Bat fly – Penicillidia (Penicillidia) dufourii (Westwood, 1835) – Nycteribiidae – Cyprus
Text by George Konstantinou.
A wingless fly adapted to bats
Penicillidia dufourii is a small, wingless parasitic fly belonging to the family Nycteribiidae. Adults live on the external surface of bats and feed on their blood. Its unusual appearance, with a small head and long legs, resembles a miniature spider more than an ordinary fly. However, it is an insect, with three pairs of legs.
The family Nycteribiidae contains more than 270 described species worldwide. The genus Penicillidia comprises approximately 25 species, in addition to subspecies, according to the world checklist of the family. Nycteribiidae belong to the superfamily Hippoboscoidea and display particularly pronounced adaptations to parasitic life.
Morphology and identification
Penicillidia dufourii measures approximately 3.4–4 mm. Its body bears bristles and strong legs that facilitate movement through the host’s fur. The head is small and, at rest, folds back towards the thorax. The absence of wings is a permanent feature of adult Nycteribiidae.
Their body structure allows these flies to move in different directions through the fur. Adults remain on their host for most of their lives.
Reliable separation from related species, such as Penicillidia conspicua, requires examination of fine morphological characters. Identification keys use head shape, the arrangement of bristles and spines, abdominal plates and genital structures. The general appearance of a wingless fly on a bat is insufficient for species identification.
Distribution and occurrence in Cyprus
The nominate subspecies, Penicillidia dufourii dufourii, occurs in continental Europe, North Africa and southwestern Asia, extending towards the western Himalayas and eastern Kazakhstan.
Its presence in Cyprus was documented in the study by Ševčík, Benda and Lučan, published in 2013. Near Kakopetria, two females were recorded from Myotis blythii on 15 October 2005, and one male from a bat identified as Myotis nattereri on 13 October 2005. The specimens came from abandoned mine galleries in the Troodos Mountains.
These constituted the first published records of the species from Cyprus. A first scientific record establishes documented presence, rather than the time at which a species arrived on the island.
Hosts and ecological relationships
The principal hosts across the species’ wider distribution are Myotis myotis and Myotis blythii. It has also been recorded from other bats, including Miniopterus schreibersii. The specific Cypriot hosts in the 2013 study were M. blythii and M. nattereri.
Shared roosts used by different bat species create opportunities for parasites to transfer between hosts. P. dufourii shows a preference for bats of the genus Myotis, but is not restricted exclusively to a single species.
Research published in 2024 on bat roosts showed that season and host-community composition can influence recorded parasitic relationships. Finding a fly on a particular host does not necessarily mean that this is its principal host throughout its distribution.
Further information about the hosts is available in the list of bats of Cyprus.
Reproduction and life cycle
Nycteribiidae have a distinctive reproductive strategy known as adenotrophic viviparity. A female retains one developing larva inside her body and nourishes it with secretions from specialised glands, known as “milk glands”. The larva completes its development there up to the third instar.
Shortly before deposition, the female temporarily leaves the bat and deposits the mature larva on a surface within the roost. The larva attaches to the substrate and pupates rapidly. The female returns to her host, while the new adult searches for a bat after emerging from its puparium.
Thus, larval development takes place inside the mother, pupation occurs in the roost, and adult life is spent mainly on the host. Development times depend on species and environmental conditions and do not provide a fixed timetable for P. dufourii in Cyprus.
Habitat and dispersal
The habitat includes both the bats’ bodies and their roosts. The Cypriot records from mine galleries demonstrate the species’ association with underground spaces used by its hosts.
Because adults cannot fly, transport over longer distances is associated with bat movements. Within colonies, proximity between animals and shared use of the roost facilitate contact with new hosts. Suitable bats and pupation sites are therefore essential for maintaining the life cycle.
Parasites on the parasite itself
Penicillidia dufourii can host microscopic fungi belonging to the order Laboulbeniales. European literature has reported Arthrorhynchus nycteribiae and A. eucampsipodae on this fly.
This is a characteristic example of hyperparasitism: the fly parasitises the bat, and the fungus parasitises the fly. Studies have shown that P. dufourii is a secondary host of A. nycteribiae compared with P. conspicua. These European observations do not constitute records of the fungi from Cyprus.
Modern scientific research
The complete mitochondrial genome of Penicillidia dufourii was published in 2024. It contains 15,354 base pairs and 37 genes. The analysis supported its placement within Nycteribiidae and provided molecular data for comparisons with related species.
Combining morphology and genetic information helps identify these specialised parasites and understand their relationships with bats.
Taxonomy
Kingdom: Animalia
Phylum: Arthropoda
Class: Insecta
Order: Diptera
Superfamily: Hippoboscoidea
Family: Nycteribiidae — more than 270 described species
Subfamily: Nycteribiinae
Genus: Penicillidia Kolenati, 1863 — approximately 25 species, in addition to subspecies
Subgenus: Penicillidia
Species: Penicillidia dufourii (Westwood, 1835)
Full name including subgenus: Penicillidia (Penicillidia) dufourii (Westwood, 1835)
Original combination: Nycteribia dufourii Westwood, 1835
Synonym: Penicillidia frauenfeldi Kolenati, 1856
Status in Cyprus: confirmed by collected scientific specimens.
Sources
Ševčík, Benda & Lučan, 2013 — Diptera Pupipara from bats of Crete and Cyprus. journals.tubitak.gov.tr
Updated Taxonomic Key of European Nycteribiidae, 2023. MDPI
Checklist of World Nycteribiidae. researchgate.net
Checklist of the Diptera of the Netherlands — Nycteribiidae morphology and biology. Nycteribiidae
Szentiványi and colleagues, 2019 — Bat Flies and Their Microparasites. frontiersin.org
Season and host-community composition inside roosts may affect host-specificity of bat flies, 2024. Scientific Reports
Laboulbeniales infection of bat flies across Europe, 2018. Springer Nature Link
Focus on Hyperparasites, 2022 — fungi and parasitic bat flies. frontiersin.org
Complete mitochondrial genome of Penicillidia dufourii, 2024. pubmed.ncbi.nlm.nih.gov
Catalogus dipterorum Germaniae: Nycteribiidae, 2025.
Diptera
Pupipara from bats of two large eastern Mediterranean islands, Crete and Cyprus
Martin
ŠEVČÍK1,*, Petr BENDA2,3, Radek Kris LUČAN3
1
Department of Zoology and Anthropology, Faculty of
Natural Sciences, Constantine the Philosopher University in Nitra,
SK-949 74 Nitra, Slovakia
2
Department of
Zoology, National Museum, CZ-115 79 Prague 1, Czech Republic 3
Department of Zoology, Faculty of Science, Charles
University, CZ-128 44 Prague 2, Czech Republic
Received:
15.12.2011 Accepted: 19.09.2012 Published Online: 24.12.2012 Printed: 21.01.2013
Abstract: This paper presents a review of records of bat flies
(families Nycteribiidae and Streblidae) from Crete and Cyprus, based on new
findings and published data. Seven species representing 4 genera belonging to
the family Nycteribiidae and 1 species from the family Streblidae from 5 bat
species in Crete are reported, and 7 Nycteribiidae species (4 genera) and 1
Streblidae species from 9 bat species in Cyprus are reported. The bat flies Nycteribia latreillii (Leach, 1817), Basilia mongolensis nudior Hůrka 1972, Penicillidia conspicua Speiser, 1900, P. dufourii Westwood, 1835, and Brachytarsina flavipennis Macquart, 1851
are reported from Crete for the first time, and Phthiridium integrum (Theodor and Moscona, 1954), P. dufourii, and B. flavipennis are reported from Cyprus for the first time. Key words: Nycteribiidae, Streblidae,
first records, distribution, hosts
1. Introduction
The family Nycteribiidae Samouelle, 1819
comprises the dipterans parasitising on bats of all zoogeographical regions.
According to the checklist by Graciolli and Dick (2008), more than 270 species
occur in the world. Seventeen species (5 genera) are known from the western
Palaearctic (Theodor, 1967; Hůrka and Soós, 1986a), and 14 species (5 genera)
of them occur in the Mediterranean area (Hůrka, 1964; Theodor, 1967).
No detailed information on Nycteribiidae has been available
from Crete, except for several records of 3 species reported by Theodor (1967)
and Kock (1974, 1989): Nycteribia
pedicularia Latreille, 1805 (collected from Myotis capaccinii (Bonaparte, 1837)), Nycteribia schmidlii Schiner, 1853 (from M. capaccinii and Miniopterus
schreibersii (Kuhl, 1817)), and Phthiridium
biarticulatum Hermann, 1804 (from Rhinolophus
ferrumequinum (Schreber, 1774)). A similar situation holds for the
nycteribiid fauna of Cyprus: only scarce data have been available from this
island, including records of 5 species, Nycteribia
pedicularia (from M. capaccinii),
N. schmidlii (from Rhinolophus euryale Blasius, 1853), Phthiridium biarticulatum (from R. euryale), Basilia daganiae Theodor and Moscona, 1954 (from Pipistrellus kuhlii (Kuhl, 1817)), and Penicilidia conspicua Speiser, 1900
(from M. schreibersii) (Bequaert,
1953; Theodor, 1956, 1967; Kock, 1974, 1984, 1989).
* Correspondence:
martin.sevcik@hotmail.sk
The family Streblidae Kolenati, 1863 is represented by 4
species of 3 genera in the Mediterranean (Jobling, 1939; Hůrka and Soós,
1986b). Just one species, Brachytarsina
flavipennis Macquart, 1851, occurs in Europe (Hůrka, 1972). So far, no
record of this species has been available either from Crete or from Cyprus.
2. Materials and methods
In October 2005, July and October 2006, and
May and June 2008, bats were collected from Crete and Cyprus in the course of
research trips (Benda et al., 2007, 2009). Mist nets were installed according
to standard procedures (Kunz and Kurta, 1988) in front of water, and bats were
also captured in their roosts (caves, mines). Each captured host was placed
into a separate cloth bag and labelled. Hosts were inspected for bat flies,
ectoparasites living in the fur of the body, which were collected with
entomological tweezers and preserved in 75% ethanol. Samples for a species of
host were stored in an Eppendorf tube. The age and sexual status (adult male (♂
ad), adult female (♀ ad), subadult female (♀
sad)) of the host is given for most samples, with the exception of the period
of October 2010, when only the number and species of host were registered. All
bat flies were observed freely under a magnifying glass, and keys were used for
the determination and comparison of parasitic species: Theodor and Moscona
(1954), Theodor (1967, 1975), and Hůrka (1972). Voucher specimens of the
parasites are deposited in the collection of the first author (cMŠ). The taxa
are arranged alphabetically. The information for each species appears in the
following format: synonyms available from the respective islands; published
data; new material examined (number and developmental stage of specimens,
locality, date, number of collection, age/number of host).
3. Results and discussion
In total, 6 species representing 4 genera
belonging to the family Nycteribiidae and 1 species of the family Streblidae
were collected from Crete, and 4 species of 2 genera of the family
Nycteribiidae and 1 species of Streblidae were collected from Cyprus.
3.1. Family Nycteribiidae (Samouelle 1819)
3.1.1. Subfamily Nycteribiinae Samouelle, 1819,
sensu Maa 1965
3.1.1.1.
Nycteribia (Nycteribia) latreillii
(Leach, 1817) Crete. New material examined: 1♂, 1♀,
Lefkogeia, at Venetian bridge ca. 3 km E of Kato Preveli Monastery, 30 May 2008
(cMŠ 3962-3963, leg. P. Benda and V. Hanák) from 2♀♀ ad
Myotis blythii.
Remarks: Until Theodor (1954a), the species was reported in
Europe under various names, as the proper name was erroneously used for N. kolenati Theodor and Moscona, 1954.
The distribution of N. latreillii
corresponds to the ranges of its primary host species, Myotis myotis (Borkhausen, 1797), M. blythii (Tomes, 1857), and M.
punicus Felten, 1977, i.e. almost all of continental Europe (northern limit
of 51°N–52°N), the Maghreb, the Middle East, and Central Asia (Hůrka, 1964;
Theodor, 1967). In addition, it can also occur on other species of the genus Myotis Kaup, 1829, Miniopterus Bonaparte, 1837, and Rhinolophus Lacépède, 1799, as well as on other cavedwelling bats
(see Theodor, 1967). Our findings represent the first evidence of this species
from Crete. Although we did not find this bat fly in Cyprus, its occurrence is
to be expected due the presence of one of the principal host species, M. blythii (Benda et al., 2007).
3.1.1.2.
Nycteribia (Nycteribia) pedicularia
Latreille, 1805 Crete. Published records: Kock (1974): Piskokefalo at Sitia
(host Myotis capaccinii).
Cyprus. Published
records: Kock (1974): Kyrenia (host Myotis
capaccinii).
Remarks: Nycteribia
pedicularia has frequently been confused with N. latreillii and/or N.
kolenati Theodor and Moscona, 1954. N.
pedicularia is distributed in south-eastern Europe, and the northern limit
of its range probably reaches the Alps and southern Carpathians (Hůrka, 1962).
In the broader scale of the western Palaearctic, its distribution range
encompasses Turkey, Palestine, Iran, Tunisia, and Algeria (Theodor and Moscona,
1954; Theodor, 1967). However, the record from Cyprus is doubtful, as the
occurrence of the host species as well as the record itself is in doubt (see
Benda et al., 2007). Host species of this bat fly are cave-dwelling bats of the
genera Myotis, Miniopterus, and Rhinolophus
(Theodor and Moscona, 1954).
3.1.1.3. Nycteribia (Listropoda) s.
schmidlii Schiner, 1853
Crete.
Published records: Theodor (1967): Limnea (host Miniopterus schreibersii). - Kock (1974): Canea = Chaniá (host Miniopterus schreibersii), Piskokefalo
bei Sitia (host Myotis capaccinii).
New material examined: 2♂♂, 1♀,
Dramia, Petres river bridge, 6 October 2006 (cMŠ 3400, leg. P. Benda and V.
Hanák) from 1♀ ad Myotis
capaccinii; 5♂♂, 9♀♀, Argyroupoli, Mouselas
river, 3 June 2008 (cMŠ 3973-3974, leg. P. Benda and V. Hanák) from 2♀♀
ad Myotis capaccinii; 3♂♂,
3♀♀,
Omalos Plateau, Tzani cave, 1 October 2006 (cMŠ 3360-3375, leg. P. Benda and V.
Hanák) from collection 13♂♂ ad and 2♀♀
sad Miniopterus schreibersii.
Cyprus.
Published records: Theodor (1967): site
unspecified (host Rhinolophus euryale).
Remarks: The distribution of the western subspecies N. s. schmidlii corresponds to the
distributional range of its principal host bat, Miniopterus schreibersii, viz. central and southern Europe, the
Middle East, Afghanistan, and North Africa.
3.1.1.4.
Phthiridium biarticulatum Hermann, 1804 Synonym: Stylidia biarticulata Hermann, 1804: Theodor (1967: 127).
Crete. Published
records: Theodor (1967): Gonia = Mount Gonias (host Rhinolophus ferrumequinum). - Kock (1989): Rethimnon (host Rhinolophus ferrumequinum).
New material examined: 1♂, Kolymvari, mine above
village, 9 October 2006 (cMŠ 3426, leg. P. Benda and V. Hanák) from 1♂
ad Rhinolophus ferrumequinum; 2♀♀,
Omalos Plateau, Tzani cave, 1 October 2006 (cMŠ 3377-3378, leg. P. Benda and V.
Hanák) from 2♂♂ ad Rhinolophus
ferrumequinum.
Cyprus.
Published records: Theodor (1967): site
undefined (host Rhinolophus euryale).
New material examined: 1♀, Kakopetria, Troodos
Forest, abandoned mine 5 km to SW, upper gallery, 13 October 2005 (cMŠ c07,
leg. R.K. Lučan) from Rhinolophus ferrumequinum
individual of unknown sex and age; 1♂, Afendrika, ruins of
Panagia Chrysiotissa church and Asomatos church, 17 October 2005 (cMŠ c02, leg.
R.K. Lučan) from 1♂ ad Rhinolophus
ferrumequinum (Schreber, 1774); 3♂♂, 6♀♀,
Kakopetria, Troodos Forest, abandoned mine 5 km to SW, upper gallery, 27 July
2006
(cMŠ 3335, 3336, leg. P. Benda) from 1♀
ad and 1♀ sad Rhinolophus
blasii (Peters, 1866).
Remarks: This cave-dwelling bat fly lives on representatives
of the genus Rhinolophus. Its
distribution range is circum-Mediterranean and reaches Tajikistan and
Kyrgyzstan (Hůrka, 1964). While Rhinolophus
ferrumequinum is the most frequent host of this bat fly (Hůrka, 1964),
other (mostly cave-dwelling) bats, e.g., Miniopterus
schreibersii, Myotis myotis and M. blythii, are less-parasitised hosts
(Hůrka, 1964).
3.1.1.5. Phthiridium integrum
(Theodor and Moscona, 1954)
Cyprus.
New material examined: 3♀♀, Kantara, Kantara castle
ruins 3 km to NE, 25 July 2006 (cMŠ 3328, leg. P. Benda) from 1♂
ad Eptesicus anatolicus.
Remarks: This species was described by Speiser (1901) and
redescribed by Karaman (1939) based on a record from Egypt. Findings are known
from host species of the genera Taphozous
Geoffroy, 1818, Rhinolophus and Hipposideros Gray, 1831 from the Middle
East: Egypt, Palestine, Jordan, Saudi Arabia, and Yemen (Sanborn and
Hoogstraal, 1953; Theodor and Moscona, 1954; Theodor, 1976; Kock and Nader,
1979; Amr and Qumsiyeh, 1993). This species is reported here for the first time
from Cyprus and from a new host, Eptesicus
anatolicus Felten, 1971.
3.1.1.6.
Basilia (Basilia) daganiae
Theodor and Moscona, 1954
Cyprus.
Published records: Bequaert (1953): Limassol (host Pipistrellus kuhlii). - Theodor (1956): Limassol, (host Pipistrellus kuhlii), site undefined,
leg. Lilford (without host). - Theodor (1967): site undefined, leg. Lilford
(host Pipistrellus kuhlii). - Kock
(1974): Larnaka (host Pipistrellus kuhlii).
Remarks: The distribution range of this species from the “bathybothyra group” covers the whole
eastern Mediterranean: Cyprus, Turkey, Syria, Jordan, Egypt, Lebanon, and
Palestine (Theodor and Moscona, 1954; Lewis and Harrison, 1962; Hůrka and Soós,
1986a; Benda et al., 2010). Its occurrence outside the eastern Mediterranean
region is not excluded, considering the much broader distribution of its main
host Pipistrellus kuhlii. B. daganiae has also been recorded from
other species of the genus Pipistrellus:
P. pipistrellus (Schreber, 1774) from
Jordan (Benda et al., 2010).
3.1.1.7.
Basilia (Basilia) mongolensis nudior Hůrka, 1972 Crete. New
material examined: 2♀♀, Meskla, above creek, 5
October 2006 (cMŠ 3393, leg. P. Benda and V.
Hanák) from 1♂ ad Myotis aurascens; 1♂, 1♀,
Katholiko monastery, Agiou Ioanni cave, 27 May 2008 (cMŠ 3958, leg. P. Benda)
from 1♀ ad Myotis
aurascens.
Remarks: This subspecies from the “bathybothyra group” was described by Hůrka (1972) from the host Plecotus austriacus Fischer, 1829. The
full expansion of this subspecies is unknown, and we expect it to be in the
same area distribution as the host Myotis
nattereri (Kuhl, 1817). Its occurrence on other arboreal host species is
not excluded, but it occurs mostly in the “mystacinus
group”, viz. M. alcathoe von
Helversen and Heller, 2001 (Heddergott, 2009), and in our records from M. aurascens Kusjakin, 1935. Previous
records are from Croatia, Greece (Hůrka, 1972), Bulgaria (Hůrka, 1984), Austria
(Kock, 1984), Turkey (Aktaş and Hasbenli, 1994; Hasbenli, 1997), and Germany
(Heddergott, 2009).
3.1.1.8. Penicillidia (Neopenicillidia)
conspicua Speiser, 1901
Crete.
New material examined: 3♂♂, 3♀♀,
Omalos Plateau, Tzani cave, 1 October 2006 (cMŠ 3360-3375, leg. P. Benda and V.
Hanák) from collection 13♂♂ ad and 2♀♀
sad Miniopterus schreibersii; 2♀♀,
Milatos, Milatou cave, 7 October 2006 (cMŠ 3422-3425, leg. P. Benda and V.
Hanák) from collection 1♀ ad and 3♀♀
sad Miniopterus schreibersii.
Cyprus. Published
records: Kock (1989): Emba, Paphos Distr. (host Miniopterus schreibersii).
New material examined: 1♂, 1♀,
Neo Horio, Smigies Trail, ca. 3 km to NW, abandoned chromite mine system
‘Magnesia Mine’, 12 October 2005 (cMŠ c03, c04, leg. R.K. Lučan) from 2 Miniopterus schreibersii individuals of
unknown sex and age.
Remarks: Penicillidia
conspicua is a species bound to
its principal host, Miniopterus
schreibersii, and its distribution range corresponds to the occurrence of
this bat (Hůrka, 1964). Within this range, it has frequently been found also on
Myotis myotis and other cave-dwelling
bat species (Hůrka, 1964). This parasite species is reported from Crete for the
first time here.
3.1.1.9.
Penicillidia (Penicillidia) dufourii
Westwood, 1835 Crete. New material examined: 2♂♂, 3♀♀,
Lefkogeia, at Venetian bridge ca. 3 km E of Kato Preveli Monastery, 30 May 2008
(cMŠ 3962-3963, leg. P. Benda and V. Hanák) from 2♀♀ ad
Myotis blythii; 1♂,
1♀,
Dramia, Petres river bridge, 6 October 2006 (cMŠ 3400, leg. P. Benda and V.
Hanák) from 1♀ ad Myotis
capaccinii; 2♂♂, 2♀♀, Argyroupoli, Mouselas
river, 3 June 2008 (cMŠ 39733974, leg. P. Benda and V. Hanák) from 2♀♀
ad Myotis capaccinii.
Cyprus. New
material examined: 2♀♀, Kakopetria, Troodos
Forest, abandoned mine 4 km to SW, lower gallery, 15 October 2005 (cMŠ c05,
leg. R.K. Lučan) from 1♂ ad Myotis blythii; 1♂, Kakopetria, Troodos
Forest, abandoned mine 5 km to SW, upper gallery, 13 October 2005 (cMŠ c06,
leg. R.K. Lučan) from Myotis nattereri
individual of unknown sex and age.
Remarks: The subspecies P.
d. dufourii is distributed over continental Europe, North Africa, and
south-western Asia to the western Himalayas and eastern Kazakhstan. Myotis myotis and M. blythii are the main hosts of this species, but it can also be
found on Miniopterus schreibersii,
though much more rarely. This parasite species is recorded here for the first
time from Crete and Cyprus.
3.2. Family Streblidae Kolenati, 1863
3.2.1. Subfamily Brachytarsininae (Speiser,
1900)
3.2.1.1. Brachytarsina (Brachytarsina)
flavipennis Macquart, 1851
Crete.
New material examined: 1♂, Patsos, Agiou Antoniou
cave, 3 October 2006 (cMŠ 3388, leg. P. Benda and V. Hanák) from 1♀
sad Rhinolophus ferrumequinum; 2♀♀,
Katholiko monastery, Agiou Ioanni cave, 27 May 2008 (cMŠ 3958, leg. P. Benda)
from 1♀ ad Myotis
aurascens.
Cyprus. New
material examined: 1♀, Kakopetria, Troodos
Forest, abandoned mine 5 km to SW, upper gallery, 13 October 2005 (cMŠ 08, leg.
R.K. Lučan) from Rhinolophus
ferrumequinum individual of unknown sex and age; 1♂, 2♀♀,
Kakopetria, Troodos Forest, abandoned mine 5 km to SW, upper gallery, 27 July
2006 (cMŠ 3336, leg. P. Benda) from 1♀ sad Rhinolophus blasii.
Remarks: B.
flavipennis is the only streblid
species occurring in the Mediterranean portion of the Palaearctic. This species
has a circum-Mediterranean distribution range, and in Europe it reaches the
latitude of 40°N–43° N, roughly corresponding to the areas of mean January
temperatures of 8 to 10 °C (Jobling, 1951;
Hůrka, 1972).
Jobling (1934, 1939) indicated the occurrence of this
species in Cyprus and Theodor’s record (1954b) is uncertain, since a direct
finding has not been reported. This bat fly is recorded here from both large
islands for the first time.
In conclusion, the combination of our new data with
previously published information has resulted in an increase in the total
number of species of the Nycteribiidae family on Crete from 3 to 7 species, and
from 5 to 7 species on Cyprus. In addition to the above discussion, we may
expect other species from this family (genera Nycteribia Latreille, 1796, Basilia
Miranda-Ribeiro, 1903, and Eucampsipoda Kolenati,
1857) on the individual islands. Nycteribia
(Acrocholidia) vexata Westwood, 1835 is a species with a relation to the extension
of the principal hosts Myotis myotis
and M. blythii (as N. latreilii), whereas the second host
was confirmed on the island of Crete (Benda et al., 2008). The principal host
of Basilia nana Theodor, 1954, in
Middle Eastern countries is M. nattereri (Turkey
- Aktaş and Hasbenli, 1994; Hasbenli, 1997; Jordan - Amr and Qumsiyeh, 1993;
Benda et al., 2010), and on this basis we can expect this parasitic species to
confirm this host from Cyprus (Benda et al., 2007). In addition, on the same
island there probably occurs the parasitic species of the host Rousettus aegyptiacus (Geoffroy, 1810),
viz. Eucampsipoda aegyptia (Macquart,
1851). The occurrence of this species on island has already been considered by
Theodor (1952, 1954a, 1955; see also Kock and Nader, 1979). This host species
has been confirmed on Cyprus by
several authors (summary in Benda et al., 2011).
The fauna of Streblidae is represented by a single species,
which was confirmed for the first time on both islands.
Acknowledgements
This research was supported by grants from
the Czech Science Foundation (#206/09/0888), the Ministry of Culture of the
Czech Republic (#DKRVO 00023272), and the University Grant Agency of the
Constantine of Philosopher University in Nitra, Slovakia (VII/36/2011).
Appendix
Appendix I. Gazetteer.
Only the originally recorded sites are
mentioned.
Crete.
Prefecture of Chania: Argyroupoli [Αργυρούπολη], Mouselas river, old
building, 35°19′N,
24°20′E, 105 m a.
s. l.; - Katholiko Monastery [Μονή
Καθολικού], Agiou Ioanni cave, 35°35′N,
24°09′E, 80 m a. s.
l.; - Kolymvari [Κολυμβάρι], mine, 35°33′N,
23°46′E, 44 m a. s.
l.; - Meskla [Μεσκλά], creek, 35°24′N,
23°57′E, 206 m a.
s. l.; - Omalos Plateau [Οροπέδιο Ομαλού], Tzani cave, 35°21′N, 23°54′E, 1025 m a. s. l.; Prefecture
of Rethymno: Dramia [Δράμια], Petre river bridge, 35°21′N, 24°21′E, 2 m a. s. l. - Lefkogeia
[Λευκόγεια], Venetian bridge ca. 3 km E of Kato Preveli Monastery, 35°10′N, 24°28′E, 45 m a. s. l.; - Patsos
[Πατσός], Agiou Antoniou cave, 35°15′N,
24°34′E, 200 m a.
s. l.; Prefecture of Lasithi: Milatos [Μίλατος], Milatou cave, 35°18′N, 25°35′E, 129 m a. s. l.
Cyprus. Northern
Cyprus: Afendrika (= Panagia Afentrika [Παναγια Αφεντρικα]), ruins, İskele
District, 35°39′N,
34°26′E, ca. 25 m
a. s. l.; - Kantara (= Kantara [Κανταρα]), Kantara castle ruins 3 km to NE,
İskele District, 35°24′N,
33°55′E, ca. 625 m
a. s. l.; - Southern Cyprus: Kakopetria [Κακοπετρια], Troodos Forest, mine 5 km
to SW (upper gallery), Lefkosia District, 34°57′N,
32°52′E, ca. 1665 m
a. s. l.; - Kakopetria [Κακοπετρια], Troodos Forest, mine 4 km to SW (lower
gallery), Lefkosia District, 34°58′N,
32°52′E, ca. 1305 m
a. s. l.; - Neo Horio [Νεο Χωριο], Smigies Trail ca. 3 km to NW, abandoned
chromite mine system ‘Magnesia Mine’, Akamas Peninsula, Pafos District, 35°03′N, 32°20′E, ca. 305 m a. s. l.
Country / Family / Subfamily / Genus / Species
Country:
CRETE
Family:
NYCTERIBIIDAE Samouelle, 1819
Subfamily:
Nycteribiinae Samouelle, 1819
Genus: Nycteribia Latreille,
1796
|
Species of
host
|
Nycteribia latreilii (Leach, 1817)*
|
Myotis blythii (Tomes, 1857)**
|
Nycteribia pedicularia Latreille, 1805
|
Myotis capaccinii (Bonaparte, 1837)
|
Nycteribia
schmidlii schmidlii Schiner, 1853
Genus: Phthiridium Hermann,
1804
|
Myotis capaccinii (Bonaparte, 1837) Miniopterus schreibersii (Kuhl, 1817)
|
Phthiridium biarticulatum Hermann,
1804 Genus: Basilia Miranda-Ribeiro, 1903
|
Rhinolophus ferrumequinum (Schreber, 1774)
|
Basilia mongolensis nudior Hůrka,
1972* Genus: Penicillidia Kolenati, 1863
|
Myotis aurascens Kusjakin, 1935**
|
Penicillidia conspicua Speiser, 1901*
|
Miniopterus schreibersii (Kuhl, 1817)**
|
Penicillidia
dufourii Westwood, 1835*
Family:
STREBLIDAE Kolenati, 1683
Subfamily:
Brachytarsininae (Speiser, 1900)
Genus: Brachytarsina
Macquart, 1851
|
Myotis
blythii (Tomes, 1857)**
Myotis capaccinii (Bonaparte, 1837)**
|
Brachytarsina
flavipennis Macquart, 1851*
Country:
CYPRUS
Family:
NYCTERIBIIDAE Samouelle, 1819
Subfamily:
Nycteribiinae Samouelle, 1819
Genus: Nycteribia Latreille,
1796
|
Rhinolophus ferrumequinum (Schreber, 1774)** Myotis aurascens Kusjakin, 1935**
|
Nycteribia pedicularia Latreille, 1805
|
Myotis capaccinii (Bonaparte, 1837)
|
Nycteribia schmidlii schmidlii Schiner,
1853 Genus: Phthiridium Hermann, 1804
|
Rhinolophus euryale Blasius, 1853
|
Phthiridium biarticulatum Hermann, 1804
|
Rhinolophus
blasii Peters, 1867**
Rhinolophus
euryale Blasius, 1853
Rhinolophus
ferrumequinum (Schreber, 1774)**
|
Phthiridium integrum (Theodor
and Moscona, 1954)* Genus: Basilia Miranda-Ribeiro, 1903
|
Eptesicus anatolicus Felten, 1971**
|
Basilia daganiae Theodor
and Moscona, 1954 Genus: Penicillidia Kolenati, 1863
|
Pipistrellus kuhlii (Kuhl, 1817)
|
Penicillidia conspicua Speiser, 1901
|
Miniopterus schreibersii (Kuhl, 1817)
|
Penicillidia
dufourii Westwood, 1835*
Family:
STREBLIDAE Kolenati, 1683
Subfamily:
Brachytarsininae (Speiser, 1900)
Genus: Brachytarsina
Macquart, 1851
|
Myotis blythii (Tomes, 1857)**
Myotis nattereri (Kuhl, 1817)**
|
Brachytarsina flavipennis Macquart, 1851*
|
Rhinolophus blasii Peters, 1867**
|
|
Rhinolophus
ferrumequinum (Schreber, 1774)**
|
Appendix
II. Checklist of Nycteribiidae and Streblidae with their hosts from the islands
of Crete and Cyprus. Checklist of 7 species, with 4 genera of bat flies Nycteribiidae and 1 species of
Streblidae known from Crete and the same number of species and genera from
both parasitic families documented from
Cyprus, based on both literature data and new records obtained during recent
field studies. The first record of the parasite (*) and the first record from the host (**) of the respective island are
indicated.
References
Amr, Z.S.
and Qumsiyeh, M.B. 1993. Records of bat flies from Jordan, Libya and Algeria.
Entomol. News 104: 43–46.
Aktaş, M.
and Hasbenli, A. 1994. Bat flies of Eastern Turkey (the east of
Samsun-İskenderun Line) (Diptera: Nycteribiidae). Gazi Üniv. Fen Bil. Enst.
Derg. 7: 48–51.
Benda, P.,
Abi-Said, M., Bartonička, T., Bilgin, R., Faizolahi, K., Lučan, R.K., Nikolaou,
H., Reiter, A., Shohdi, W.M., Uhrin, M. and Horáček, I. 2011. Rousettus aegyptiacus (Pteropodidae) in
the Palaearctic: list of records and revision of the distribution range.
Vespertilio 15: 3–36.
Benda, P.,
Georgiakakis, P., Dietz, C., Hanák, V., Galanaki, K., Markantonatou, V.,
Chudárková, A., Hulva, P. and Horáček, I. 2009. Bats (Mammalia: Chiroptera) of
the Eastern Mediterranean and Middle East. Part 7. The bat fauna of Crete,
Greece. Acta Soc. Zool. Bohem. 72: 105–190.
Benda, P., Hanák, V.,
Horáček, I., Hulva, P., Lučan, R. and Ruedi,
M. 2007. Bats (Mammalia:
Chiroptera) of the Eastern Mediterranean. Part 5. Bat fauna of Cyprus: review
of records with confirmation of 6 species new for the island and description of
a new subspecies. Acta Soc. Zool. Bohem. 71: 71–130.
Benda, P.,
Lučan, R.K., Obuch, J., Reiter, A., Andreas, M., Bačkor, P., Bohnenstengel, T.,
Eid, E.K., Ševčík, M., Vallo, P. and Amr, Z.S. 2010. Bats (Mammalia:
Chiroptera) of the Eastern Mediterranean and Middle East. Part 8. Bats of
Jordan: fauna, ecology, echolocation, ectoparasites. Acta Soc. Zool. Bohem. 74:
185–353.
Bequaert,
J.G. 1953. Exploration du Parc National Albert. Mission G.F. de Witte
(1933-1935). Hippoboscidae and Nycteribiidae. Fasc. 79(7): 89–94. Inst. Parcs
Nation. Congo Belge, Bruxelles.
Graciolli,
G. and Dick, C.W. 2008. Checklist of World Nycteribiidae (Diptera:
Hippoboscoidea). Available at
http://fm1.fieldmuseum.org/aa/Files/cdick/Nycteribiidae_ Checklist_1jul08.pdf
(accessed November 2011).
Hasbenli, A.
1997. Contributions to bat flies of Turkey (Diptera: Nycteribiidae,
Streblidae). Gazi Üniv. Fen Bil. Enst. Derg. 10(4): 533–544.
Heddergott,
M. 2009. Erstnachweis der Fledermausfliege Basilia
mongolensis nudior Hůrka, 1972 in Deutschland (Diptera: Nycteribiidae).
Studia dipterologica 15(1–2): 301–304.
Hůrka, K.
1962. Beitrag zur Nycteribien- und Streblidenfauna Albaniens nebst Bemerkungen
zur Fauna von Bulgarien, Ungarn und UdSSR. Čas. Českoslov. Spol. Entomol. 59:
156– 164.
Hůrka, K.
1964. Distribution, bionomy and ecology of the European bat flies with special
regard to the Czechoslovak fauna (Diptera, Nycteribidae). Acta Univ. Carol.-
Biol. 1964(3): 167–231.
Hůrka, K.
1972. Basilia mongolensis nudior
subsp. n. nebst Bemerkungen zur Nycteribien- und Streblidenfauna Thrakiens
(Diptera: Pupipara). Ann. Naturhist. Mus. Wien 76: 709–713.
Hůrka, K.
1984. New taxa and new records of Palearctic Nycteribiidae and Streblidae
(Diptera, Pupipara). Acta Soc. Zool. Bohemoslov. 48: 90–101.
Hůrka, K.
and Soós, Á. 1986a. Nycteribiidae. In: Catalogue of Palaearctic Diptera. Volume
11 (Eds. Á. Soós and L. Papp). Elsevier Science Publishers, Amsterdam, and
Akadémiai Kiadó, Budapest, Hungary, pp. 226–234.
Hůrka, K.
and Soós, Á. 1986b. Streblidae. In: Catalogue of Palaearctic Diptera. Volume 11
(Eds. Á. Soós and L. Papp). Elsevier Science Publishers, Amsterdam, and
Akadémiai Kiadó, Budapest, Hungary, pp. 234–236.
Jobling, B.
1934. A revision of the genus Nycteribosca
Speiser (Diptera Pupipara, Streblidae). Parasitology 26: 64–97.
Jobling, B.
1939. On the African Streblidae (Diptera Acalypterae) including the morphology
of the genus Ascodipteron Adens.
and a description of a new
species. Parasitology 31: 147–165.
Jobling, B.
1951. A record of the Streblidae from the Philippines and other Pacific
islands, including morphology of the abdomen, host-parasite relationship and
geographical distribution, and with descriptions of five new species. Trans.
Royal Entomol. Soc. London 102(4): 211–246.
Kock, D.
1974. Pupipare Dipteren von Säugetieren des nordöstlichen Mittelmeerraumens
(Ins.: Diptera). Senckenberg. Biol. 17: 87–104.
Kock, D.
1984. Basilia mongolensis nudior
Hůrka 1972 in Österreich (Insecta: Diptera: Nycteribiidae). Senckenberg. Biol.
65(1–2): 43–46.
Kock, D.
1989. Fledermaus-Fliegen aus der E-Mediterraneis (Diptera: Nycteribiidae).
Entomol. Zeitsch. 99: 56–58.
Kock, D. and
Nader, I.A. 1979. Two bat flies from the Kingdom of Saudi Arabia, their
nomenclature, host specificity and zoogeography (Insecta: Diptera:
Nycteribiidae). Senckenberg. Biol. 60: 65–73.
Kunz, T.H.
and Kurta, A. 1988. Capture methods and holding devices. In: Ecological and
Behavioral Methods for the Study of Bats (Ed. T.H. Kunz). Smithsonian Institution
Press, Washington, D.C., pp. 1–29.
Lewis, R.E.
and Harrison, D.L. 1962. Notes on bats from the Republic of Lebanon. Proc.
Royal Entomol. Soc. London 138: 473–486.
Sanborn,
C.C. and Hoogstraal, H. 1953. Some mammals of Yemen and their ectoparasites.
Field. Zool. 34: 229–252.
Speiser, P.
1907. Preliminary description of three new Nycteribiidae from India. Rec. Ind.
Mus. 1: 295–296.
Theodor, O.
1952. On the zoogeography of some groups of Diptera in the Middle East. Rév.
Fac. Sci. Univ. Istanbul (B) 17(2): 107–119.
Theodor, O.
1954a. 66a. Nycteribiidae. In: Die Fliegen der palaearktischen Region. Volume
12 (Ed. E. Lindner). E. Schweizerbart’sche Verlagsbuchhandlung, Stuttgart,
Germany, pp. 1–44.
Theodor, O.
1954b. 66b. Streblidae. In: Die Fliegen der palaearktischen Region. Volume 12
(Ed. E. Lindner). E. Schweizerbart’sche Verlagsbuchhandlung, Stuttgart,
Germany, pp. 1–12.
Theodor, O.
1955. On the genus Eucampsipoda Kol.
and Dipseliopoda
n.g. (Nycteribiidae,
Diptera). Parasitology 45: 195–229.
Theodor, O.
1956. On the genus Tripselia and the
group of Basilia bathybothyra
(Nycteribiidae, Diptera). Parasitology 46: 353– 394.
Theodor, O.
1967. An Illustrated Catalogue of the ROTHSCHILD Collection of Nycteribiidae in
the British Museum (Natural History), with Keys and Short Descriptions for the
Identification of Subfamilies, Genera, Species and Subspecies.
British Museum (Natural History) Publication, London.
Theodor, O.
1975. Diptera Pupipara; Fauna Palaestina-Insecta I. The Israel Academy of
Sciences and Humanities, Jerusalem, Israel.
Theodor, O.
and Moscona, A. 1954. On bat parasites in Palestine. I. Nycteribiidae,
Streblidae, Hemiptera, Siphonaptera.
Parasitology 44: 157–245.
