAT FLY HOST ASSOCIATIONS
TABLE 1. (Continued)
Bat fly species Host species, references and new data Distribution of bat fly
species
Nycteribia pedicularia
Latreille
Eptesicus serotinus—Falcoz (1926) (doubtful record), Scott (1936),
Theodor (1954), Hůrka (1970)
Albania, Bosnia and
Herzegovina, Bulgaria,
Croatia, Cyprus,
European Turkey,
France (incl. Corsica),
Greece, Italy (incl.
Sardinia and Sicily),
Latvia, Romania,
Slovenia, Spain
(Mallorca) Switzerland,
ex-Yugoslavia
Miniopterus schreibersii—Falcoz (1926) (doubtful record), Scott
(1936), Theodor (1954), Hůrka (1970)
Myotis blythii—Falcoz (1926) (doubtful record), Theodor (1954),
Hůrka (1962), Scheffler (2011b)
Myotis capaccinii—Theodor & Moscona (1954), Aellen (1963),
Hůrka (1970), Kock (1974), Hůrka (1984), Noblet (1995), Kock &
Quetglas (2003), Scheffler (2011b), Scheffler et al. (2013)
Myotis dasycneme—Falcoz (1926) (doubtful record), Theodor
(1954), Jaunbauere et al. (2008)
Myotis daubentonii—Falcoz (1926) (doubtful record), Theodor
(1954), Jaunbauere et al. (2008)
Myotis emarginatus—Theodor & Moscona (1954)
Myotis myotis—Falcoz (1926) (doubtful record), Theodor (1954),
Theodor & Moscona (1954), Hutson (1984), Güttinger et al. (2011),
Scheffler (2011b), Scheffler et al. (2013)
Myotis mystacinus—Theodor (1954)
Myotis nattereri—Theodor (1954)
Nyctalus noctula—Falcoz (1926) (doubtful record), Scott (1936),
Theodor (1954), Hutson (1984)
Rhinolophus ferrumequinum—Hutson (1984)
Rhinolophus hipposideros—Falcoz (1926) (doubtful record), Scott
(1936), Theodor (1954)
Rousettus aegyptiacus—[Theodor & Moscona (1954; Palestine)]
[Tadarida teniotis—Theodor & Moscona (1954; Palestine)]
Nycteribia schmidlii Schiner
Syn: Nycteribia blasii
Kolenati
Barbastella barbastellus—Hutson (1984) Albania, Bosnia and
Herzegovina, Bulgaria,
Croatia, Cyprus,
European Turkey,
France (incl. Corsica),
Germany, Greece (incl.
Crete), Hungary, Italy
(incl. Sicily
andSardinia),
Macedonia, Portugal,
Romania, Slovakia,
Spain (incl. Mallorca),
Switzerland, Ukraine,
ex-Yugoslavia
Miniopterus schreibersii—Falcoz (1924), Dudich (1925), Falcoz
(1926), Theodor (1954), Grulich & Povolný (1955), Soós (1955),
Aellen (1963), Hůrka (1964), Theodor (1967), Balcells (1968b),
Kock (1974), Krištofík (1982), Hůrka (1984), Estrada-Peña &
Serra-Cobo (1991), Beaucournu & Noblet (1996), Imaz et al.
(1999), Kock & Quetglas (2003), Krištofík & Danko (2012),
Scheffler et al. (2013), Ševčík et al. (2013). Present study: 10 ♀ and
3 ♂, Hungary, [Baranya county], Szársomlyó, 1998, P. Paulovics
leg.; 1 ♀ and 1 ♂, Hungary, [Heves county], Felsőtárkány,
13.IX.2012, P. Estók leg.; 5 ♀ and 2 ♂, Hungary, [Heves county],
Felsőtárkány, 02.X.2012, P. Estók leg.; 11 ♀ and 2 ♂, Hungary,
[Heves county], Felsőtárkány, 04.X.2012, P. Estók leg.
Myotis bechsteinii—Grulich & Povolný (1955), Hutson (1984),
Baagøe (2011b)
Myotis blythii—Hůrka (1962), Krištofík (1982)
Myotis capaccinii—Theodor & Moscona (1954), Theodor (1967),
Kock (1974), Ševčík et al. (2013)
Myotis daubentonii—Falcoz (1924, 1926), Theodor (1954), Theodor
& Moscona (1954), Soós (1955), Hutson (1984). Present study: 1 ♂,
Hungary, [Pest county], Szentendre, 29.VI.2014, P. Estók leg.; 1 ♂,
Hungary, [Heves county], Felsőtárkány, 08.IX.2014, P. Estók leg.
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TABLE 1. (Continued)
Bat fly species Host species, references and new data Distribution of bat fly
species
Myotis myotis—Falcoz (1924), Dudich (1925), Falcoz (1926), Theodor
(1954), Theodor & Moscona (1954), Grulich & Povolný (1955), Soós
(1955), Aellen (1963), Hůrka (1964), Balcells (1968b), Hůrka (1980),
Krištofík (1982), Hutson (1984), Güttinger et al. (2011). Present study: 1 ♀
and 1 ♂, Hungary, [Baranya county], Szársomlyó, 1998, P. Paulovics leg.
Myotis mystacinus—Aellen (1963)
[Pipistrellus kuhlii—Aktaş & Hasbenli (1994; Turkey)]
Pipistrellus pipistrellus—Falcoz (1924, 1926), Theodor (1954), Soós
(1955), Hutson (1984), Nowosad et al. (1987)
Plecotus auritus—Theodor (1967), Krištofík (1982)
Rhinolophus euryale—Soós (1955), Theodor (1967)
Rhinolophus ferrumequinum—Theodor (1954), Theodor & Moscona
(1954), Balcells (1968b), Hutson (1984)
Rhinolophus hipposideros—Grulich & Povolný (1955), Hutson (1984)
Rhinolophus mehelyi—Falcoz (1924, 1926), Theodor (1954), Soós (1955)
[Rousettus aegyptiacus—Theodor & Moscona (1954; Palestine)]
Vespertilio murinus—Theodor (1967)
Nycteribia vexata Wes twood
Miniopterus schreibersii—Falcoz (1926), Theodor (1954), Grulich &
Povolný (1955), Soós (1955), Krištofík (1982)
Albania, Austria, Belgium,
Bosnia and Herzegovina,
Syn.: Nycteribia montaguei
Kolenati
Myotis bechsteinii—Soós (1955), Hutson (1984), Baagøe (2011b)
Bulgaria, Croatia, Czech
Republic, European
Acrocholidia bechsteinii
Kolenati
Myotis blythii—Theodor (1954), Hůrka (1962, 1964), Balcells (1968a,
1968b), Czuppon (2001), Scheffler et al. (2013)
Turkey, France, Cermany,
Greece, Hungary, Italy
Nycteribia ercolanii Rondani
Myotis daubentonii—Hutson (1984)
(incl.Sardinia and Sicily),
Myotis myotis—Falcoz (1926), Theodor (1954), Grulich & Povolný (1955),
Soós (1955), Aellen (1963), Hůrka (1964), Kock (1973), Krištofík (1982),
Hůrka (1984), Hutson (1984), Nowosad (1990), Beaucournu & Noblet
(1996), Czuppon (2001), Kock & Quetglas (2003), Uhrin et al. (2010),
Güttinger et al. (2011), Scheffler (2011b), Scheffler et al. (2013). Present
study: 3 ♀, Hungary, [Veszprém county], Kislőd, 19.VII.1999, P. Paulovics
leg.; 1 ♂, Hungary, [Veszprém county ], Eplény, 21.VII.1999, P. Paulovics
leg.; 1 ♀ and 1 ♂, Hungary, [Heves county], Noszvaj, 17.VIII.2012, P.
Estók leg.
Macedonia, Poland,
Portugal, Romania,
Slovakia, Slovenia,
Spain, Switzerland,
Ukraine, ex-Yugoslavia
Myotis mystacinus—Aellen (1963), Kock (1999)
Nyctalus leisleri—Kaňuch et al. (2005)
[Pipistrellus kuhlii—Aktaş & Hasbenli (1994; Turkey)]
Plecotus auritus—Theodor (1954), Theodor & Moscona (1954)
Rhinolophus euryale—Falcoz (1926), Theodor (1954), Soós (1955)
Rhinolophus ferrumequinum—Falcoz (1926), Theodor (1954), Soós (1955),
Hutson (1984)
Rhinolophus hipposideros—Falcoz (1926), Theodor (1954), Theodor &
Moscona (1954), Soós (1955), Hutson (1984)
Vespertilio murinus—Westwood (1835) (doubtful record), Kolenati (1857)
(doubtful record)
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TABLE 1. (Continued)
Bat fly species Host species, references and new data Distribution of bat fly
species
Penicillidia conspicua
Speiser
Miniopterus schreibersii—Falcoz (1923), Dudich (1925), Falcoz
(1926), Theodor (1954), Grulich & Povolný (1955), Soós (1955),
Aellen (1963), Hůrka (1964, 1970), Krištofík (1982), Hůrka (1984),
Kock (1989), Beaucournu & Noblet (1996), Krištofík & Danko
(2012), Scheffler et al. (2013), Ševčík et al. (2013). Present study: 2
♂, Hungary, [Baranya county], Szársomlyó, 19.V.1998, P. Paulovics
leg.; 2 ♀ and 1 ♂, Hungary, [Baranya county], Szársomlyó, 1998, P.
Paulovics leg.; 2 ♀ and 1 ♂, Hungary, [Heves county],
Felsőtárkány, 13.IX.2012, P. Estók leg.; 2 ♀ and 1 ♂, Hungary,
[Heves county], Felsőtárkány, 02.X.2012, P. Estók leg.; 1 ♀,
Hungary, [Borsod-Abaúj-Zemplén county], Miskolc, 15.IX.2013, P.
Estók leg.; 4 ♀ and 3 ♂, Hungary, [Heves county], Felsőtárkány,
23.VIII.2014, P. Estók leg.
Albania, Bulgaria,
Croatia, European
Turkey, France (inc.
Corsica), Greece,
Hungary, Italy (Sicily
and Sardinia),
Macedonia, Portugal,
Romania, Slovakia,
Spain (incl. Mallorca),
Switzerland, ex-
Yug o s la v ia
Myotis blythii—Nowosad et al. (1987), Scheffler (2011b)
Myotis capaccinii—Theodor (1967)
Myotis myotis—Dudich (1925), Falcoz (1926), Theodor (1954),
Soós (1955), Hůrka (1964), Hutson (1984), Güttinger et al. (2011),
Scheffler (2011b)
Myotis mystacinus—Theodor (1954)
[Pipistrellus kuhlii—Aktaş & Hasbenli (1994; Turkey)]
Plecotus auritus—Theodor (1967)
Rhinolophus blasii—Falcoz (1926), Theodor (1954)
Rhinolophus euryale—Falcoz (1926), Theodor (1954), Grulich &
Povolný (1955), Soós (1955). Present study: 1 ♀, Hungary,
[Borsod-Abaúj-Zemplén county], Edelény, 13.XI.2014, P. Estók
leg.
Rhinolophus ferrumequinum—Grulich & Povolný (1955), Hutson
(1984)
Rhinolophus hipposideros—Theodor (1954), Hutson (1984)
Vespertilio murinus—Theodor (1967)
Penicillidia dufourii
(Westwood)
Syn.: Nycteribia westwoodi
Guérin-Méneville
Nycteribia frauenfeldi
Kolenati
Megistopoda leachii Kolenati
Eptesicus serotinus—Hůrka (1970) Albania, Austria,
Belgium, Bosnia and
Herzegovina, Bulgaria,
Croatia, Czech
Republic, European
Turkey, France (incl.
Corsica), Germany,
Greece, Hungary, Italy
(incl. Sardinia and
Sicily), Macedonia,
Poland, Portugal,
Romania, Slovakia,
Spain (incl. Mallorca),
Switzerland, The
Netherlands, Ukraine,
ex-Yugoslavia
Miniopterus schreibersii—Dudich (1925), Theodor (1954), Grulich
& Povolný (1955), Aellen (1963), Hůrka (1970, 1980), Krištofík
(1982), Hůrka (1984), Estrada-Peña & Serra-Cobo (1991),
Beaucournu & Noblet (1996), Imaz et al. (1999), Kock & Quetglas
(2003, as “P. dufourii dufourii”), Krištofík & Danko (2012),
Scheffler et al. (2013). Present study: 7 ♀ and 6 ♂, Hungary,
[Baranya county], Szársomlyó, 19.V.1998, P. Paulovics leg.; 1 ♂,
Hungary, [Baranya county], Szársomlyó, 1998, P. Paulovics leg.
Myotis blythii—Falcoz (1924, 1926), Theodor (1954), Soós (1955),
Hůrka (1962, 1964), Balcells (1968a, 1968b), Hůrka (1970, 1984),
Czuppon (2001), Scheffler (2011b), Scheffler et al. (2013), Ševčík
et al. (2013). Present study: 1 ♀ and 1 ♂, Hungary, [Baranya
county], Szársomlyó, 1998, P. Paulovics leg.
Myotis capaccinii—Falcoz (1924, 1926), Theodor (1954), Theodor
& Moscona (1954), Soós (1955), Aellen (1963), Hůrka (1970,
1984), Nowosad et al. (1987), Noblet (1995), Beaucournu & Noblet
(1996), Kock & Quetglas (2003, as “P. dufourii dufourii”), Scheffler
(2011b), Scheffler et al. (2013), Ševčík et al. (2013)
Myotis dasycneme—Walter (2004)
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TABLE 1. (Continued)
Bat fly species Host species, references and new data Distribution of bat fly
species
Myotis daubentonii*—Aellen (1963), Hutson (1984), Scheffler et
al. (2013). Present study: 1 ♀, Hungary, [Borsod-Abaúj-Zemplén
county], Miskolc, 29.VIII.2012, P. Estók leg.
Myotis emarginatus—Theodor & Moscona (1954), Hůrka (1980),
Imaz et al. (1999)
Myotis myotis—Dudich (1925), Falcoz (1926), Theodor (1954),
Theodor & Moscona (1954), Aellen (1955), Grulich & Povolný
(1955), Soós (1955), Aellen (1963), Hůrka (1964), Balcells (1968a),
Kock (1973), Krištofík (1982), Hůrka (1984), Hutson (1984),
Noblet (1995), Beaucournu & Noblet (1996), Imaz et al. (1999),
Czuppon (2001), Kock & Quetglas (2003, as “P. dufourii dufourii”),
Güttinger et al. (2011), Scheffler (2011b), Krištofík & Danko
(2012), Scheffler et al. (2013). Present study: 2 ♀ and 2 ♂, Hungary,
[Baranya county], Szársomlyó, 1998, P. Paulovics leg.; 1 ♀ and 1 ♂,
Hungary, [Heves/Nógrád county], Mátra Mts. 05.VI.1999, P.
Paulovics leg.; 1 ♀, Hungary, [Borsod-Abaúj-Zemplén county],
Miskolc, 10.IX.2012, P. Estók leg.; 1 ♀, Hungary, [Borsod-Abaúj-
Zemplén county], Miskolc, 19.VIII.2014, P. Estók leg.; 1 ♀,
Hungary, [Heves county], Noszvaj, 06.IX.2014, P. Estók leg.; 1 ♂,
Hungary, [Heves county], Nagyvisnyó, 17.IX.2014, P. Estók leg.; 1
♀ and 1 ♂, Hungary, [Borsod-Abaúj-Zemplén county], Miskolc,
21.IX.2014, P. Estók leg.
Myotis mystacinus—Theodor & Moscona (1954) (doubtful record)
Myotis nattereri—Theodor (1954), Hutson (1984), Topál (2011),
Ševčík et al. (2013)
Plecotus auritus—Theodor (1967)
Rhinolophus blasii—[Theodor & Moscona (1954; Palestine)], Soós
(1955)
Rhinolophus euryale—Falcoz (1926), Grulich & Povolný (1955),
Soós (1955), Hůrka (1980)
Rhinolophus ferrumequinum—Falcoz (1926), Theodor & Moscona
(1954), Soós (1955), Hutson (1984)
Rhinolophus hipposideros—Falcoz (1926), Theodor & Moscona
(1954), Soós (1955), Hutson (1984)
Rhinolophus mehelyi—Scott (1936)
Tadarida teniotus—Scott (1936)
Vespertilio murinus—Westwood (1835) (doubtful record), Theodor
(1967)
Penicillidia monoceros
Speiser
Myotis dasycneme—Skuratowicz (1962), Czuppon & Molnár
(2001), Kock (2004), Scheffler (2009), [Orlova (2011; Russia)]
Czech Republic,
Denmark, Finland,
Germany, Hungary,
Kaliningrad Region,
Norway, Poland,
Sweden
Myotis daubentonii—Theodor (1954), Nowosad (1974), Hůrka
(1984), Hutson (1984), Müller & Ohlendorf (1984), Nowosad
(1987, 1990), Kock (1999), Rupp (1999), Heddergott & Claußen
(2004), Kock (2004), Rupp et al. (2004), Scheffler & Ressler
(2007), Scheffler (2008), Lindecke & Scheffler (2011)
Myotis myotis—Heddergott & Claußen (2004)
Myotis mystacinus—Nowosad (1987)
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TABLE 1. (Continued)
Bat fly species Host species, references and new data Distribution of bat fly
species
Myotis nattereri—Skuratowicz (1962), Hutson (1984), Topál (2011)
Nyctalus noctula—Czuppon & Molnár (2001), Rupp et al. (2004)
Plecotus auritus—Theodor (1954)
Vespertilio murinus—Hutson (1984)
Phthiridium biarticulatum
Hermann
Syn.:
Celeripes vespertilionis
Montagu
Phthiridium hermanni Leach
Barbastella barbastellus—Krištofík (1982) Albania, Austria,
Bulgaria, Croatia,
Cyprus, Czech
Republic, European
Turkey, France (incl.
Corsica), Germany,
Great Britain, Greece
(incl. Crete), Hungary,
Ireland, Italy (incl.
Sicily and Sardinia),
Latvia, Macedonia,
Portugal, Romania,
Slovakia, Slovenia,
Spain, Switzerland, ex-
Yug o s la v ia
Miniopterus schreibersii—Falcoz (1926), Theodor (1954), Theodor
& Moscona (1954), Soós (1955), Hůrka (1964, 1980)
Myotis blythii—Hůrka (1962, 1964)
Myotis dasycneme—Jaunbauere et al. (2008)
Myotis daubentonii—[Theodor & Moscona (1954; Morocco)],
Theodor (1967)
Myotis emarginatus—Theodor (1967)
Myotis myotis—Falcoz (1926), Theodor (1954), [Theodor &
Moscona (1954; Palestine)], Grulich & Povolný (1955), Soós
(1955), Hůrka (1964, 1980), Krištofík (1982), Hutson (1984),
Güttinger et al. (2011)
Pipistrellus pipistrellus—Colyer & Hammond (1951), Theodor
(1967), Hutson (1984)
Plecotus auritus—Falcoz (1926), Theodor (1954), Theodor &
Moscona (1954), Soós (1955)
Rhinolophus blasii—Falcoz (1926), Theodor (1954), Soós (1955),
[Amr & Qumsiyeh (1993; Jordan)], Scheffler et al. (2013), Ševčík
et al. (2013)
Rhinolophus euryale—Falcoz (1926), Theodor (1954), Grulich &
Povolný (1955), Soós (1955), Hůrka (1964), Theodor (1967), Imaz
et al. (1999), Scheffler et al. (2013)
Rhinolophus ferrumequinum—Westwood (1835), Falcoz (1923,
1926), Colyer & Hammond (1951), Theodor (1954), Theodor &
Moscona (1954), Aellen (1955), Grulich & Povolný (1955), Soós
(1955), Humphries (1959), Aellen (1963), Hůrka (1964), Theodor
(1967), Balcells (1968b), Krištofík (1982), Hůrka (1984), Hutson
(1984), Kock (1989), Imaz et al. (1999), Krištofík & Danko (2012),
Scheffler et al. (2013), Ševčík et al. (2013)
Rhinolophus hipposideros—Austen (1906), Dudich (1925), Falcoz
(1926), Colyer & Hammond (1951), Theodor (1954), Theodor &
Moscona (1954), Grulich & Povolný (1955), Soós (1955),
Humphries (1959), Aellen (1963), Hůrka (1964), Rageau &
Mouchet (1967), Balcells (1968b), Krištofík (1982), Hutson (1984),
Ševčík et al. (2011)
Rhinolophus mehelyi—Falcoz (1926), Theodor (1954), Soós (1955)
Phthiridium integrum
(Theodor & Moscona)
Eptesicus anatolicus—Ševčík et al. (2013) Cyprus
Rhinolophus blasii—Theodor (1954) (locality unknown), [Theodor
& Moscona (1954; Palestine)]
Rhinolophus ferrumequinum—Theodor (1954) (locality unknown),
[Theodor & Moscona (1954; Palestine)]
[Rhinolophus hipposideros—Amr & Qumsiyeh (1993; Jordan)]
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TABLE 2. List of bat species and their associated bat flies (Nycteribiidae and Streblidae) in Europe, based on the
literature and previously unpublished data. The main bat fly species are in bold font; records from outside Europe are
listed in square brackets. Host associations marked with and asterisk (*) are new for Hungary.
TABLE 1. (Continued)
Bat fly species Host species, references and new data Distribution of bat fly
species
Streb li da e Kolenati
Subfamily Brachytarsininae (Speiser)
Brachytarsina flavipennis
Macquart
Syn.:
Raymondia kollari
Frauenfeld
Miniopterus schreibersii—Falcoz (1926), Theodor (1954),
[Theodor & Moscona (1954; Palestine)]
Albania, European
Turkey, France, Greece,
Italy (incl. Sardinia),
Portugal, Spain
Myotis aurascens—Ševčík et al. (2013)
Myotis myotis—Theodor (1954)
Rhinolophus blasii—Theodor (1954), [Theodor & Moscona (1954;
Palestine)], [Amr & Qumsiyeh (1993; Jordan)], Ševčík et al. (2013)
Rhinolophus euryale—Falcoz (1926), Theodor (1954), [Theodor &
Moscona (1954; Palestine)], [Benda et al. (2010; Jordan)]
Rhinolophus ferrumequinum—Falcoz (1926), Theodor (1954),
[Theodor & Moscona (1954; Palestine)], Beaucournu & Noblet
(1996), Ševčík et al. (2013)
Rhinolophus hipposideros—Falcoz (1926), Theodor (1954),
[Theodor & Moscona (1954; Palestine)]
Rhinolophus mehelyi—Falcoz (1926), Theodor (1954), [Walter &
Ebenau (1997; Syria)]
Vespertilio murinus—Jobling (1934; doubtful record)
Bat species Bat fly species
Pteropodidae Gray
Rousettus aegyptiacus (E. Geoffroy) [Nycteribia pedicularia—Theodor & Moscona (1954; Palestine)]
[Nycteribia schmidlii—Theodor & Moscona (1954; Palestine)]
Rhinolophidae Gray
Rhinolophus blasii Peters Penicillidia dufourii—[Theodor & Moscona (1954; Palestine)], Soós (1955)
Penicillidia conspicua—Falcoz (1926), Theodor (1954)
Phthiridium biarticulatum—Falcoz (1926), Theodor (1954), Soós (1955), [Amr
& Qumsiyeh (1993; Jordan)], Scheffler et al. (2013), Ševčík et al. (2013)
Phthiridium integrum—Theodor (1954) (locality unknown), [Theodor &
Moscona (1954; Palestine)]
Brachytarsina flavipennis [Streblidae]—Theodor (1954), [Theodor & Moscona
(1954; Palestine)], [Amr & Qumsiyeh (1993; Jordan)], Ševčík et al. (2013)
Rhinolophus euryale Blasius Nycteribia latreillii—Theodor & Moscona (1954)
Nycteribia schmidlii—Soós (1955), Theodor (1967)
Nycteribia vexata—Falcoz (1926), Theodor (1954), Soós (1955)
Penicillidia conspicua—Falcoz (1926), Theodor (1954), Grulich & Povolný
(1955), Soós (1955). Present study: see Table 1 for details.
Penicillidia dufourii—Falcoz (1926), Grulich & Povolný (1955), Soós (1955),
Hůrka (1980)
Phthiridium biarticulatum—Falcoz (1926), Theodor (1954), Grulich & Povolný
(1955), Soós (1955), Hůrka (1964), Theodor (1967), Imaz et al. (1999), Scheffler
et al. (2013)
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TABLE 2. (Continued)
Bat species Bat fly species
Brachytarsina flavipennis [Streblidae]—Falcoz (1926), Theodor (1954),
[Theodor & Moscona (1954; Palestine)], [Benda et al. (2010; Jordan)]
Rhinolophus ferrumequinum (Schreber) Basilia nana—Hutson (1984)
Nycteribia latreillii—Theodor (1954, 1967), Hutson (1984)
Nycteribia pedicularia—Hutson (1984)
Nycteribia schmidlii—Theodor (1954), Theodor & Moscona (1954), Balcells
(1968b), Hutson (1984)
Nycteribia vexata—Falcoz (1926), Theodor (1954), Soós (1955), Hutson (1984)
Penicillidia conspicua—Grulich & Povolný (1955), Hutson (1984)
Penicillidia dufourii—Falcoz (1926), Theodor & Moscona (1954), Soós (1955),
Hutson (1984)
Phthiridium biarticulatum—Westwood (1835), Falcoz (1923, 1926), Colyer &
Hammond (1951), Theodor (1954), Theodor & Moscona (1954), Aellen (1955),
Grulich & Povolný (1955), Soós (1955), Humphries (1959), Aellen (1963),
Hůrka (1964), Theodor (1967), Balcells (1968b), Krištofík (1982), Hůrka (1984),
Hutson (1984), Kock (1989), Imaz et al. (1999), Krištofík & Danko (2012),
Scheffler et al. (2013), Ševčík et al. (2013)
Phthiridium integrum—Theodor (1954) (locality unknown), [Theodor &
Moscona (1954; Palestine)]
Brachytarsina flavipennis [Streblidae]—Falcoz (1926), Theodor (1954),
[Theodor & Moscona (1954; Palestine)], Beaucournu & Noblet (1996), Ševčík et
al. (2013)
Rhinolophus hipposideros (Bechstein) Basilia nana—Theodor & Moscona (1954)
Basilia nattereri—Hutson (1984)
Nycteribia kolenatii—Hůrka (1964), Hutson (1984)
Nycteribia latreillii—Theodor & Moscona (1954), Theodor (1967)
Nycteribia pedicularia—Falcoz (1926) (doubtful record), Scott (1936), Theodor
(1954)
Nycteribia schmidlii—Grulich & Povolný (1955), Hutson (1984)
Nycteribia vexata—Falcoz (1926), Theodor (1954), Theodor & Moscona (1954),
Soós (1955), Hutson (1984)
Penicillidia conspicua—Theodor (1954), Hutson (1984)
Penicillidia dufourii—Falcoz (1926), Theodor & Moscona (1954), Soós (1955),
Hutson (1984)
Phthiridium biarticulatum—Austen (1906), Dudich (1925), Falcoz (1926),
Colyer & Hammond (1951), Theodor (1954), Theodor & Moscona (1954),
Grulich & Povolný (1955), Soós (1955), Humphries (1959), Aellen (1963),
Hůrka (1964), Rageau & Mouchet (1967), Balcells (1968b), Krištofík (1982),
Hutson (1984), Ševčík et al. (2011)
[Phthiridium integrum—Amr & Qumsiyeh (1993; Jordan)]
Brachytarsina flavipennis [Streblidae]—Falcoz (1926), Theodor (1954),
[Theodor & Moscona (1954; Palestine)]
Rhinolophus mehelyi Matschie Nycteribia schmidlii—Falcoz (1924, 1926), Theodor (1954), Soós (1955)
Penicillidia dufourii—Scott (1936)
Phthiridium biarticulatum—Falcoz (1926), Theodor (1954), Soós (1955)
Brachytarsina flavipennis [Streblidae]—Falcoz (1926), Theodor (1954), [Walter
& Ebenau (1997; Syria)]
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TABLE 2. (Continued)
Bat species Bat fly species
Vespertilionidae Gray
Barbastella barbastellus (Schreber) Basilia italica—Czuppon & Molnár (2001). Present study: see Table 1 for
details.
Basilia nana—Grulich & Povolný (1955)
Nycteribia kolenatii—Haitlinger (1978), Nowosad (1987), Scheffler (2011a)
Nycteribia schmidlii—Hutson (1984)
Phthiridium biarticulatum—Krištofík (1982)
Eptesicus anatolicus Felten Phthiridium integrum—Ševčík et al. (2013)
Eptesicus bottae (Peters) –
Eptesicus isabellinus Temminck –
Eptesicus nilssonii (Keyserling & Bläsius) –
Eptesicus serotinus (Schreber) Basilia italica—Czuppon & Molnár (2001)
Basilia mediterranea—Kock & Quetglas (2003, as “E. serotinus boscai”)
Basilia nattereri—Kolenati (1857) (doubtful record), Theodor (1954), Theodor
& Moscona (1954), Hůrka (1980), Hutson (1984), Baagøe (2011a)
Nycteribia kolenatii—Theodor (1954), Theodor & Moscona (1954), Hůrka
(1964), Hutson (1984), Baagøe (2011a)
Nycteribia latreillii—Gil Collado (1934), Theodor (1954), Hutson (1984),
Baagøe (2011a)
Nycteribia pedicularia—Falcoz (1926) (doubtful record), Scott (1936), Theodor
(1954), Hůrka (1970)
Penicillidia dufourii—Hůrka (1970)
Hypsugo savii (Bonaparte) Basilia mediterranea—Beaucournu & Noblet (1994)
Myotis alcathoe Helversen & Heller Basilia italica*—Danko et al. (2010). Present study: see Table 1 for details.
Basilia mongolensis nudior—Heddergott (2009)
Myotis aurascens Kusjakin Basilia mongolensis nudior—Ševčík et al. (2013)
Brachytarsina flavipennis [Streblidae]—Ševčík et al. (2013)
Myotis bechsteinii (Kuhl) Basilia nana—Theodor (1954), Theodor & Moscona (1954), Grulich & Povolný
(1955), Aellen (1963), Hůrka (1964), Kock (1973), Hůrka (1984), Hutson
(1984), Nowosad (1987), Nowosad (1990), Beaucournu & Noblet (1996), Kock
(1999), Rupp (1999), Czuppon (2001), Czuppon & Molnár (2001), Rupp et al.
(2004), Reckardt & Kerth (2006), Scheffler (2010), Scheffler & Hiller (2010),
Baagøe (2011b), Krištofík & Danko (2012), Szentiványi et al. (2016). Present
study: see Table 1 for details.
Basilia nattereri—Dudich (1925) (doubtful record), Soós (1955). Present study:
see Table 1 for details.
Nycteribia kolenatii*—Aellen (1963), Hůrka (1964, 1984), Hutson (1984),
Nowosad (1987), Scheffler (2010), Baagøe (2011b). Present study: see Table 1
for details.
Nycteribia schmidlii—Grulich & Povolný (1955), Hutson (1984), Baagøe
(2011b)
Nycteribia vexata—Soós (1955), Hutson (1984), Baagøe (2011b)
Myotis blythii (Tomes) Basilia nana—Hůrka (1964, 1980), Czuppon & Molnár (2001)
Nycteribia latreillii—Hůrka (1962, 1964), Balcells (1968a, 1968b), Hůrka
(1984), Nowosad (1990), Beaucournu & Noblet (1996), Czuppon (2001),
Scheffler (2011b), Ševčík et al. (2013). Present study: see Table 1 for details.
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TABLE 2. (Continued)
Bat species Bat fly species
Nycteribia pedicularia—Falcoz (1926) (doubtful record), Theodor (1954),
Hůrka (1962), Scheffler (2011b)
Nycteribia schmidlii—Hůrka (1962), Krištofík (1982)
Nycteribia vexata—Theodor (1954), Hůrka (1962, 1964), Balcells (1968a,
1968b), Czuppon (2001), Scheffler et al. (2013)
Penicillidia conspicua—Nowosad et al. (1987), Scheffler (2011b)
Penicillidia dufourii—Falcoz (1924, 1926), Theodor (1954), Soós (1955),
Hůrka (1962, 1964), Balcells (1968a, 1968b), Hůrka (1970, 1984), Czuppon
(2001), Scheffler (2011b), Scheffler et al. (2013), Ševčík et al. (2013). Present
study: see Table 1 for details.
Phthiridium biarticulatum—Hůrka (1962, 1964)
Myotis brandtii (Eversmann) Basilia italica—Hůrka (1964, as “Myotis mystacinus”), Hůrka (1980, 1984),
Czuppon (2001), Czuppon & Molnár (2001). Present study: see Table 1 for
details.
Nycteribia kolenatii—Rupp (1999), Rupp et al. (2004), Walter (2004)
Myotis capaccinii (Bonaparte) Nycteribia latreillii—Theodor & Moscona (1954), Theodor (1967)
Nycteribia pedicularia—Theodor & Moscona (1954), Aellen (1963), Hůrka
(1970), Kock (1974), Hůrka (1984), Noblet (1995), Kock & Quetglas (2003),
Scheffler (2011b), Scheffler et al. (2013)
Nycteribia schmidlii—Theodor & Moscona (1954), Theodor (1967), Kock
(1974), Ševčík et al. (2013)
Penicillidia conspicua—Theodor (1967)
Penicillidia dufourii—Falcoz (1924, 1926), Theodor (1954), Theodor &
Moscona (1954), Soós (1955), Aellen (1963), Hůrka (1970, 1984), Nowosad et
al. (1987), Noblet (1995), Beaucournu & Noblet (1996), Kock & Quetglas
(2003,as “P. dufourii dufourii”), Scheffler (2011b), Scheffler et al. (2013),
Ševčík et al. (2013)
Myotis dasycneme (Boie) Basilia nana—Theodor (1954), Theodor & Moscona (1954), Theodor (1967),
Hutson (1984), Nowosad (1987)
Nycteribia kolenatii—Theodor (1954), Theodor & Moscona (1954), Hůrka
(1964), Hutson (1984), [Orlova (2011; Russia)]
Nycteribia pedicularia—Falcoz (1926) (doubtful record), Theodor (1954),
Jaunbauere et al. (2008)
Penicillidia dufourii—Walter (2004)
Penicillidia monoceros—Skuratowicz (1962), Czuppon & Molnár (2001), Kock
(2004), Scheffler (2009), [Orlova (2011; Russia)]
Phthiridium biarticulatum—Jaunbauere et al. (2008)
Myotis daubentonii (Kuhl) Basilia nana—Haitlinger (1978), Czuppon (2001), Czuppon & Molnár (2001).
Present study: see Table 1 for details.
Basilia nattereri—Soós (1955), Czuppon & Molnár (2001), Krištofík & Danko
(2012)
Nycteribia kolenatii—Theodor (1954), Theodor & Moscona (1954), Grulich &
Povolný (1955), Aellen (1963), Hůrka (1964), Beaucournu (1972), Kock (1973),
Nowosad (1974), Hůrka (1984), Hutson (1984), Müller & Ohlendorf (1984),
Nowosad (1987), Walter (1987), Nowosad (1990), Haitlinger & Ruprecht
(1992), Dietz & Walter (1995), Beaucournu & Noblet (1996), Kock (1999),
Rupp (1999), Czuppon (2001), Heddergott & Claußen (2004), Rupp et al.
(2004), Scheffler & Ressler (2005, 2007), Scheffler (2008, 2009), Lindecke &
Scheffler (2011), [Orlova (2011; Russia)], Krištofík & Danko (2012). Present
study: see Table 1 for details.
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TABLE 2. (Continued)
Bat species Bat fly species
Nycteribia latreillii—Colyer & Hammond (1951), Walter (2004)
Nycteribia pedicularia—Falcoz (1926) (doubtful record), Theodor (1954),
Jaunbauere et al. (2008)
Nycteribia schmidlii—Falcoz (1924, 1926), Theodor (1954), Theodor &
Moscona (1954), Soós (1955), Hutson (1984). Present study: see Table 1 for
details.
Nycteribia vexata—Hutson (1984)
Penicillidia dufourii*—Aellen (1963), Hutson (1984), Scheffler et al. (2013).
Present study: see Table 1 for details.
Penicillidia monoceros—Theodor (1954), Hůrka (1984), Hutson (1984), Müller
& Ohlendorf (1984), Nowosad (1987, 1990), Kock (1999), Rupp (1999),
Heddergott & Claußen (2004), Kock (2004), Rupp et al. (2004), Scheffler &
Ressler (2007), Scheffler (2008), Lindecke & Scheffler (2011)
Phthiridium biarticulatum—[Theodor & Moscona (1954; Morocco)], Theodor
(1967)
Myotis emarginatus (E. Geoffroy) Basilia italica—Theodor (1954), Hůrka (1980)
Basilia nana—Theodor & Moscona (1954), Theodor (1967), Beaucournu &
Noblet (1996)
Basilia nattereri—Dudich (1925)
Nycteribia kolenatii—Aellen (1963)
Nycteribia latreillii—Krištofík (1982)
Nycteribia pedicularia—Theodor & Moscona (1954)
Penicillidia dufourii—Theodor & Moscona (1954), Hůrka (1980), Imaz et al.
(1999)
Phthiridium biarticulatum—Theodor (1967)
Myotis escalerai Cabrera –
Myotis myotis (Borkhausen) Basilia italica—Theodor (1954), Hutson (1984)
Basilia nana—Theodor & Moscona (1954), Grulich & Povolný (1955), Aellen
(1963), Hůrka (1964, 1980), Krištofík (1982), Hůrka (1984), Hutson (1984),
Czuppon (2001), Czuppon & Molnár (2001), Güttinger et al. (2011)
Basilia nattereri—Soós (1955), Hůrka (1980)
Nycteribia kolenatii—Dudich (1925) (doubtful record), Hůrka (1964), Nowosad
(1974), Hutson (1984), Nowosad (1987), Czuppon (2001), Heddergott &
Claußen (2004), Rupp et al. (2004), Güttinger et al. (2011), Scheffler (2012)
Nycteribia latreillii—Theodor (1954), Theodor & Moscona (1954), Grulich &
Povolný (1955), Aellen (1963), Hůrka (1964), Kock (1973), Krištofík (1982),
Hůrka (1984), Hutson (1984), Nowosad (1990), Beaucournu & Noblet (1996),
Rupp (1999), Czuppon (2001), Rupp et al. (2004), Scheffler (2009), Uhrin et al.
(2010), Güttinger et al. (2011), Scheffler (2011b), Krištofík & Danko (2012),
Scheffler et al. (2013). Present study: see Table 1 for details.
Nycteribia pedicularia—Falcoz (1926) (doubtful record), Theodor (1954),
Theodor & Moscona (1954), Hutson (1984), Güttinger et al. (2011), Scheffler
(2011b), Scheffler et al. (2013)
Nycteribia schmidlii—Falcoz (1924), Dudich (1925), Falcoz (1926), Theodor
(1954), Theodor & Moscona (1954), Grulich & Povolný (1955), Soós (1955),
Aellen (1963), Hůrka (1964), Balcells (1968b), Hůrka (1980), Krištofík (1982),
Hutson (1984), Güttinger et al. (2011). Present study: see Table 1 for details.
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TABLE 2. (Continued)
Bat species Bat fly species
Nycteribia vexata—Falcoz (1926), Theodor (1954), Grulich & Povolný (1955),
Soós (1955), Aellen (1963), Hůrka (1964), Kock (1973), Krištofík (1982), Hůrka
(1984), Hutson (1984), Nowosad (1990), Beaucournu & Noblet (1996), Czuppon
(2001), Kock & Quetglas (2003), Uhrin et al. (2010), Güttinger et al. (2011),
Scheffler (2011b), Scheffler et al. (2013). Present study: see Table 1 for details.
Penicillidia conspicua—Dudich (1925), Falcoz (1926), Theodor (1954), Soós
(1955), Hůrka (1964), Hutson (1984), Güttinger et al. (2011), Scheffler (2011b)
Penicillidia dufourii—Dudich (1925), Falcoz (1926), Theodor (1954), Theodor
& Moscona (1954), Aellen (1955), Grulich & Povolný (1955), Soós (1955),
Aellen (1963), Hůrka (1964), Balcells (1968a), Kock (1973), Krištofík (1982),
Hůrka (1984), Hutson (1984), Noblet (1995), Beaucournu & Noblet (1996), Imaz
et al. (1999), Czuppon (2001), Kock & Quetglas (2003,as “P. dufourii dufourii”),
Güttinger et al. (2011), Scheffler (2011b), Krištofík & Danko (2012), Scheffler et
al. (2013). Present study: see Table 1 for details.
Penicillidia monoceros—Heddergott & Claußen (2004)
Phthiridium biarticulatum—Falcoz (1926), Theodor (1954), [Theodor &
Moscona (1954; Palestine)], Grulich & Povolný (1955), Soós (1955), Hůrka
(1964, 1980), Krištofík (1982), Hutson (1984), Güttinger et al. (2011)
Brachytarsina flavipennis [Streblidae]—Theodor (1954)
Myotis mystacinus (Kuhl) Basilia italica—Aellen (1963), Nowosad (1990), Beaucournu & Noblet (1996),
Czuppon (2001), Czuppon & Molnár (2001), Krištofík & Danko (2012),
Szentiványi et al. (2016)
Basilia nana—Theodor (1954), Theodor & Moscona (1954), Theodor (1967),
Kock (1973)
Basilia nattereri—Aellen (1963), Hůrka (1980)
Nycteribia kolenatii—Theodor (1954), Theodor & Moscona (1954), Aellen
(1963), Hůrka (1964), Walter (1987)
Nycteribia pedicularia—Theodor (1954)
Nycteribia schmidlii—Aellen (1963)
Nycteribia vexata—Aellen (1963), Kock (1999)
Penicillidia conspicua—Theodor (1954)
Penicillidia dufourii—Theodor & Moscona (1954) (doubtful record)
Penicillidia monoceros—Nowosad (1987)
Myotis nattereri (Kuhl) Basilia italica—Czuppon (2001), Czuppon & Molnár (2001)
Basilia nana—Theodor & Moscona (1954), Grulich & Povolný (1955), Hůrka
(1980, 1984), Hutson (1984), [Amr & Qumsiyeh (1993; Jordan)], Imaz et al.
(1999), Czuppon (2001), Czuppon & Molnár (2001), Topál (2011). Present
study: see Table 1 for details.
Basilia nattereri—Soós (1955), Aellen (1963), Hutson (1984), Masson (1989),
Imaz et al. (1999), Czuppon (2001), Czuppon & Molnár (2001), Topál (2011),
Krištofík & Danko (2012). Present study: see Table 1 for details.
Nycteribia kolenatii—Theodor (1954), Theodor & Moscona (1954), Hůrka (1964),
Kock (1973), Nowosad (1974), Walter & Benk (1982), Hutson (1984), Nowosad
(1987), Czuppon (2001), Scheffler & Ressler (2005, 2007), Topál (2011)
Nycteribia latreillii—Beaucournu (1961)
Nycteribia pedicularia—Theodor (1954)
Penicillidia dufourii—Theodor (1954), Hutson (1984), Topál (2011), Ševčík et
al. (2013)
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TABLE 2. (Continued)
Bat species Bat fly species
Penicillidia monoceros—Skuratowicz (1962), Hutson (1984), Topál (2011)
Myotis punicus Felten –
Nyctalus azoreum (Thomas) –
Nyctalus lasiopterus (Schreber) –
Nyctalus leisleri (Kuhl) Nycteribia kolenatii—Walter (2004)
Nycteribia latreillii—Kaňuch et al. (2005)
Nycteribia vexata—Kaňuch et al. (2005)
Nyctalus noctula (Schreber) Nycteribia kolenatii—Walter (1987, 2004)
Nycteribia pedicularia—Falcoz (1926) (doubtful record), Scott (1936), Theodor
(1954), Hutson (1984)
Penicillidia monoceros—Czuppon & Molnár (2001), Rupp et al. (2004)
Pipistrellus kuhlii (Kuhl) Basilia daganiae—Bequaert (1953), Theodor & Moscona (1954), Theodor
(1956, 1967), Kock (1974)
[Nycteribia schmidlii—Aktaş & Hasbenli (1994; Turkey)]
[Nycteribia vexata—Aktaş & Hasbenli (1994; Turkey)]
[Penicillidia conspicua—Aktaş & Hasbenli (1994; Turkey)]
Pipistrellus maderensis (Dobson) –
Pipistrellus nathusii (Keyserling &
Bläsius)
–
Pipistrellus pipistrellus (Schreber) [Basilia daganiae—Benda et al. (2010; Jordan)]
Basilia mediterranea—Beaucournu & Noblet (1985, 1994)
Basilia nana—Hutson (1984)
Nycteribia kolenatii—Theodor & Moscona (1954), Hutson (1984), Walter (1987, 2004)
Nycteribia schmidlii—Falcoz (1924, 1926), Theodor (1954), Soós (1955),
Hutson (1984), Nowosad et al. (1987)
Phthiridium biarticulatum—Colyer & Hammond (1951), Theodor (1967),
Hutson (1984)
Pipistrellus pygmaeus (Leach) Basilia mediterranea—Bauer (1957, as “P. pipistrellus mediterraneus”)
Plecotus auritus (Linnaeus) Basilia italica*—Present study: see Table 1 for details.
Basilia nana*—Theodor (1954), Theodor & Moscona (1954), Beaucournu
(1961), Walter (2004). Present study: see Table 1 for details.
Basilia nattereri—Balcells (1968b), Hůrka (1980)
Nycteribia kolenatii—Hůrka (1964), Nowosad (1974, 1987)
Nycteribia latreillii—Theodor & Moscona (1954)
Nycteribia schmidlii—Theodor (1967), Krištofík (1982)
Nycteribia vexata—Theodor (1954), Theodor & Moscona (1954)
Penicillidia conspicua—Theodor (1967)
Penicillidia dufourii—Theodor (1967)
Penicillidia monoceros—Theodor (1954)
Phthiridium biarticulatum—Falcoz (1926), Theodor (1954), Theodor &
Moscona (1954), Soós (1955)
Plecotus austriacus (Fischer) Basilia mongolensis nudior—Hůrka (1972)
Plecotus kolombatovici Dulic –
Plecotus macrobullaris Kuzjakin –
......continued on the next page
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EUROPEAN BAT FLY HOST ASSOCIATIONS
TABLE 2. (Continued)
Bat species Bat fly species
Plecotus sardus Mucedda, Kiefer,
Pidinchedda & Veith
–
Plecotus teneriffae Barrett–Hamilton –
Vespertilio murinus Linnaeus Nycteribia kolenatii—Theodor & Moscona (1954) (doubtful record), Hutson
(1984)
Nycteribia latreillii—Westwood (1835) (doubtful record), Theodor & Moscona
(1954) (doubtful record), Theodor (1967)
Nycteribia schmidlii—Theodor (1967)
Nycteribia vexata—Westwood (1835) (doubtful record), Kolenati (1857)
(doubtful record)
Penicillidia conspicua—Theodor (1967)
Penicillidia dufourii—Westwood (1835) (doubtful record), Theodor (1967)
Penicillidia monoceros—Hutson (1984)
Brachytarsina flavipennis [Streblidae]—Jobling (1934) (doubtful record)
Miniopteridae Dobson
Miniopterus schreibersii (Kuhl) Basilia nana—Aellen (1963)
Nycteribia latreillii—Grulich & Povolný (1955), Hůrka (1964, 1980), Krištofík
(1982)
Nycteribia pedicularia—Falcoz (1926) (doubtful record), Scott (1936), Theodor
(1954), Hůrka (1970)
Nycteribia schmidlii—Falcoz (1924), Dudich (1925), Falcoz (1926), Theodor
(1954), Grulich & Povolný (1955), Soós (1955), Aellen (1963), Hůrka (1964),
Theodor (1967), Balcells (1968b), Kock (1974), Krištofík (1982), Hůrka (1984),
Estrada-Peña & Serra-Cobo (1991), Beaucournu & Noblet (1996), Imaz et al.
(1999), Kock & Quetglas (2003), Krištofík & Danko (2012), Scheffler et al.
(2013), Ševčík et al. (2013). Present study: see Table 1 for details.
Nycteribia vexata—Falcoz (1926), Theodor (1954), Grulich & Povolný (1955),
Soós (1955), Krištofík (1982)
Penicillidia conspicua—Falcoz (1923), Dudich (1925), Falcoz (1926), Theodor
(1954), Grulich & Povolný (1955), Soós (1955), Aellen (1963), Hůrka (1964,
1970), Krištofík (1982), Hůrka (1984), Kock (1989), Beaucournu & Noblet
(1996), Krištofík & Danko (2012), Scheffler et al. (2013), Ševčík et al. (2013).
Present study: see Table 1 for details.
Penicillidia dufourii—Dudich (1925), Theodor (1954), Grulich & Povolný
(1955), Aellen (1963), Hůrka (1970, 1980), Krištofík (1982), Hůrka (1984),
Estrada-Peña & Serra-Cobo (1991), Beaucournu & Noblet (1996), Imaz et al.
(1999), Kock & Quetglas (2003, as “P. dufourii dufourii”), Krištofík & Danko
(2012), Scheffler et al. (2013). Present study: see Table 1 for details.
Phthiridium biarticulatum—Falcoz (1926), Theodor (1954), Theodor &
Moscona (1954), Soós (1955), Hůrka (1964, 1980)
Brachytarsina flavipennis [Streblidae]– Falcoz (1926), Theodor (1954),
[Theodor & Moscona (1954; Palestine)]
Molossidae Gervais in de Castelnau
Tadarida teniotis (Rafinesque) [Nycteribia pedicularia—Theodor & Moscona (1954; Palestine)]
Penicillidia dufourii—Scott (1936)
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Acknowledgements
We are grateful to Péter Paulovics (Szeged, Hungary), Krisztina Szőke (Department of Parasitology and Zoology,
Szent István University,Budapest, Hungary), Sándor Boldogh (Aggtelek National Park Directorate,Jósvafő,
Hungary), Péter Gombkötő (Bükk National Park Directorate, Eger, Hungary), Imre Dombi (Duna-Dráva National
Park Directorate,Pécs, Hungary) and Tamás Görföl (Department of Zoology, Hungarian Natural History Museum,
Budapest, Hungary) for their help in the collection of bat flies, and to Patrik Katona (Department of Evolutionary
Zoology and Human Biology, University of Debrecen, Debrecen, Hungary) for producing the map in Figure 1.
This study was supported by the National Talent Program (NTP-EFÖ-P-15-0234) of the Hungarian Ministry of
Human Resources, and co-financed by the European Social Fund in the framework of TÁMOP-4.2.4.A/ 2-11/
1-2012-0001 ‘National Excellence Program’. The suggestions made by Thomas Pape (Zoological Museum,
Natural History Museum of Denmark, University of Copenhagen, Copenhagen, Denmark), Danilo Russo
(Department of Agriculture, University of Naples “Federico II”, Naples, Italy and School of Biological Sciences,
University of Bristol, Bristol, United Kingdom), Daniel Whitmore (Department of Life Sciences, Natural History
Museum, London, United Kingdom) and an anonymous referee greatly improved the manuscript.
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