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Zootaxa 3893 (1): 077–100 ISSN 1175-5326 (print edition)

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Copyright © 2014 Magnolia Press
Article ZOOTAXA
ISSN 1175-5334 (online edition)
http://dx.doi.org/10.11646/zootaxa.3893.1.3
http://zoobank.org/urn:lsid:zoobank.org:pub:7ABF7414-EC3E-4072-885A-FAD073AC239F

A review of the ectoparasitic mites (Acari: Dermanyssoidea) associated with


birds and their nests in Slovakia, with notes on identification of some species

PETER MAŠÁN1, PETER FENĎA2, JÁN KRIŠTOFÍK1 & BRUCE HALLIDAY3


1
Institute of Zoology, Slovak Academy of Sciences, Dúbravská cesta 9, 845 06 Bratislava, Slovakia.
E-mails: uzaepema@savba.sk; uzaekris@savba.sk
2
Department of Zoology, Faculty of Natural Sciences, Comenius University, Mlynská dolina B-1, 842 15 Bratislava, Slovakia.
E-mail: fenda@fns.uniba.sk
3
CSIRO Ecosystem Sciences, GPO Box 1700, Canberra ACT 2601, Australia. E-mail: Bruce.Halliday@csiro.au

Abstract

We review the parasitic mites of the superfamily Dermanyssoidea found in the nests of Slovakian domestic and wild birds
(and a few mammals), compiled from new data and literature sources. Three mite genera are included, namely Dermany-
ssus Dugès (Dermanyssidae), Ornithonyssus Sambon and Pellonyssus Clark & Yunker (Macronyssidae), in which we rec-
ognised nine reliably documented species. Pellonyssus is represented by one species, Dermanyssus by five species, and
Ornithonyssus by three species. We compiled information on the ecological requirements and host preferences of these
mite species, including data on geographic distribution, altitudinal distribution, and occurrence in different habitats and
nest types. An identification key to the species occurring in Slovakia is provided, together with taxonomic remarks on the
identification and external morphology of some selected species. Parasitic mites have been reported from 119 taxa of Slo-
vakian vertebrates (including man), of which 108 belong to 17 orders of birds. Dermanyssus americanus Ewing and Or-
nithonyssus bursa (Berlese) are reported from Slovakia for the first time. Examination of older voucher material collected
in Slovakia revealed many misidentified specimens: the name Dermanyssus hirundinis (Hermann) has been widely used
for misidentified specimens of Dermanyssus carpathicus Zeman; D. americanus was confused with D. hirundinis; while
O. bursa has been confused with Ornithonyssus sylviarum (Canestrini & Fanzago). Slovakia is the second country in Eu-
rope with known occurrence of the genus Pellonyssus.

Key words: Acari, Mesostigmata, parasitic mites, Dermanyssus, Ornithonyssus, Pellonyssus, bird hosts, Slovakia

Introduction

The mite superfamily Dermanyssoidea is the most ecologically diverse group of Mesostigmata (Walter & Shaw
2005). It includes many species of free-living predators that are abundant in soil, especially in the Hypoaspidinae
(Laelapidae). A basic free-living stock of this type appears to have given rise to a diverse range of families and
genera that are closely associated with vertebrate and invertebrate hosts (Radovsky 1969; Lindquist et al. 2009).
This large assemblage of derivative groups includes commensals, phoretics, and symbionts, as well as obligate
blood-feeding ectoparasites of mammals and birds. Parasitism in these families may be either temporary or
permanent, and some species have mixed feeding behaviour that includes blood-feeding as well as predation and
scavenging. Both parasitic and non-parasitic species can be found in large numbers in the nests and burrows made
by their vertebrate hosts. At least one species, Dermanyssus gallinae (de Geer), attacks the domestic hen and other
captive birds, and can cause huge economic losses (Sparagano et al. 2014).
Some of these species of mites occasionally infest man, and some transmit diseases to humans. Mite-borne
rickettsial pox, scrub typhus, and tick-borne pathogens have been relatively well studied, but mites have also been
found to transmit other rickettsial agents such as Rickettsia akari and Orientia tsutsugamushi. Reeves et al. (2006)
screened 25 species of dermanyssoid mites and found that they contained Anaplasma spp., Bartonella sp.,
Spiroplasma sp., Wolbachia sp., and other unclassified Rickettsiales. Non-tick acarines can also transmit or harbour

Accepted by O. Seeman: 28 Oct. 2014; published: 5 Dec. 2014 77


Anaplasma phagocytophilum, arboviruses, coccidians, Coxiella burnetii, filarial nematodes, hantaviruses,
Rickettsia spp., tapeworms, and other pathogens of wildlife (Huebner et al. 1946; Allred 1954; Yunker 1964;
Zemskaya 1967; Yunker et al. 1975; Moss 1978; Traub et al. 1978; Renz & Wenk 1981; Mohamed et al. 1987;
Fernández-Soto et al. 2001; Walter & Shaw 2005). Study of the systematics and host relationships of
mesostigmatic mites is therefore of medical and veterinary importance, aside from its intrinsic value in studies of
host-parasite co-evolution and the evolution of parasitism.
The present paper summarises records of dermanyssid and macronyssid mites from extensive Slovakian
investigations of birds and their ectoparasites, including rich collections of nest fauna made by J. Krištofík, A.
Darolová, M. Krumpál, D. Cyprich, Z. Országhová, M. Lichard and M. Jurík, who with the help of acarologists (P.
Mašán, P. Fenďa, M. Ambros, M. Mrciak and P. Zeman) made significant contributions to the knowledge of the
nidicolous fauna of mesostigmatic mites. The results of these studies form the basis of an extensive literature of
major papers (Nosek & Lichard 1962; Zeman 1979; Zeman & Jurík 1981; Ambros et al. 1992; Cyprich et al. 1992,
2000; Mašán & Krištofík 1993, 1995; Krištofík et al. 1993, 1996, 2001, 2002, 2003, 2005, 2007, 2009, 2013;
Mašán & Országhová 1995a, 1995b; Krištofík & Mašán 1996; Janiga 1997; Mašán 1997; Fenďa et al. 1998a,
1998b, 2011; Krumpál et al. 1998, 2001; Fenďa & Schniererová 2004, 2005, 2010; Švaňa et al. 2006; Fenďa &
Lengyel 2007; Fenďa 2009; Fenďa & Kicková 2009; Kaľavský et al. 2009), as well a large number of other papers
of marginal importance in relation to the bird hosts. These papers focus on specific birds or specific localities and
habitats, and there has been no large-scale systematic study of parasitic mites in Slovakia. This paper is the first
review of obligatorily haematophagous mesostigmatans of the superfamily Dermanyssoidea specifically associated
with wild birds in Slovakia. It combines the results of previous studies, and adds new previously unpublished
collection data. The main aim of this work is the improvement of the knowledge of the Slovak dermanyssoid fauna,
and contributing to the present knowledge of biodiversity of the European region, particularly with regard to the
West Carpathians and adjacent regions. This is the first host list for bird parasites from Slovakia, despite a long
tradition of study of mites living in association with birds and their nests.

Material and methods

This paper summarises all published data on parasitic mesostigmatic mites specifically associated with birds and
their nests in Slovakia, including both wild and domestic species. We include a few records of these mite species
from soil. Non-parasitic and free-living nidicolous species, and parasitic species specifically associated with
mammals or other vertebrates and occasionally found in the nests of birds, are generally excluded (except those
classified in the genus Ornithonyssus). In addition, we include extensive new data on various mite-bird
associations on the basis of our own collections. Wherever possible we re-examined older material used in previous
publications, to confirm the identifications. For all species, including those with dubious taxonomic status, short
taxonomic notes are given. In Slovakia, representatives of only three genera of Mesostigmata are adapted for an
ectoparasitic mode of life on birds—Dermanyssus, Ornithonyssus, and Pellonyssus (excluding avian nasal
dermanyssoid mites of the family Rhinonyssidae, which includes obligate haematophagous endoparasites of non-
ratite birds). For each species, a bibliography of basic taxonomic works and a list of the most important synonyms
are provided, together with some basic ecological data, such as host and habitat preferences and altitudinal
distribution.
For new collections, mites were extracted from nests using a modified Berlese-Tullgren funnel. Specimens
were mounted on permanent microscope slides using Swan’s gum-chloral medium. If accessible, older material
published by previous authors was re-examined and revised. The terminology of dorsal and ventral chaetotactic
pattern used in this paper follows Evans & Till (1979). Taxonomy of the birds generally follows Boyd (2010) and
Gill & Donsker (2014), and taxonomy of the mammals that of Wilson & Reeder (2005).

78 · Zootaxa 3893 (1) © 2014 Magnolia Press MAŠÁN ET AL.


Systematic account of Slovak taxa

Family: Dermanyssidae Kolenati, 1859

Genus: Dermanyssus Dugès, 1834

Dermanyssus Dugès, 1834: 5. Type species Acarus gallinae De Geer, 1778.

Historical reviews of the genus Dermanyssus can be found in Evans & Till (1962), Moss (1968, 1978) and Roy &
Chauve (2007, 2010). The genus includes about 30 valid species from all climatic zones of the world, all of which
are obligate haematophagous ectoparasites of birds. Five of these species are credibly recorded from Slovakia,
namely D. americanus, D. carpathicus, D. chelidonis, D. gallinae, and D. hirundinis. Roy et al. (2009a, 2010)
recorded D. longipes from Slovakia, but we here consider those specimens to belong to D. hirundinis (see
"Dubious species from Slovakia" below). Nosek & Lichard (1962) identified D. passerinus from Slovakia, but that
record is also regarded as dubious.

Dermanyssus americanus Ewing, 1923

Dermanyssus americanus Ewing, 1923: 24.


Dermanyssus oti Ewing, 1925: 21. Synonymy by Oudemans (1936).
Dermanyssus americanus.—Evans & Till, 1962: 281; Moss, 1968: 71, 1978: 634.

Taxonomic remarks. This is a clearly recognisable and well-defined species. It has only two pairs of J-setae (J1
and J3) on the opisthonotal portion of the dorsal shield, two pairs of small, suboval plates on the posterodorsal soft
integument, and it lacks metasternal setae. These characters cannot be found in any other species that is reliably
documented from Slovakia.
Occurrence and habitat. This species is known from Slovakia on the basis of one specimen deposited in our
mite collection (see below). It was originally reported as Dermanyssus hirundinis (by Krumpál et al. 2001).
Host distribution. Bird nest: Passer sp. (Krumpál et al. 2001: as Dermanyssus hirundinis).

Revised data. Jablunkovské Medzihorie Mts.: 1 ♀, 30 December 1985, Svrčinovec Village, Passer sp.

Dermanyssus carpathicus Zeman, 1979

Dermanyssus carpathicus Zeman, 1979: 173.

Taxonomic remarks. Dermanyssus carpathicus was originally described by Zeman (1979), based on specimens
from Slovakia (type locality: Urpín Mt., Banská Bystrica Town, Central Slovakia), and it has never been re-
described by subsequent authors. Some authors may have confused this species with the strikingly similar D.
hirundinis.
The high level of phenotypic variability in the genus Dermanyssus makes species identification difficult
(Evans & Till 1962; Moss 1978; Roy et al. 2009a), and has led to many misinterpretations and misidentifications.
In Slovakia, the name D. hirundinis has been widely used for misidentified specimens of D. carpathicus, which is
very common in nests of various birds in Slovakia (almost all the published records below). We compared
specimens of D. hirundinis from the specific host Hirundo rustica with other specimens that had been identified as
this species, and collected from other bird hosts. This comparison showed that most of the Slovak specimens
available for our re-examination and published in the past under the name D. hirundinis are actually D.
carpathicus. These two species differ in several small details (e.g., in length of peritreme and shape of anal shield),
but the most obvious differences are in the chaetotaxy of some leg segments, the position of the humeral platelets,
the length of some leg setae, and the posteriormost setae inserted in the soft caudal integument (see the
identification key).

SLOVAKIAN DERMANYSSOIDEA ASSOCIATED WITH BIRDS & NESTS Zootaxa 3893 (1) © 2014 Magnolia Press · 79
We have not followed the diagnostic key for identification of the gallinae-group species from France provided
by Roy et al. (2009a), because it includes several morphological inconsistencies. We cannot confirm the character
states used there for separation of Dermanyssus carpathicus from the other species found in France, in particular
(1) seta al1 on the palpgenu is not characteristically formed, and does not differ from that in D. hirundinis; and (2)
the two dorsal setae on femur I are longer than the other three setae in both species, not only in D. carpathicus
(these two elongated setae are slightly longer and more robust in D. carpathicus). There are many European
publications on mites collected from the nests of various bird hosts in which the name D. hirundinis is used, but the
species is not illustrated or described in these publications, so its true identity is therefore uncertain. Confusion
between these two species is probably widespread.
Occurrence and habitat. Dermanyssus carpathicus is the most frequent and abundant representative of the
genus Dermanyssus in Slovakia, and is widely distributed throughout the territory. It is a polyxenous ectoparasite,
which is distributed from lowlands to subalpine grasslands, with an optimum in the areas up to 800 m a.s.l., and
with an altitudinal range from 100 to 1,440 m a.s.l. This species was collected from a variety of host species as
listed below, but was also found in soil, litter, and detritus taken from caves. Despite its low host specificity (found
in 12 bird orders and 58 host species, see below), it apparently prefers the nests of passeriform birds (69% of all
identified birds), especially normal cup nests, either on the ground or elevated, including specific pendant or sphere
nests which are completely enclosed (Remiz pendulinus). It can also be found in mound or floating platform nests
of some aquatic birds (Charadriiformes, Gruiformes) in very wet situations.
Host distribution. Bird nests: Acrocephalus arundinaceus (Krištofík et al. 2001; Fenďa & Schniererová 2004,
2005), Acrocephalus arundinaceus with Cuculus canorus, Aegithalos caudatus, Alauda arvensis, Alcedo atthis,
Anas platyrhynchos (Fenďa & Schniererová 2005, new data), Anthus trivialis, Aquila chrysaetos, Carduelis
carduelis, Certhia brachydactyla, Chloris chloris, Chroicocephalus ridibundus, Ciconia ciconia, Circus
aeruginosus, Delichon urbicum, Dendrocopos major, Emberiza schoeniclus, Falco cherrug, Falco tinnunculus,
Ficedula albicollis (Švaňa et al. 2006), Fringilla coelebs, Fulica atra, Himantopus himantopus, Lanius collurio
(Krištofík et al. 2002), Lanius minor (Krištofík et al. 2002), Limosa limosa, Linaria cannabina, Locustella
luscinioides, Merops apiaster (Krištofík et al. 1996, Fenďa 2008), Motacilla alba, Motacilla cinerea, Motacilla
flava, Muscicapa striata, Nannus troglodytes, Notiocichla palustris (Krištofík et al. 2005), Notiocichla scirpacea
(Fenďa & Schniererová 2004, 2005), Numenius arquata, Oriolus oriolus, Parus major (Zeman 1979, new data),
Parus sp., Passer montanus (Krumpál et al. 2001, Švaňa et al. 2006, Fenďa et al. 2011, new data), Phoenicurus
ochruros, Phoenicurus phoenicurus (Zeman 1979), Phoenicurus sp., Phylloscopus collybita, Pica pica, Podiceps
grisegena, Prunella modularis, Remiz pendulinus (Krištofík et al. 1993, Krištofík & Mašán 1996, Mašán 1997),
Riparia riparia (Mašán & Krištofík 1993, Krištofík et al. 1994, Fenďa 2008), Saxicola rubetra, Serinus serinus,
Strix aluco, Sylvia atricapilla, Taeniopygia castanotis (Cyprich et al. 2000), Turdus merula, Turdus philomelos,
Turdus pilaris, Turdus sp., Turdus torquatus (Krumpál et al. 1998), Vanellus vanellus. Mammals: Apodemus
flavicollis (Krumpál et al. 1998), Canis familiaris, Glis glis (Mašán & Krištofík 1996), Talpa europaea (Mašán et
al. 1994). Other findings: soil and litter (Fenďa & Mašán 2003), detritus of caves.
Based on our revision (PM), all of the above published specimens have been misidentified, and originally
reported under the name Dermanyssus hirundinis, except those of Zeman (1979). In general, the new hosts (and
findings) of this section are listed in this paper with no author’s reference, but their complete collection data are
available in the paragraphs labelled "New data" (see below).

New data. Borská Nížina Lowland: 2 NN, 15 May 1991, Jakubov Village, Jakubovské Rybníky Fishponds, Numenius arquata;
2 ♀♀, 1 ♂, 7 NN, 15 May 1991, Vysoká Pri Morave Village (floodplain forest), Aegithalos caudatus; 1 ♀, 12 June 1991, Malé
Leváre Village (meadow), Saxicola rubetra; 4 ♀♀, 1 N, 21 June 1991, Vysoká Pri Morave Village (meadow), Alauda arvensis;
1 ♀, 21 June 1991, Vysoká Pri Morave Village (meadow), Motacilla flava; 1 N, 7 June 1992, Stupava Town (floodplain forest),
Phylloscopus collybita; 2 ♀♀, 4 NN, 16 June 1992, Malé Leváre Village (floodplain forest), Certhia brachydactyla; 1 N, 28
June 1992, Vysoká Pri Morave Village (reed stand), Emberiza schoeniclus; 3 ♀♀, 1 ♂, 12 NN, 30 June 1992, Jakubov Village,
Jakubovské Rybníky Fishponds, Chloris chloris; 1 ♀, 3 NN, 30 June 1992, Jakubov Village, Jakubovské Rybníky Fishponds,
Serinus serinus; 1 ♀, 5 August 1992, Vysoká Pri Morave Village (meadow), Anthus trivialis; 1 N, 16 June 1993, Vysoká Pri
Morave Village (canal), Anas platyrhynchos; 4 ♀♀, 19 July 1993, Malé Leváre Village (floodplain forest), Prunella modularis;
2 ♀♀, 1 ♂, 2 NN, 26 May 2002, Bratislava Capital, Devínske Jazero Settlement, Linaria cannabina; 3,044 ♀♀, 1,129 ♂♂, 488
NN, 28 May 2002, Bratislava Capital, Devínske Jazero Settlement, Phoenicurus ochruros; 19 ♀♀, 1 ♂, 3 June 2008, Bratislava
Capital, Devínske Jazero Settlement, Falco tinnunculus. Malá Fatra Mts.: 3 ♀♀, 3 ♂♂, 1 N, 21 August 1992, Aquila
chrysaetos. Malé Karpaty Mts.: 1 ♀, 2 NN, 25 June 1990, Bratislava Capital, Železná Studnička Forest, Nannus troglodytes; 2

80 · Zootaxa 3893 (1) © 2014 Magnolia Press MAŠÁN ET AL.


♀♀, 12 July 1990, Bratislava Capital, zoological garden, Parus major; 5 ♀♀, 6 NN, 30 August 1990, Bratislava Capital,
Železná Studnička Forest, Alcedo atthis; 2 ♀♀, 1 ♂, 23 July 1991, Bratislava Capital, Železná Studnička Forest, Motacilla
cinerea; 39 ♀♀, 12 ♂♂, 140 NN, 3 March 1992, Bratislava Capital, zoological garden, Passer montanus; 1 ♀, 29 July 1993,
Falco cherrug. Myjavská Pahorkatina Hills: 5 ♀♀, 1 N, 14 May 1992, Sobotište Village (forest), Strix aluco. Podbeskydská
Brázda Mts.: 97 ♀♀, 20 ♂♂, 10 NN, 18 July 1989, Mútne Village, Phoenicurus sp. Podunajská Rovina Lowland: 2 ♀♀, 8 NN,
28 June 1989, Veľké Blahovo Village, Veľkoblahovské Rybníky Fishponds, Fulica atra; 9 ♀♀, 1 N, 3 August 1989, Bodíky
Village, Ciconia ciconia; 1 ♀, 1 September 1989, Dunajská Lužná Village (windbreak), Falco tinnunculus; 1 ♀, 1 September
1989, Dunajská Lužná Village (windbreak), Pica pica; 31 ♀♀, 3 November 1989, Veľké Blahovo Village, Veľkoblahovské
Rybníky Fishponds, Oriolus oriolus; 4 ♀♀, 3 ♂♂, 47 NN, 5 June 1990, Dobrohošť Village (floodplain forest), Sylvia
atricapilla; 144 ♀♀, 25 ♂♂, 129 NN, 5 June 1990, Trstená Na Ostrove Village, Kráľovská Lúka Arm (floodplain forest),
Turdus philomelos; 10 ♀♀, 1 ♂, 10 NN, 2 July 1990, Trstená Na Ostrove Village, Kráľovská Lúka Arm (floodplain forest),
Turdus merula; 1 ♀, 1 N, 2 July 1990, Trstená Na Ostrove Village, Kráľovská Lúka Arm (floodplain forest), Fringilla coelebs;
1 ♀, 1 ♂, 17 August 1990, Gabčíkovo Village (floodplain forest), Carduelis carduelis; 1 ♀, 1 ♂, 4 September 1990, Komárno
Town, Apáli Nature Reserve, Dendrocopos major; 1 ♀, 5 September 1990, Gabčíkovo Village, Delichon urbicum; 4 ♀♀, 1 N,
23 July 1991, Dolný Štál Village, Hroboňovské Rybníky Fishponds, Chroicocephalus ridibundus; 1 ♀, 7 June 1992, Veľké
Blahovo Village, Vanellus vanellus; 1 N, 19 June 1992, Veľké Blahovo Village, Veľkoblahovské Rybníky Fishponds, Limosa
limosa; 233 ♀♀, 16 ♂♂, 349 NN, 23 June 1992, Vojka Nad Dunajom Village (canal), Himantopus himantopus; 6 NN, 15 July
1992, Šuľany Village (floodplain forest), Muscicapa striata; 2 ♀♀, 4 August 1992, Dolný Štál Village, Hroboňovské Rybníky
Fishponds, Acrocephalus arundinaceus + Cuculus canorus; 4 ♀♀, 1 ♂, 16 July 1993, Veľké Blahovo Village, Veľkoblahovské
Rybníky Fishponds, Circus aeruginosus; 1 ♀, 25 July 1994, Dolný Štál Village, Hroboňovské Rybníky Fishponds, Oriolus
oriolus; 24 ♀♀, 3 ♂♂, 8 NN, 18 July 2006, Veľké Blahovo Village, Veľkoblahovské Rybníky Fishponds, Locustella
luscinioides. Trnavská Pahorkatina Hills: 2 ♀♀, 1 N, VI. 1992, Trnava Town, Trnavské Rybníky Fishponds, Podiceps
grisegena. Veľká Fatra Mts.: 22 ♀♀, 3 NN, 22 June 1989, Stankovany Village, Turdus sp.; 3 ♀♀, 1 ♂, ibid., Parus sp.; 5 ♀♀,
ibid., Canis familiaris; 18 ♀♀, 762 NN, 27 August 1993, Stankovany Village, Podšíp Settlement, Motacilla alba. Veporské
Vrchy Mts.: 1 ♀, 6 July 1993, Hriňová Village, Pivnička Settlement, Turdus pilaris.

Dermanyssus chelidonis Oudemans, 1939

Dermanyssus hirundinis.—Berlese, 1889: 3. Misidentification.


Dermanyssus chelidonis Oudemans, 1939: 306. New name for Dermanyssus hirundinis Hermann, 1804 sensu Berlese (1889).
Dermanyssus chelidonis.—Evans & Till, 1962: 283, 1966: 353; Moss, 1968: 79, 1978: 634.

Taxonomic remarks. The species may be easily separated from its congeners by its markedly reduced peritremes,
which reach only the mid level of the coxae II. Other distinguishing characters can be found in the identification
key below.
Occurrence and habitat. There are only a few records of this species from the western and central parts of
Slovakia, but the species is undoubtedly more common and widely distributed than has been reported. The
recorded vertical distribution of the Slovakian specimens ranges from 128 to 600 m a.s.l. Dermanyssus chelidonis
appears to be almost host-specific for the common house martin, Delichon urbicum (86.2% of all specimens
reported from Slovakia), although it occasionally occurs in the nests of other bird species (see below).
Host distribution. Bird nests: Delichon urbicum, Linaria cannabina, Riparia riparia. These hosts have been
previously listed by Fenďa (2009), without complete collection data, which is added below. Another host, the
Eurasian blue tit Cyanistes caeruleus, is not included among the hosts because Zeman & Jurík (1981) did not
record D. chelidonis with this host in the territory of the former Czechoslovakia.

New data. Borská Nížina Lowland: 9 ♀♀, 5 ♂♂, 1 N, 8 April 1993, Bratislava Capital, Devínske Jazero Settlement, Linaria
cannabina. Malé Karpaty Mts.: 11 ♀♀, 1 ♂, 10 September 1989, Bratislava Capital, Rača Settlement, Delichon urbicum.
Podunajská Rovina Lowland: 3 ♀♀, 1 ♂, 15 October 1986, Šamorín Village, Riparia riparia; 4 ♀♀, 5 September 1990,
Gabčíkovo Village, Delichon urbicum. Starohorské Vrchy Mts.: 23 ♀♀, 1 N, 13 September 1989; 13 ♀♀, 1 ♂, 11 NN, 17
September 1989, Staré Hory Village, Delichon urbicum. Štiavnické Vrchy Mts.: 52 ♀♀, 2 ♂♂, 30 September 1986, Štiavnické
Bane Village, Počúvadlo Lake, Delichon urbicum.

SLOVAKIAN DERMANYSSOIDEA ASSOCIATED WITH BIRDS & NESTS Zootaxa 3893 (1) © 2014 Magnolia Press · 81
Dermanyssus gallinae (De Geer, 1778)

Acarus gallinae De Geer, 1778: 111.


Dermanyssus avium Dugès, 1834: 18. Synonymy by Oudemans (1936).
Dermanyssus evotomydis Ewing, 1933: 11. Synonymy by Evans & Till (1962).
Dermanyssus gallinae.—Evans & Till, 1962: 283; 1966: 350; Moss, 1968: 78; 1978: 634.

Taxonomic remarks. This species may be distinguished from other species of Dermanyssus in Slovakia by the
presence of four pairs of J-setae on the opisthonotal portion of the dorsal shield, and peritremes that are not
strongly reduced in length.
Occurrence and habitat. Dermanyssus gallinae is of the greatest economic and veterinary importance among
the studied species because it occurs worldwide and is closely associated with domesticated birds. Based on
published Slovak records, the most frequent infestation was found in the nests of Columba livia f. domestica
(66.5% of all specimens, 85 mites per nest), Passer montanus (7.8%, 31 mites per nest) and Gallus gallus var.
domesticus (5.9%, 93 mites per nest). Among the wild birds, D. gallinae is more frequently found in the nests of
birds that are common in synanthropic and urban habitats, namely Anas platyrhynchos, Corvus frugilegus, Cygnus
olor, Delichon urbicum, Falco tinnunculus, Hirundo rustica, Motacilla alba, Passer spp., Phoenicurus ochruros,
and Turdus merula. A unusual finding was documented by Mrciak (1963) who observed this species on a dung
beetle, Geotrupes stercorarius. It has been recorded at altitudes between 100 and 1,150 m a.s.l.
Host distribution. Bird nests: Acrocephalus arundinaceus (Fenďa & Schniererová 2004, 2005), Anas
platyrhynchos (Fenďa & Schniererová 2005, new data), Asio otus, Aves sp. (Dudich et al. 1940), Circus
aeruginosus, Columba livia f. domestica (Ambros et al. 1992, Cyprich et al. 1992, Janiga 1997, Fenďa & Kicková
2009, new data), Coracias garrulus, Corvus frugilegus, Cygnus olor, Delichon urbicum (Ambros et al. 1992, new
data), Falco tinnunculus (Kaľavský et al. 2009, new data), Ficedula albicollis, Fringilla coelebs, Fulica atra
(Fenďa & Schniererová 2010, new data), Gallus gallus var. domesticus (Mrciak & Rosický 1956: as Dermanyssus
gallinarum De Geer, Mrciak 1963, Fenďa & Mašán 2003, new data), Hirundo rustica (Ambros et al. 1992), Lanius
collurio, Merops apiaster (Krištofík et al. 1996), Motacilla alba, Nannus troglodytes, Parus major (Ambros et al.
1992, new data), Parus sp., Passer domesticus (Krumpál et al. 2001), Passer montanus (Krumpál et al. 2001,
Švaňa et al. 2006, Fenďa et al. 2011, new data), Phoenicurus ochruros, Remiz pendulinus (Krištofík et al. 1993),
Riparia riparia (Ambros et al. 1992, Mašán & Krištofík 1993, Fenďa 2008), Sturnus vulgaris (Švaňa et al. 2006,
new data), Sylvia atricapilla, Sylvia sp., Turdus merula, Turdus philomelos, Turdus sp. Combined nests:
Dendrocopos sp. and Micromammalia sp., Parus major and Apodemus sp., Parus sp. and Apodemus sp. Mammals:
Apodemus sp., Mus musculus (Mrciak 1963), Muscardinus avellanarius, Mustela putorius (Mrciak 1963), Rattus
norvegicus (Mrciak 1963), on human. Other findings: on scarabaeid Geotrupes stercorarius (Mrciak 1963), soil
detritus (Kalúz & Žuffa 1986, Fenďa 2008).
Ficedula albicollis, Jynx torquilla Linnaeus, Periparus ater (Linnaeus), Parus major, Phoenicurus
phoenicurus, Sitta europaea—the host birds given for D. gallinae from the former Czechoslovakia by Zeman &
Jurík (1981), are not included here due to lack of specific collection data.

New data. Borská Nížina Lowland: 3 ♀♀, 3 NN, 4 June 1992, Malé Leváre Village (canal), Cygnus olor; 10 ♀♀, 1 ♂, 2 NN,
17 August 1992, Malé Leváre Village (floodplain forest), Nannus troglodytes; 19 ♀♀, 28 ♂♂, 12 NN, 5 July 1994, Borský
Mikuláš Village, Phoenicurus ochruros. Chvojnická Pahorkatina Hills: 9 ♀♀, 8 ♂♂, 14 NN, 9 April 1989, Skalica Town,
Columba livia f. domestica. Malé Karpaty Mts.: 105 ♀♀, 36 ♂♂, 7 NN, 20 June 1989, Bratislava Capital, zoological garden,
Parus major; 2 ♀♀, 16 September 1989, Bratislava Capital, zoological garden, Passer montanus; 1 ♀, 12 July 1990, Bratislava
Capital, zoological garden, Turdus merula; 5 ♀♀, 20 April 2005, Bratislava Capital, on human; 9 ♀♀, 8 ♂♂, 1 July 2009,
Bratislava Capital, Karlova Ves Settlement, Falco tinnunculus. Oravská Vrchovina Highland: 3 ♀♀, 6 NN, 26 February 1987,
Dolný Kubín Town, Parus major. Podunajská Rovina Lowland: 1 ♂, 15 January 1985, Svätý Jur Village, Šúr Nature Reserve,
Ficedula albicollis; 1 ♀, 5 NN, 5 December 1985, ibid., Muscardinus avellanarius; 4 NN, 17 April 1986, ibid., Anas
platyrhynchos; 1 ♀, 5 April 1989, Pataš Village, Corvus frugilegus; 9 ♀♀, 1 ♂, 1 N, 28 June 1989, Veľké Blahovo Village,
Veľkoblahovské Rybníky Fishponds, Fulica atra; 1 ♀, 3 August 1989, Dunajská Lužná Village (windbreak), Asio otus; 63 ♀♀,
12 ♂♂, 1 N, 8 September 1989, Gabčíkovo Village, Delichon urbicum; 2 ♀♀, 5 June 1990, Dobrohošť Village (floodplain
forest), Sylvia atricapilla; 11 ♀♀, 2 ♂♂, 1 N, 5 June 1990, Gabčíkovo Village, Motacilla alba; 1 N, 5 June 1990, Trstená Na
Ostrove Village, Kráľovská Lúka Arm (floodplain forest), Turdus philomelos; 1 ♀, 2 July 1990, Trstená Na Ostrove Village,
Kráľovská Lúka Arm (floodplain forest), Fringilla coelebs; 71 ♀♀, 19 ♂♂, 29 NN, 5 September 1990, Gabčíkovo Village,
Delichon urbicum; 3 ♀♀, 1 ♂, 5 NN, 10 September 1991, Veľké Blahovo Village, Veľkoblahovské Rybníky Fishponds, Circus

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aeruginosus; 1 ♀, 7 NN, 25 November 1994, Dobrohošť Village, Dunajské Kriviny (floodplain forest), Sturnus vulgaris; 1 ♀,
ibid., Parus major + Apodemus sp.; 1 ♀, ibid., Apodemus sp.; 3 ♀♀, 4 December 1995, ibid., Parus major; 1 ♀, ibid., Parus sp.
+ Apodemus sp.; 263 ♀♀, 169 ♂♂, 97 NN, ibid., Sturnus vulgaris; 5 ♀♀, 13 ♂♂, 12 NN, ibid., Sylvia sp.; 2 ♂♂, ibid., Turdus
sp.; 1 ♀, 27 February 2000, Rusovce Village, Sturnus vulgaris; 10 ♀♀, 8 ♂♂, 14 NN, 20 April 2004, Dobrohošť Village,
Gallus gallus var. domesticus; 55 ♀♀, 14 ♂♂, 3 NN, 9 June 2007, 3 ♀♀, 1 ♂, 16 July 2007, Martovce Village, Coracias
garrulus; 6 ♀♀, 1 N, 12 August 2007, Vojka Nad Dunajom Village, Gallus gallus var. domesticus. Popradská Kotlina Basin: 1
♀, 15 July 1994, Stará Lesná Village, Parus sp.; 22 ♀♀, 9 ♂♂, 3 NN, 15 March 1995, Stará Lesná Village, Aves sp. Slanské
Vrchy Mts.: 1 ♀, 14 February 1989, Zemplínska Teplica Village, Krčmárka Forest, Parus major. Starohorské Vrchy Mts.: 3
♀♀, 1 ♂, 3 NN, 17 September 1989, Staré Hory Village, Delichon urbicum. Veľká Fatra Mts.: 1 ♀, 2 NN, 25 February 1987,
Ľubochňa Village, Ľubochnianska Dolina Valley, Dendrocopos sp. + Micromammalia sp. Veporské Vrchy Mts.: 1 ♀, 1 ♂, 6
July 1993, Hriňová Village, Pivnička Settlement, Lanius collurio. Zvolenská Kotlina Basin: 6 ♀♀, 6 July 1993, Kováčová
Village (forest), Ficedula albicollis.

Dermanyssus hirundinis (Hermann, 1804)

Acarus hirundinis Hermann, 1804: 83.


Dermanyssus hirundinis.—Evans & Till, 1962: 285, 1966: 356; Moss, 1968: 75, 1978: 634.

Taxonomic remarks. This species is often confused with Dermanyssus carpathicus (see taxonomic remarks for
that species). It shows phenotypic variability in the length of the dorsal shield setae and the width of posterior
portion of the dorsal shield. In Slovakia, populations identified here as Dermanyssus hirundinis may be a mixture
of two morphologically similar species.
Occurrence and habitat. This species is widely distributed throughout Slovakia, and has been reported from
the collection sites at altitudes between 100 and 1,650 m a.s.l., and from the nests of 41 host species (see below)
representing nine orders of birds. Its host range differs from that of D. carpathicus, despite the fact that both
species can occasionally be found together in the same nest, and relatively often in the nests of the same host
species (22.8% of all identified birds). Its relative frequency in the nests of passeriform hosts (58.5%) is lower than
that for D. carpathicus (69%). However this number is inflated by the affinity of D. hirundinis for nests built in
artificial nest boxes, which are mostly exploited by small passeriform birds. Generally, it preferentially inhabits
nests with dry and unlined depression to house the eggs, including (1) closed or open cup nests made from mud
pellets by swallows and martins (Hirundinidae); (2) nest boxes used by cavity-nesting birds; (3) elevated platform
nests of diurnal birds of prey (Accipitriformes, Falconiformes); (4) stick nests of crows, magpies and some owls;
and (5) mound or floating platform nests of some water birds, mainly waterfowl (Anseriformes).
Host distribution. Bird nests: Accipiter gentilis, Acrocephalus arundinaceus (Krištofík et al. 2001; Fenďa &
Schniererová 2004, 2005), Anas crecca, Anas platyrhynchos (Fenďa & Schniererová 2005, new data), Anser anser
(Fenďa & Schniererová 2005), Anthus spinoletta, Asio otus, Aythya ferina (Fenďa & Schniererová 2005), Aythya
fuligula (Fenďa & Schniererová 2005), Carduelis carduelis, Chroicocephalus ridibundus (Fenďa & Schniererová
2010, new data), Ciconia ciconia, Circus aeruginosus, Coracias garrulus, Cyanistes caeruleus, Delichon urbicum,
Erithacus rubecula, Falco tinnunculus (Kaľavský et al. 2009, new data), Falco vespertinus, Fulica atra (Fenďa &
Schniererová 2010, new data), Hirundo rustica (Mrciak & Rosický 1956, Mašán & Országhová 1995, Országhová
et al. 2006, new data), Ictinaetus pomarinus, Lanius minor, Luscinia megarhynchos, Merops apiaster (Krištofík et
al. 1996, Fenďa 2008), Milvus migrans, Motacilla alba, Motacilla cinerea, Nannus troglodytes, Parus major
(Švaňa et al. 2006, new data), Passer domesticus (Mrciak 1977), Passer montanus (Krumpál et al. 2001, Švaňa et
al. 2006, Fenďa et al. 2011, new data), Passer sp. (Krumpál et al. 2001), Phoenicurus ochruros (Krumpál et al.
1998), Pica pica, Riparia riparia (Mašán & Krištofík 1993, Krištofík et al. 1994, Fenďa 2008), Sitta europaea,
Sturnus vulgaris (Švaňa et al. 2006), Sylvia atricapilla, Turdus merula, Turdus philomelos. Combined nests: Aves
sp. and Apodemus sp., Ficedula albicollis and Glis glis, Parus major and Apodemus sp. Mammals: Glis glis
(Mašán & Krištofík 1996), Mus musculus (Mrciak 1963). Other findings: moss, soil detritus of caves.
In former Czechoslovakia, Zeman & Jurík (1981) cited the species additionally from Ficedula albicollis,
Poecile palustris (Linnaeus) and Strix aluco, with no geographic specification of individual records.

New data. Borská Nížina Lowland: 1 ♂, 16 August 1991, Malé Leváre Village (floodplain forest), Milvus migrans; 10 ♀♀, 3
June 2008, Bratislava Capital, Devínske Jazero Settlement, Falco tinnunculus. Jablunkovské Medzihorie Mts.: 3 ♀♀, 4 NN, 29
December 1985, Svrčinovec Village, Carduelis carduelis. Košická Kotlina Basin: 118 ♀♀, 13 NN, 22 October 1987, Moldava

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Nad Bodvou Village, Paňovský Les Forest, Glis glis; 2 ♀♀, ibid., Cyanistes caeruleus; 11 ♀♀, 6 NN, ibid., Aves sp. +
Apodemus sp.; 6 ♀♀, 13 NN, ibid., Ficedula albicollis + Glis glis; 5 ♀♀, ibid., Parus major + Apodemus sp. Kysucké Beskydy
Mts.: 16 NN, 3 June 1987, Oščadnica Village, Anas crecca. Laborecká Vrchovina Highland: 1 ♀, 31 October 1998, 6 ♀♀, 2
January 1999, Svetlice Village, Ictinaetus pomarinus; 1 ♀, 31 October 1998, Palota Village, Ictinaetus pomarinus; 19 ♀♀, 31
October 1998, Rokytov Pri Humennom Village, Ictinaetus pomarinus; 15 ♀♀, 2 January 1999, Palota Village, Ictinaetus
pomarinus; 11 ♀♀, 2 January 1999, Rokytovce Village, Ictinaetus pomarinus. Malá Fatra Mts.: 1 ♀, 2 NN, 9 July 2008, 1 N, 6
July 2009, 1 ♀, 31 October 2010, Turany Village, Veľký Kriváň Mt., Anthus spinoletta; 1 N, 6 July 2009; 2 ♀♀, 31 October
2010, Turany Village, Snilovské Sedlo Mountain Pass, Anthus spinoletta; 1 ♀, 31 October 2010, Šútovo Village, Chleb Mt.,
Anthus spinoletta. Malé Karpaty Mts.: 2 ♀♀, 16 September 1989, Bratislava Capital, zoological garden, Passer montanus; 17
♀♀, 2 ♂♂, 9 NN, 12 July 1990, Bratislava Capital, zoological garden, Passer montanus; 3 ♀♀, 1 N, 25 July 1990, Bratislava
Capital, Železná Studnička Forest, Nannus troglodytes; 2,002 ♀♀, 85 ♂♂, 211 NN, 23 July 1991, Bratislava Capital, Železná
Studnička Forest, Hirundo rustica; 10 ♀♀, 23 July 1991, Bratislava Capital, Železná Studnička Forest, Motacilla cinerea; 1 ♀,
3 June 1992, Bratislava Capital, zoological garden, Turdus merula; 14 ♀♀, 17 December 1998, Borinka Village, Trojuholník
Cave (soil). Moravsko-sliezske Beskydy Mts.: 1 ♀, 30 June 1997, Klokočov Village, Malý Polom Mt. (moss). Podunajská
Rovina Lowland: 2 ♀♀, 2 ♂♂, 15 January 1985, Svätý Jur Village, Šúr Nature Reserve, Parus major; 1 ♀, 4 July 1989,
Dunajská Lužná Village (windbreak), Pica pica; 11 ♀♀, 3 August 1989, Bodíky Village, Ciconia ciconia; 214 ♀♀, 5 ♂♂, 5
NN, 3 August 1989, Dunajská Lužná Village (windbreak), Asio otus; 1 ♀, 2 July 1990, Trstená Na Ostrove Village, Kráľovská
Lúka Arm (floodplain forest), Sylvia atricapilla; 25 ♀♀, 6 ♂♂, 6 July 1990, Šamorín Town (windbreak), Pica pica; 3 ♀♀, 5
September 1990, Gabčíkovo Village, Delichon urbicum; 3 ♀♀, 23 July 1991, Dolný Štál Village, Hroboňovské Rybníky
Fishponds, Chroicocephalus ridibundus; 2 ♂♂, 2 NN, 26 July 1991, Accipiter gentilis; 1 ♀, 4 September 1991, Gabčíkovo
Village, Motacilla alba; 1 ♀, 10 September 1991, Veľké Blahovo Village, Veľkoblahovské Rybníky Fishponds, Circus
aeruginosus; 5 NN, 3 August 1992, Jarovce Village (windbreak), Turdus philomelos; 103 ♀♀, 14 ♂♂, 14 NN, 17 July 1993,
Veľké Blahovo Village (windbreak), Falco vespertinus; 57 ♀♀, 5 ♂♂, 10 NN, 17 May 1994, Bratislava Capital, Krasňany
Settlement, Erithacus rubecula; 1 ♀, 2 August 1994, Rusovce Village (windbreak), Falco vespertinus; 1 ♀, 30 August 1996,
Medveďov Village (floodplain forest), Luscinia megarhynchos; 1 ♀, 7 NN, 23 June 1998, Rusovce Village (windbreak), Falco
tinnunculus; 15 ♀♀, 4 ♂♂, 3 NN, 23 August 1998, Rusovce Village (windbreak), Falco vespertinus; 1 ♀, 4 NN, 27 June 2001,
Bratislava Capital, Podunajské Biskupice Settlement, Lanius minor; 3 ♀♀, 16 July 2007, Martovce Village, Coracias garrulus;
406 ♀♀, 78 ♂♂, 24 NN, 3 July 2009, Alžbetin Dvor Village, Falco tinnunculus. Považské Podolie Basin: 3 ♀♀, 13 August
1989, Dubnica Nad Váhom Town, Parus major. Slovenský Raj Mts.: 1 ♀, 28 July 2000, Spišská Nová Ves Town, Medvedia
Jaskyňa Cave (soil); 16 NN, 11 July 2012, Stratená Village, Duča Cave (soil and litter). Starohorské Vrchy Mts.: 8 ♀♀, 13
September 1989, 327 ♀♀, 12 ♂♂, 329 NN, 17 September 1989, Staré Hory Village, Delichon urbicum. Trnavská Pahorkatina
Hills: 547 ♀♀, 112 ♂♂, 75 NN, 23 September 2012, Horná Streda Village, Hirundo rustica; 8 ♀♀, 12 August 1982,
Budmerice Village, Passer domesticus. Veľká Fatra Mts.: 6,510 ♀♀, 2,185 ♂♂, 13,517 NN, 29 August 1991, Stankovany
Village, Podšíp Settlement, Parus major; 9 ♀♀, 2 ♂♂, 27 NN, 26 June 1993, Stankovany Village, Podšíp Settlement, Sitta
europaea. Vihorlatské Vrchy Mts.: 8 ♀♀, 28 December 1998, Ptičie Village, Ictinaetus pomarinus.

Family: Macronyssidae Oudemans, 1936

Genus: Ornithonyssus Sambon, 1928

Ornithonyssus Sambon, 1928: 105. Type species: Dermanyssus sylviarum Canestrini & Fanzago, 1877.

The genus Ornithonyssus includes obligatorily haematophagous mites that are ectoparasites of birds and mammals.
It contains up to 30 named species, distributed especially throughout the Neotropical and Nearctic Regions. The
concept of Ornithonyssus used here follows the classification by Micherdziński (1980). That concept is
considerably wider than that of Radovsky (2010), which divides Ornithonyssus into several separate genera. Three
identified species are found in Slovakia, and all are considered to be cosmopolitan, namely O. bacoti, O. bursa and
O. sylviarum (Radovsky 2010). The other Slovak member of the genus, Ornithonyssus pipistrelli, is essentially a
parasite of vespertilionid bats, and can occasionally be found on some birds and dormice (Rodentia, Gliridae) and
their nests, mainly on those nesting in natural tree hollows and artificial nest boxes used by bats for shelter.

Ornithonyssus bacoti (Hirst, 1913)

Leiognathus bacoti Hirst, 1913: 122.


Haemogamasus sanguineus Ewing & Stover, 1915: 109. Synonymy by Strandtmann & Wharton (1958).
Liponyssus tenuiscutatus Ewing, 1923: 22. Synonymy by Strandtmann & Wharton (1958).

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Liponyssus nagayoi Yamada, 1931: 258. Synonymy by Strandtmann & Wharton (1958).
Liponyssus hirsti Fonseca, 1935: 94. Synonymy by Micherdziński (1980).
Liponyssus brasiliensis Fonseca, 1939: 147. Synonymy by Micherdziński (1980).
Liponyssus lutzi Fonseca, 1942: 104. Synonymy by Micherdziński (1980).
Liponyssus monteiroi Fonseca, 1942: 107. Synonymy by Micherdziński (1980).
Liponyssus vitzthumi Fonseca, 1942: 110. Synonymy by Micherdziński (1980).
Liponyssus meprai Manso Soto & Pletneff, 1951: 15. Synonymy by Micherdziński (1980).
Bdellonyssus bacoti.—Zemskaya, 1955: 350.
Ornithonyssus bacoti.—Evans & Till, 1966: 321; Wilson, 1967: 135; Micherdziński, 1980: 41.

Taxonomic remarks. Our specimens from Slovakia are generally in good agreement with those described by
Zemskaya (1955), Evans & Till (1966), Wilson (1967) and Micherdziński (1980), except for the length of the
peritremes. The anterior sections of the peritremes in our specimens are slightly shorter than in the specimens
previously described—they reach to the mid level of coxae II, while the peritremes in this species are usually
described as reaching at least to the posterior margin of the coxae I.
Occurrence and habitat. The European distribution of this species was originally centered mainly around
ports and associated primarily with rats of the genus Rattus Fischer (Beran et al. 1956; Evans & Till 1966). There
are only a few reports of this mite in Slovakia, mostly from associations with the house mouse, Mus musculus. The
finding of some specimens in an aviary with the Australian zebra finch, Taeniopygia castanotis, is presumed to be
an accidental but remarkable occurrence.
Host distribution. Bird nests: Taeniopygia castanotis (Cyprich et al. 2000). Mammals: Mus musculus (Mrciak
1963, Ambros & Stanko 1989, Stanko 1995, Fenďa & Mašán 2003, new data).

New data. Východoslovenská Rovina Lowland: 5 ♀♀, 30 October 1994, Markovce Village, Mus musculus.

Ornithonyssus bursa (Berlese, 1888)

Leiognathus bursa Berlese, 1888: 208.


Leiognathus morsitans Hirst, 1915: 56. Synonymy by Strandtmann & Wharton (1958).
Leiognathus constrictus Ewing, 1923: 16. Synonymy by Strandtmann & Wharton (1958).
Liponyssus eruditus Fonseca, 1935: 85. Synonymy by Micherdziński (1980).
Liponyssus iheringi Fonseca, 1935: 89. Synonymy by Fonseca (1958).
Ornithonyssus bursa.—Micherdziński, 1980: 53.

Taxonomic remarks. Despite the fact that this species is easily recognisable, the Slovak specimens have been
misidentified (Mašán & Országhová 1995) and incorrectly listed under the name Ornithonyssus sylviarum.
Occurrence and habitat. This species is found throughout the temperate and tropical regions of the world. In
Europe, it is relatively rare, but can be massively abundant in some colonies of the barn swallow Hirundo rustica
(Gjelstrup & Møller 1986; Møller 2002). Ornithonyssus bursa is newly recorded for the Slovak fauna.
Host distribution. Bird nests: Hirundo rustica (Mašán & Országhová 1995; Országhová et al. 2006; both as
Ornithonyssus sylviarum).

Revised data. Podunajská rovina lowland: 42 ♀♀, 1 ♂, 3 NN, 28 July 1993; 105 ♀♀, 11 ♂♂ 7 NN, 4 August 1993; 4,147 ♀♀,
736 ♂♂, 597 NN, 15 August 1993; 8,835 ♀♀, 1,513 ♂♂, 669 NN, 18 August 1993; 260 ♀♀, 22 ♂♂, 20 NN, 26 August 1993;
82 ♀♀, 12 ♂♂, 3 NN, 29 August 1993; 20,906 ♀♀, 9,182 ♂♂, 11,629 NN, 2 September 1993; 7,108 ♀♀, 6,039 ♂♂, 10,960
NN, 8 September 1993, Rusovce Village, Hirundo rustica.

Ornithonyssus pipistrelli (Oudemans, 1904)

Liponyssus musculi.—Oudemans, 1902: 17. Misidentification.


Liponyssus pipistrelli Oudemans, 1904: 111. New name for Liponyssus musculi (C. L. Koch, 1836) sensu Oudemans (1902).
Ornithonyssus pavlovskii Lange, 1959: 484. Synonymy by Micherdziński (1980).
Ornithonyssus pavlovskii.—Zemskaya, 1966: 50.
Ornithonyssus pipistrelli.—Evans & Till, 1966: 326; Micherdziński, 1980: 104.

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Liponyssus pipistrelli.—Radovsky, 1967: 209.
Cryptonyssus pipistrelli.—Radovsky, 1969: 462; 2010: 93.

Occurrence and habitat. Although Ornithonyssus pipistrelli has been widely reported from a variety of
vespertilionid bats (Stanyukovich 1997; Baker & Craven 2003), no Slovak specimens were found in association
with these hosts (see host distribution). It has been collected at altitudes between 113 and 760 m a.s.l.
Host distribution. Bird nests: Columba livia f. domestica (Fenďa & Kicková 2009), Emberiza citrinella,
Nannus troglodytes, Parus sp., Passer montanus (Krumpál et al. 2001, Švaňa et al. 2006, Fenďa et al. 2011), Sitta
europaea (Fenďa & Mašán 2003), Turdus sp. Combined nests: Cyanistes caeruleus and Muscardinus avellanarius,
Parus sp. and Apodemus flavicollis. Mammal nests: Canis familiaris, Glis glis (Mašán & Krištofík 1996),
Micromys minutus, Muscardinus avellanarius.

New data. Branisko Mts.: 9 ♀♀, 21 June 1989, Bijacovce Village, Branisko Mountain Pass, Muscardinus avellanarius.
Košická Kotlina Basin: 44 ♀♀, 3 ♂♂, 1 N, 22 October 1987, Moldava Nad Bodvou Village, Paňovský Les Forest, Cyanistes
caeruleus + Muscardinus avellanarius; 3 ♀♀, 22 October 1987, Moldava Nad Bodvou Village, Paňovský Les Forest, Parus sp.
+ Apodemus flavicollis; 1 ♀, 12 NN, 22 October 1987, Moldava Nad Bodvou Village, Paňovský les Forest, Muscardinus
avellanarius. Malé Karpaty Mts.: 1 N, 10 February 1986, Bratislava Capital, Vajnorská Dolina Valley, Turdus sp.; 1 ♀, 1 N, 28
July 1992, Bratislava Capital, Železná Studnička Forest, Nannus troglodytes. Podunajská Rovina Lowland: 1 N, 17 May 1987,
Sap Village, Erced Forest, Emberiza citrinella; 1 ♀, 25 July 1992, Svätý Jur Village, Šúr Nature Reserve, Micromys minutus.
Veľká Fatra Mts.: 2 ♀♀, 1 N, 24 May 1985, Stankovany Village, Aves sp.; 2 NN, 22 June 1989, Stankovany Village, Parus sp.;
1 N, 22 June 1989, Stankovany Village, Canis familiaris.

Ornithonyssus sylviarum (Canestrini & Fanzago, 1877)

Dermanyssus sylviarum Canestrini & Fanzago, 1877: 124.


Liponyssus americanus Banks, 1905: 136. Synonymy by Strandtmann & Wharton (1958).
Liponyssus canadensis Banks, 1909: 134. Synonymy by Strandtmann & Wharton (1958).
Liponyssus pacificus Ewing, 1923: 19. Synonymy by Strandtmann & Wharton (1958).
Bdellonyssus ondatrae Willmann, 1952: 410. Synonymy by Micherdziński (1980).
Ornithonyssus banksi Strandtmann & Wharton, 1958: 85. Synonymy by Whitaker & Wilson (1974).
Ornithonyssus sylviarum.—Evans & Till, 1966: 318; Micherdziński, 1980: 115.
Ornithonyssus sylviarum utrechtensis Micherdziński, 1980: 318. Synonymy by Radovsky & Estébanes-Gonzáles (2001).

Occurrence and habitat. This cold tolerant species is distributed mainly in the north temperate zone, and in
Slovakia it is the commonest dermanyssoid associate of birds. It is very frequently and abundantly found in many
bird taxa, including species that build various types of nests and occur in diverse habitats. It is very widely
distributed in Slovakia, ranging from lowlands to the alpine zone of mountains (2,120 m a.s.l.). Ornithonyssus
sylviarum is reported here from 71 identified hosts representing 14 orders of Aves (as listed below). It does not
seem to be host-specific for any of the bird species or their higher taxa. It shows only a certain degree of positive
habitat preference at the level of an ecological guild associated with riparian biotopes, due to its apparent affinity
for the nests of the passeriform bird communities nesting in sites with littoral reed vegetation and locally high air
humidity (e.g., Acrocephalus arundinaceus, Emberiza schoeniclus, Notiocichla spp., and Panurus biarmicus). It is
collected also in a variety of microhabitats not directly associated with the bodies and nests of birds and mammals,
e.g., hygrophilous vegetation, moss, organic detritus of herbs, soil, litter, and substrates taken from a cave
environment.
Host distribution. Bird nests: Accipiter gentilis, Acrocephalus arundinaceus (Krištofík et al. 2001, Fenďa &
Schniererová 2004), Acrocephalus arundinaceus with Cuculus canorus, Aegithalos caudatus, Alauda arvensis,
Alcedo atthis, Anas platyrhynchos (Fenďa & Schniererová 2005), Anthus pratensis, Anthus spinoletta, Anthus
trivialis, Aquila chrysaetos, Aythya ferina, Bubo bubo, Buteo buteo, Carduelis carduelis, Certhia sp., Chloris
chloris (Ambros et al. 1992), Chroicocephalus ridibundus, Ciconia ciconia, Circus aeruginosus, Circus pygargus,
Corvus frugilegus, Cygnus olor, Delichon urbicum, Dendrocopos major, Emberiza schoeniclus, Falco cherrug,
Falco tinnunculus, Fringilla coelebs, Fulica atra, Garrulus glandarius (Mrciak & Rosický 1956: as Dermanyssus
sylviarum), Glaucidium passerinum, Himantopus himantopus, Hippolais icterina (Ambros et al. 1992), Ictinaetus
pomarinus (Krištofík et al. 2009), Ixobrychus minutus (Fenďa & Schniererová 2005), Lanius collurio (Ambros et

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al. 1992, Krištofík et al. 2002), Lanius minor (Krištofík et al. 2002), Luscinia megarhynchos, Merops apiaster
(Krištofík et al. 1996, Fenďa 2008), Milvus migrans, Milvus milvus, Motacilla alba (Ambros et al. 1992),
Motacilla cinerea, Motacilla cinereocaphilla, Motacilla feldegg, Motacilla flava, Muscicapa striata, Nannus
troglodytes, Notiocichla palustris (Krištofík et al. 2005), Notiocichla scirpacea (Fenďa & Schniererová 2004,
2005), Oriolus oriolus, Panurus biarmicus (Krištofík et al. 2007), Parus major, Parus sp. (Fenďa & Mašán 2003),
Passer montanus (Krumpál et al. 2001, Švaňa et al. 2006, Fenďa et al. 2011), Phoenicurus ochruros, Phylloscopus
collybita, Phylloscopus sibilatrix, Pica pica, Podiceps cristatus, Prunella modularis, Remiz pendulinus (Ambros et
al. 1992, Krištofík et al. 1993, Krištofík & Mašán 1996, Mašán 1997), Riparia riparia (Mašán & Krištofík 1993,
Fenďa 2008), Serinus serinus, Sitta europaea (Fenďa & Mašán 2003), Sylvia atricapilla, Sylvia nisoria, Sylvia sp.,
Turdus merula, Turdus philomelos (Micherdziński 1980), Turdus pilaris, Upupa epops (Krištofík et al. 2013).
Mammals: Apodemus flavicollis (Mrciak 1963), Glis glis (Mašán & Krištofík 1996), Mus musculus (Mrciak 1963).
Other findings: leaves of Phragmites australis (Fenďa & Schniererová 2004, 2005), soil (Fenďa & Mašán 2003,
Fenďa & Ciceková 2005), litter, moss, cave habitat.

New data. Borská Nížina Lowland: 1 ♀, 15 May 1991, Vysoká Pri Morave Village (floodplain forest), Aegithalos caudatus; 1
♀, 10 June 1991, Vysoká Pri Morave Village (floodplain forest), Certhia sp.; 223 ♀♀, 1 N, 27 June 1991, Jakubov Village,
Jakubovské Rybníky Fishponds, Serinus serinus; 20 ♀♀, 8 July 1991, Vysoká Pri Morave Village (floodplain forest), Sylvia
atricapilla; 230 ♀♀, 3 ♂♂, 10 NN, 8 July 1991, Vysoká Pri Morave Village (floodplain forest), Turdus merula; 3 ♀♀, 16
August 1991, Malé Leváre Village (floodplain forest), Milvus migrans; 1 ♀, 1 N, 13 May 1992, Vysoká Pri Morave Village
(floodplain forest), Accipiter gentilis; 1 ♀, 4 June 1992, Malé Leváre Village (canal), Cygnus olor; 23 ♀♀, 5 August 1992,
Vysoká Pri Morave Village (meadow), Anthus trivialis; 4,880 ♀♀, 341 ♂♂, 989 NN, 7 June 1993, Bratislava Capital, Devínske
Jazero Settlement, Motacilla flava; 1 ♀, 9 June 1993, Bratislava Capital, Devínske Jazero Settlement, Turdus merula; 22 ♀♀,
17 June 1993, Bratislava Capital, Devínske Jazero Settlement, Motacilla feldegg; 95 ♀♀, 3 NN, 26 July 1993, Jakubov Village,
Jakubovské Rybníky Fishponds, Ixobrychus minutus; 6 ♀♀, 2 August 1993, Bratislava Capital, Devínske Jazero Settlement,
Falco tinnunculus; 2 ♀♀, 12 August 1993, Moravský Svätý Ján Village (forest), Buteo buteo; 1 ♀, 28 June 1994, Vysoká Pri
Morave Village (floodplain forest), Phylloscopus sibilatrix; 1,152 ♀♀, 13 ♂♂, 28 NN, 18 July 1991. Kysucké Beskydy Mts.: 1
♀, 4 June 1987, Oščadnica Village, Parus major; 1 ♀, 1 July 1997, Nová Bystrica Village, Chmúra Valley (moss). Malá Fatra
Mts.: 1 ♀, 21 August 1992, Aquila chrysaetos; 56 ♀♀, 1 N, 15 June 1995, Párnica Village, Osnica Mt., Anthus spinoletta; 4
♀♀, 23 June 1995, Terchová Village, Stoh Mt., Anthus pratensis. Malé Karpaty Mts.: 96 ♀♀, 30 August 1990, Bratislava
Capital, Železná Studnička Forest, Alcedo atthis; 102 ♀♀, 24 June 1991, Bratislava Capital, Železná Studnička Forest,
Prunella modularis; 24 ♀♀, 25 June 1991, Falco cherrug; 21 ♀♀, 25 June 1991, Milvus milvus; 7 ♀♀, 23 July 1991,
Bratislava Capital, Železná Studnička Forest, Motacilla cinerea; 4 ♀♀, 23 July 1991, Bratislava Capital, Železná Studnička
Forest, Phoenicurus ochruros; 6 ♀♀, 23 July 1991, Bratislava Capital, Železná Studnička Forest, Nannus troglodytes. Oravská
Vrchovina Highland: 3 ♀♀, 21 July 1989, Nižná Na Orave Village, Turdus merula. Podunajská Rovina Lowland: 2 ♀♀, 5 April
1989, Pataš Village, Corvus frugilegus; 3 ♀♀, 4 July 1989, Dobrohošť Village (floodplain forest), Muscicapa striata; 864 ♀♀,
3 ♂♂, 15 NN, 23 July 1989, Baka Village (floodplain forest), Hippolais icterina; 11 ♀♀, 1 ♂, 3 August 1989, Bodíky Village,
Ciconia ciconia; 127 ♀♀, 1 ♂, 5 NN, 6 August 1990, Čierna Voda Village, Luscinia megarhynchos; 2,818 ♀♀, 120 ♂♂, 1,238
NN, 17 August 1990, Dobrohošť Village (floodplain forest), Carduelis carduelis; 37 ♀♀, 2 ♂♂, 4 September 1990, Komárno
Town, Apáli Nature Reserve, Dendrocopos major; 1 ♀, 24 May 1991, Komárno Town, Apáli Nature Reserve, Phylloscopus
collybita; 29 ♀♀, 25 June 1991, Veľké Blahovo Village, Veľkoblahovské Rybníky Fishponds, Aythya ferina; 286 ♀♀, 70 ♂♂,
122 NN, 25 June 1991, Veľké Blahovo Village, Veľkoblahovské Rybníky Fishponds, Fulica atra; 68 ♀♀, 25 June 1991, Veľké
Blahovo Village, Veľkoblahovské Rybníky Fishponds, Podiceps cristatus; 33 ♀♀, 3 NN, 3 July 1991, Dobrohošť Village
(floodplain forest), Chloris chloris; 6,413 ♀♀, 882 ♂♂, 2,580 NN, 3 July 1991, Dobrohošť Village (floodplain forest), Turdus
philomelos; 6 ♀♀, 12 July 1991, Kľúčovec Village (floodplain forest), Fringilla coelebs; 14 ♀♀, 23 July 1991, Dolný Štál
Village, Hroboňovské Rybníky Fishponds, Chroicocephalus ridibundus; 148 ♀♀, 2 NN, 14 August 1991, Dolný Štál Village,
Hroboňovské Rybníky Fishponds, Acrocephalus arundinaceus + Cuculus canorus; 1 ♀, 10 September 1991, Veľké Blahovo
Village, Veľkoblahovské Rybníky Fishponds, Circus aeruginosus; 1 ♀, 23 June 1992, Vojka Nad Dunajom Village (canal),
Himantopus himantopus; 2 ♀♀, 15 July 1993, Lehnice Village (windbreak), Turdus merula; 3 ♀♀, 27 July 1993, Dolný Štál
Village, Hroboňovské Rybníky Fishponds, Oriolus oriolus; 6 ♀♀, 14 September 1993, Gabčíkovo Village, Delichon urbicum;
1 ♀, 25 June 1994, Bratislava Capital, Ostrov Kopáč Nature Reserve, Sylvia nisoria; 4 ♀♀, 3 July 1994, Dolný Štál Village,
Hroboňovské Rybníky Fishponds, Motacilla alba; 2 ♀♀, 25 November 1994, Dobrohošť Village, Dunajské Kriviny
(floodplain forest), Glis glis; 2,239 ♀♀, 184 ♂♂, 560 NN, 30 May 1995, Dolný Štál Village, Hroboňovské Rybníky Fishponds,
Emberiza schoeniclus; 29 ♀♀, 3 ♂♂, 2 June 1995, Bratislava Capital, Ostrov Kopáč Nature Reserve, Alauda arvensis; 4 ♀♀,
25 June 1995, Dolný Štál Village, Hroboňovské Rybníky Fishponds, Pica pica; 2 ♀♀, 23 June 1997, Číčov Village, Číčovské
Mŕtve Rameno Nature Reserve, Ixobrychus minutus; 4 ♀♀, 29 June 1999, Šamorín Village (windbreak), Falco cherrug.
Považské Podolie Basin: 1 ♀, 13 August 1989, Dubnica Nad Váhom Town, Parus major. Považský Inovec Mts.: 44 ♀♀, 7 July
1991, Hubina Village, Bubo bubo. Slovenský Raj Mts.: 1 ♀, 13 August 1999, Stratená Village, Koniarova Jaskyňa Cave (bait);
4 ♀♀, 11 July 2012, Stratená Village, Duča Cave (soil and litter). Starohorské Vrchy Mts.: 2 ♀♀, 20 August 1990, Staré Hory
Village, Sylvia sp. Strážovské Vrchy Mts.: 1 ♀, 5 July 1999, Malinová Village, Ictinaetus pomarinus; 4 ♀♀, 19 July 1999,

SLOVAKIAN DERMANYSSOIDEA ASSOCIATED WITH BIRDS & NESTS Zootaxa 3893 (1) © 2014 Magnolia Press · 87
Chvojnica Village, Ictinaetus pomarinus. Turčianska Kotlina Basin: 1 ♀, 11 June 2000, Diviaky Village, Motacilla
cinereocaphilla. Veľká Fatra Mts.: 2 ♀♀, 26 June 1993, Stankovany Village, Podšíp Settlement, Sitta europaea. Veporské
Vrchy Mts.: 91 ♀♀, 5 ♂♂, 2 NN, 6 July 1993, Hriňová Village, Pivnička Settlement, Turdus pilaris. Vihorlatské Vrchy Mts.: 8
♀♀, 17 July 1999, Ptičie Village, Ictinaetus pomarinus. Východoslovenská Rovina Lowland: 28 ♀♀, 4 August 1993, Blatné
Revištia Village, Circus pygargus. Vysoké Tatry Mts.: 1 ♀, 13 July 2005, Tatranská Javorina Village, Zadná Javorová Dolina
Valley (organic plant detritus under stone). Zvolenská Kotlina Basin: 49 ♀♀, 12 July 1993, Kováčová Village (forest),
Glaucidium passerinum.

Genus: Pellonyssus Clark & Yunker, 1956

Pellonyssus Clark & Yunker, 1956: 93. Type species: Pellonyssus passeri Clark & Yunker, 1956.

According to Radovsky (2010) the genus Pellonyssus contains 13 valid species, all parasites of birds, and widely
distributed mainly in warmer latitudes. He did not record any occurrence of this genus in Europe, but there are at
least four papers published on European records of Pellonyssus from the nests of the house sparrow (Passer
domesticus) in Hungary and the Eurasian tree sparrow (Passer montanus) in Slovakia (Szabó et al. 2002, 2008;
Švaňa et al. 2006; Fenďa et al. 2011).

Pellonyssus reedi (Zumpt & Patterson, 1952)

Steatonyssus reedi Zumpt & Patterson, 1952: 163.


Steatonyssus malurus Womersley, 1956: 214. Synonymy by Till (1964).
Pellonyssus passeri Clark & Yunker, 1956: 94. Synonymy by Till (1964).
Steatonyssus stenosternus Wang, 1963: 54. Synonymy by Domrow (1966).
Pellonyssus nidicolus Baker, Delfinado & Abbatiello, 1976: 56. Synonymy by Radovsky & Estébanes-Gonzáles (2001).
Pellonyssus reedi.—Micherdziński, 1980: 190; Radovsky & Estébanes-Gonzáles, 2001: 20.

Taxonomic remarks. Our specimens are in very good agreement with those described by Clark & Yunker (1956)
under the name Pellonyssus passeri and later synonymised with Pellonyssus reedi by Till (1964). The female of the
species is recognised by the following character states (1) peritremes reaching the mid-level of coxae II; (2) coxae
II with a distinctive spur on its anterior margin; (3) basitarsi IV with an elongated setae pd3; (4) sternal shield
reduced in length, subcrescentic; (5) sternal setae st1 short, subequal with anterio-posterior length of the sternal
shield; (6) podonotal shield with ten pairs of setae and opisthonotal shield with six pairs of setae.
Occurrence and habitat. Pellonyssus reedi seems to be host-specific for sparrows, Passer spp., but it has also
occasionally been recorded in the nests of other bird species (Micherdziński 1980). In Slovakia, it was found as a
nest associate of the water pipit (Anthus spinoletta) and Eurasian tree sparrow (Passer monatnus), in collection
sites at altitudes between 130 and 1,650 m a.s.l. Our detection of one heavily infested sparrow’s nest with 2,598
specimens of this mite confirms its high reproductive ability on this host. Szabó et al. (2008) found 12,213
individuals of P. reedi on 188 young sparrows in Hungary.
Host distribution. Bird nests: Anthus spinoletta, Passer montanus (Švaňa et al. 2006, as Pellonyssus sp.,
Fenďa et al. 2011, as Pellonyssus aff. reedi).

New data. Malá Fatra Mts.: 1 ♂, 9 July 2008, Turany Village, Snilovské Sedlo Mountain Pass, Anthus spinoletta; 2 ♀♀, 9 July
2008, Turany Village, Veľký Kriváň Mt., Anthus spinoletta.

Dubious species

Dermanyssus longipes Berlese &Trouessart, 1889

Dermanyssus longipes Berlese & Trouessart, 1889: 128.

Published record. Trnavská Pahorkatina Hills: 8 ♀♀, 12 August 1982, Budmerice Village, Passer domesticus (Roy et al.
2009a). We believe this record is based on a misidentification of D. hirundinis.

88 · Zootaxa 3893 (1) © 2014 Magnolia Press MAŠÁN ET AL.


Remarks. The original description of Dermanyssus longipes by Berlese & Trouessart (1889) is not adequate for
accurate recognition of this species (type locality "France"). The type specimens deposited in the Berlese
Collection at Florence are not suitable for a detailed study (Evans & Till 1962; Moss 1978), so most modern
authors have considered this species as a “species incertae sedis” and not included it in their keys to identification.
However, Roy et al. (2009a) assigned some specimens to D. longipes on the basis of two morphological features
(anal shield "slightly more" elongated and subrectangular, dorsal shield with a subapical shrinkage and rounded
posterior margin) and geographic locality. Roy et al. (2009a) based their concept of this species on specimens from
France, and from M. Mrciak’s specimens from Slovakia, in the collection of P. Fenďa at Comenius University. We
have examined the Mrciak specimens (labelled as Dermanyssus passerinus), and can confirm that they belong to
D. hirundinis. Therefore we do not follow the concept of D. longipes as given by Roy et al. (2009a).

Dermanyssus passerinus Berlese & Trouessart, 1889

Dermanyssus passerinus Berlese & Trouessart, 1889: 127.

Published records. Malé Karpaty Mts.: 982 specimens., 10 November 1958, Trstín Village, Ficedula albicollis; 3 specimens.,
ibid., Parus major; 6 specimens., ibid., Sturnus vulgaris; 56 specimens., ibid., Ficedula albicollis + Sturnus vulgaris (Nosek &
Lichard 1962).

Remarks. As for the previous species, Dermanyssus passerinus is usually referred to as "incertae sedis" due to the
inadequate description and damaged type material (Evans & Till 1962; Moss 1978). The voucher specimens of
Nosek & Lichard (1962) are presumably lost and not available for revision, therefore they cannot be taken into
consideration. They are probably misidentified specimens of Dermanyssus hirundinis and/or D. carpathicus.

Key to genera and species of Dermanyssidae and Macronyssidae associated with birds occurring in Slo-
vakia (females)

1. Chelicerae fine and conspicuously elongated, stylet-like, almost subequal in length to legs I; cheliceral digits minute, hardly
discernible. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Dermanyssus Dugès, 1834. . . . . . . . . . . . . . . . . . . . . . . . . . 2
- Chelicerae at least moderately thick, much shorter than legs I; cheliceral digits well developed, prominent . . . . . . . . . . . . . . 6
2. Metasternal setae st4 absent; dorsal shield slightly shortened posteriorly, with two pairs of J-setae on opisthonotal surface (J1
and J3); soft integument with two pairs of small suboval platelets close to posterior margin of dorsal shield . . . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Dermanyssus americanus Ewing, 1923
- Metasternal setae st4 present; dorsal shield not reduced in length, with at least three pairs of J-setae on opisthonotal surface (J1,
J3 and J4 always present); soft integument never with small scutal elements close to posterior margin of dorsal shield . . . . 3
3. Dorsal shield with four pairs of J setae on opisthonotal surface (J2 present), and with at least nine pairs of setae on podonotal
surface (z6 present) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 4
- Dorsal shield with three pairs of J setae on opisthonotal surface (J2 absent), and with eight pairs of setae on podonotal surface
(z6 absent or placed on soft integument) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 5
4. Anterior end of peritremes reaching mid-level of coxae II; dorsal shield with eleven pairs of opisthonotal setae (j1 and s1 pres-
ent); genu IV with one posterolateral seta. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Dermanyssus gallinae (De Geer, 1778)
- Peritremes strongly reduced in length and with anterior end reaching to mid-level of coxae III; dorsal shield with nine pairs of
opisthonotal setae (j1 and s1 off the shield); genu IV with two posterolateral setae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Dermanyssus chelidonis Oudemans, 1939
5. Tibia I, genu I and femur II with two ventral setae; trochanter I without dorsal seta; femur II with one anterolateral seta;
humeral platelets fused to or adjacent to anterolateral margins of dorsal shield; idiosomal setae placed on soft caudal integu-
ment longer, more robust . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Dermanyssus carpathicus Zeman, 1979
- Tibia I, genu I and femur II with three ventral setae; trochanter I with one dorsal seta; femur II with two anterolateral setae;
humeral platelets well distant from anterolateral margins of dorsal shield; idiosomal setae placed on soft caudal integument
shorter, relatively thin . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Dermanyssus hirundinis (Hermann, 1804)
6. Dorsal shield divided, with subequal podonotal and opisthonotal portions . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Pellonyssus Clark & Yunker, 1956. . . . . . Pellonyssus reedi Zumpt & Patterson, 1952
- Dorsal shield undivided, entire . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Ornithonyssus Sambon, 1928. . . . . 7
7. Dorsal shield setae j3 present; anterior end of peritremes reaching mid-level of coxae II; sternal setae st1 shorter than setae st2
and st3 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Ornithonyssus pipistrelli Oudemans, 1903
- Dorsal shield setae j3 absent; anterior end of peritremes reaching posterior margin of coxae I; sternal setae st1 at least subequal

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to setae st2 and st3 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 8
8. Dorsal shield behind J3 with three pairs of setae (Z4, Z5 and Jx) . . . . Ornithonyssus sylviarum (Canestrini & Fanzago, 1877)
- Dorsal shield behind J3 with four pairs of setae (Z4, Z5, S5 and Jx) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 9
9. J setae relatively longer, J1 with tip reaching the following setae J2; opisthogastric soft integument densely setose . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Ornithonyssus bacoti (Hirst, 1913)
- J setae relatively shorter, J1 with tip reaching between J1 and J2; opisthogastric soft integument sparsely setose . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Ornithonyssus bursa (Berlese, 1888)

Discussion

Dermanyssoid mites include a large number of parasitic genera and are common in the nests and on the bodies of
various vertebrates. The change in morphology from chelate-dentate chelicerae towards more slender and elongate
piercing forms is a characteristic adaptation of these mites for penetrating the skin and regular feeding on blood
(Radovsky 1985). Generally, obligate blood feeders are relatively more frequent and abundant on the bodies of
their hosts than in their nests, and that is true for bird-associate parasites that show a high level of host specificity.
That may explain why we did not collect several strictly monoxenous European species in this survey.
Dermanyssus alaudae (Schrank) is a parasite of the Eurasian skylark Alauda arvensis, D. apodis Roy et al. is a
parasite of the common swift Apus apus, D. quintus Vitzthum and D. grochovskae Zemskaya are both parasites of
woodpeckers (Piciformes). We examined several nests of each of these hosts, but did not find these species of
Dermanyssus. The most common Slovak species of Dermanyssus tend to have low host specificity (D. carpathicus,
D. gallinae, D. hirundinis, and Ornithonyssus sylviarum), while others are found on a narrower range of hosts (D.
americanus, D. chelidonis, O. bursa, and Pellonyssus reedi).
According to Roy et al. (2009a), Dermanyssus species morphologically fall into two groups, those possessing
a compact and more heavily sclerotised integument with shorter and stouter legs (D. americanus), and those having
a soft body adapted for sporadic and large engorgement with reduced shields and slender legs (all other species
found in Slovakia). We agree with Roy et al. (2009a) that the second species group is more common and has wider
host ranges than the more host-specific species in the first group.
The host distribution of mites based on the records from Slovakia includes 100 identified species of Aves
representing 17 orders: Accipitriformes, Anseriformes, Apodiformes, Ardeiformes, Bucerotiformes, Charadriiformes,
Ciconiiformes, Columbiformes, Coraciiformes, Cuculiformes, Falconiformes, Galliformes, Gruiformes,
Passeriformes, Piciformes, Podicipediformes, and Strigiformes (see the appendix below). For example, Slovak
populations of the most common species of the genus Dermanyssus, D. carpathicus, were collected from the nests
of 12 different orders of Aves, while D. chelidonis was collected only from the nests of three passerine host species.
This is not consistent with the statement of Roy et al. (2009a), based on much more limited sampling, who found
D. carpathicus to be rather host specific in France (found only in the nests of two genera, namely Parus sp. and
Phoenicurus sp.). The most common Slovak species, Ornithonyssus sylviarum, Dermanyssus carpathicus, D.
hirundinis and D. gallinae were found in association with 71 (14 orders), 58 (12), 41 (9 orders), and 30 (10 orders)
bird species, respectively. The occurrence and findings of other mite species were quite sporadic.

Acknowledgments

We are grateful to many colleagues who have, in various ways, helped us to write this paper. We thank all
collectors of the bird nests examined in this study, and especially ornithologist Dr. A. Darolová (Institute of
Zoology, Bratislava), also for providing the data on identification of the bird hosts. We also thank Lise Roy for her
very useful conversations about some of our findings. This study was fully supported by the Scientific Grant
Agency of the Ministry of Education of Slovak Republic and the Academy of Sciences [VEGA Grant No. 2/0012/
11: Dermanyssoid mites (Acari, Mesostigmata) associated with birds and their nests (Aves) in Slovakia, with
consideration on taxonomy, ecology and chorology of individual species.].

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APPENDIX. Host distribution summary (based on records from Slovakia):

Class: Aves
Order: Accipitriformes Vieillot, 1816
Accipiter gentilis (Linnaeus, 1758), Northern Goshawk
Dermanyssus hirundinis, Ornithonyssus sylviarum
Aquila chrysaetos (Linnaeus, 1758), Golden Eagle
Dermanyssus carpathicus, Ornithonyssus sylviarum
Aquila heliaca Savigny, 1809, Eastern Imperial Eagle
Dermanyssus sp.
Buteo buteo (Linnaeus, 1758), Common Buzzard

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Dermanyssus sp., Ornithonyssus sylviarum
Circus aeruginosus (Linnaeus, 1758), Western Marsh Harrier
Dermanyssus carpathicus, Dermanyssus gallinae, Dermanyssus hirundinis, Ornithonyssus sylviarum
Circus pygargus (Linnaeus, 1758), Montagu's Harrier
Ornithonyssus sylviarum
Ictinaetus pomarinus C. L. Brehm, 1831, Lesser Spotted Eagle
Dermanyssus hirundinis, Ornithonyssus sylviarum
Milvus migrans (Boddaert, 1783), Black Kite
Dermanyssus hirundinis, Ornithonyssus sylviarum
Milvus milvus (Linnaeus, 1758), Red Kite
Ornithonyssus sylviarum

Order: Anseriformes Wagler, 1831


Anas crecca Linnaeus, 1758, Eurasian Teal
Dermanyssus hirundinis
Anas platyrhynchos Linnaeus, 1758, Mallard
Dermanyssus carpathicus, Dermanyssus gallinae, Dermanyssus hirundinis, Ornithonyssus sylviarum
Anser anser (Linnaeus, 1758), Greylag Goose
Dermanyssus hirundinis
Aythya ferina (Linnaeus, 1758), Common Pochard
Dermanyssus hirundinis, Ornithonyssus sylviarum
Aythya fuligula (Linnaeus, 1758), Tufted Duck
Dermanyssus hirundinis
Cygnus olor (Gmelin, 1789), Mute Swan
Dermanyssus gallinae, Ornithonyssus sylviarum

Order: Apodiformes Peters, 1940


Apus apus (Linnaeus, 1758), Common Swift
Ornithonyssus sp.

Order: Ardeiformes Wagler, 1831


Ixobrychus minutus (Linnaeus, 1766), Little Bittern
Ornithonyssus sylviarum

Order: Bucerotiformes Fürbringer, 1888


Upupa epops Linnaeus, 1758, Eurasian Hoopoe
Ornithonyssus sylviarum

Order: Charadriiformes Huxley, 1867


Chroicocephalus ridibundus (Linnaeus, 1766), Black-headed Gull
Dermanyssus carpathicus, Dermanyssus hirundinis, Ornithonyssus sylviarum
Himantopus himantopus (Linnaeus, 1758), Black-winged Stilt
Dermanyssus carpathicus, Ornithonyssus sylviarum
Limosa limosa (Linnaeus, 1758), Black-tailed Godwit
Dermanyssus carpathicus
Numenius arquata (Linnaeus, 1758), Eurasian Curlew
Dermanyssus carpathicus
Vanellus vanellus (Linnaeus, 1758), Northern Lapwing
Dermanyssus carpathicus

Order: Ciconiiformes Garrod, 1874


Ciconia ciconia (Linnaeus, 1758), White Stork
Dermanyssus carpathicus, Dermanyssus hirundinis, Ornithonyssus sylviarum

Order: Columbiformes Latham, 1790


Columba livia f. domestica Gmelin, 1789, Domestic Pigeon
Dermanyssus gallinae, Ornithonyssus pipistrelli

Order: Coraciiformes Forbes, 1884


Alcedo atthis (Linnaeus, 1758), Common Kingfisher
Dermanyssus carpathicus, Ornithonyssus sylviarum

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Coracias garrulus Linnaeus, 1758, European Roller
Dermanyssus gallinae, Dermanyssus hirundinis
Merops apiaster Linnaeus, 1758, European Bee-eater
Dermanyssus carpathicus, Dermanyssus gallinae, Dermanyssus hirundinis, Ornithonyssus sylviarum

Order: Cuculiformes Wagler, 1830


Cuculus canorus Linnaeus, 1758, Common Cuckoo
Dermanyssus carpathicus, Ornithonyssus sylviarum

Order: Falconiformes Bonaparte, 1831


Falco cherrug J. E. Gray, 1834, Saker Falcon
Dermanyssus carpathicus, Ornithonyssus sylviarum
Falco tinnunculus Linnaeus, 1758, Common Kestrel
Dermanyssus carpathicus, Dermanyssus gallinae, Dermanyssus hirundinis, Ornithonyssus sylviarum
Falco vespertinus Linnaeus, 1766, Red-footed Falcon
Dermanyssus hirundinis

Order: Galliformes Temminck, 1820


Gallus gallus var. domesticus (Linnaeus, 1758), Domestic Chicken
Dermanyssus gallinae

Order: Gruiformes Bonaparte, 1854


Fulica atra Linnaeus, 1758, Eurasian Coot
Dermanyssus carpathicus, Dermanyssus gallinae, Dermanyssus hirundinis, Ornithonyssus sylviarum

Order: Passeriformes Linnaeus, 1758


Acrocephalus arundinaceus (Linnaeus, 1758), Great Reed Warbler
Dermanyssus carpathicus, Dermanyssus gallinae, Dermanyssus hirundinis, Ornithonyssus sylviarum
Aegithalos caudatus (Linnaeus, 1758), Long-tailed Tit
Dermanyssus carpathicus, Ornithonyssus sylviarum
Alauda arvensis Linnaeus, 1758, Eurasian Skylark
Dermanyssus carpathicus, Ornithonyssus sylviarum
Anthus pratensis (Linnaeus, 1758), Meadow Pipit
Ornithonyssus sylviarum
Anthus spinoletta (Linnaeus, 1758), Water Pipit
Dermanyssus hirundinis, Ornithonyssus sylviarum, Pellonyssus reedi
Anthus trivialis (Linnaeus, 1758), Tree Pipit
Dermanyssus carpathicus, Ornithonyssus sylviarum
Carduelis carduelis (Linnaeus, 1758), European Goldfinch
Dermanyssus carpathicus, Dermanyssus hirundinis, Ornithonyssus sylviarum
Certhia brachydactyla C. L. Brehm, 1820, Short-toed Treecreeper
Dermanyssus carpathicus
Certhia sp., Treecreeper
Ornithonyssus sylviarum
Chloris chloris (Linnaeus, 1758), European Greenfinch
Dermanyssus carpathicus, Ornithonyssus sylviarum
Cinclus cinclus (Linnaeus, 1758), White-throated Dipper
Dermanyssus sp.
Corvus frugilegus Linnaeus, 1758, Rook
Dermanyssus gallinae, Ornithonyssus sylviarum
Cyanistes caeruleus (Linnaeus, 1758), Eurasian Blue Tit
Dermanyssus hirundinis, Ornithonyssus pipistrelli
Delichon urbicum (Linnaeus, 1758), Common House Martin
Dermanyssus carpathicus, Dermanyssus chelidonis, Dermanyssus gallinae, Dermanyssus hirundinis, Ornithonyssus
sylviarum
Emberiza citrinella Linnaeus, 1758, Yellowhammer
Ornithonyssus pipistrelli
Emberiza schoeniclus (Linnaeus, 1758), Common Reed Bunting
Dermanyssus carpathicus, Ornithonyssus sylviarum
Erithacus rubecula (Linnaeus, 1758), European Robin
Dermanyssus hirundinis

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Ficedula albicollis (Temminck, 1815), Collared Flycatcher
Dermanyssus carpathicus, Dermanyssus gallinae, Dermanyssus hirundinis
Fringilla coelebs Linnaeus, 1758, Common Chaffinch
Dermanyssus carpathicus, Dermanyssus gallinae, Ornithonyssus sylviarum
Garrulus glandarius (Linnaeus, 1758), Eurasian Jay
Ornithonyssus sylviarum
Hippolais icterina (Vieillot, 1817), Icterine Warbler
Ornithonyssus sylviarum
Hirundo rustica Linnaeus, 1758, Barn Swallow
Dermanyssus gallinae, Dermanyssus hirundinis, Ornithonyssus bursa
Lanius collurio Linnaeus, 1758, Red-backed Shrike
Dermanyssus carpathicus, Dermanyssus gallinae, Ornithonyssus sylviarum
Lanius minor Gmelin, 1788, Lesser Grey Shrike
Dermanyssus carpathicus, Dermanyssus hirundinis, Ornithonyssus sylviarum
Linaria cannabina (Linnaeus, 1758), Common Linnet
Dermanyssus carpathicus, Dermanyssus chelidonis
Locustella luscinioides (Savi, 1824), Savi's Warbler
Dermanyssus carpathicus
Luscinia megarhynchos C. L. Brehm, 1831, Common Nightingale
Dermanyssus hirundinis, Ornithonyssus sylviarum
Motacilla alba Linnaeus, 1758, White Wagtail
Dermanyssus carpathicus, Dermanyssus gallinae, Dermanyssus hirundinis, Ornithonyssus sylviarum
Motacilla cinerea Tunstall, 1771, Grey Wagtail
Dermanyssus carpathicus, Dermanyssus hirundinis, Ornithonyssus sylviarum
Motacilla cinereocaphilla Savi, 1831, Ashy-headed Wagtail
Ornithonyssus sylviarum
Motacilla feldegg Michahelles, 1830, Black-headed Wagtail
Ornithonyssus sylviarum
Motacilla flava Linnaeus, 1758, Western Yellow Wagtail
Dermanyssus carpathicus, Ornithonyssus sylviarum
Muscicapa striata (Pallas, 1764), Spotted Flycatcher
Dermanyssus carpathicus, Ornithonyssus sylviarum
Nannus troglodytes (Linnaeus, 1758), Eurasian Wren
Dermanyssus carpathicus, Dermanyssus gallinae, Dermanyssus hirundinis, Ornithonyssus pipistrelli, Ornithonyssus
sylviarum
Notiocichla palustris (Bechstein, 1798), Marsh Warbler
Dermanyssus carpathicus, Ornithonyssus sylviarum
Notiocichla scirpacea (Hermann, 1804), Eurasian Reed Warbler
Dermanyssus carpathicus, Ornithonyssus sylviarum
Oriolus oriolus (Linnaeus, 1758), Eurasian Golden Oriole
Dermanyssus carpathicus, Ornithonyssus sylviarum
Panurus biarmicus (Linnaeus, 1758), Bearded Reedling
Ornithonyssus sylviarum
Parus major Linnaeus, 1758, Great Tit
Dermanyssus carpathicus, Dermanyssus gallinae, Dermanyssus hirundinis, Ornithonyssus sylviarum
Parus sp., Tit
Dermanyssus carpathicus, Dermanyssus gallinae, Ornithonyssus pipistrelli, Ornithonyssus sylviarum
Passer domesticus (Linnaeus, 1758), House Sparrow
Dermanyssus hirundinis, Dermanyssus gallinae
Passer montanus (Linnaeus, 1758), Eurasian Tree Sparrow
Dermanyssus carpathicus, Dermanyssus gallinae, Dermanyssus hirundinis, Ornithonyssus pipistrelli, Ornithonyssus
sylviarum, Pellonyssus reedi
Passer sp., Sparrow
Dermanyssus americanus, Dermanyssus hirundinis
Phoenicurus ochruros (S. G. Gmelin, 1774), Black Redstart
Dermanyssus carpathicus, Dermanyssus gallinae, Dermanyssus hirundinis, Ornithonyssus sylviarum
Phoenicurus phoenicurus (Linnaeus, 1758), Common Redstart
Dermanyssus carpathicus
Phoenicurus sp., Redstart
Dermanyssus carpathicus
Phylloscopus collybita (Vieillot, 1817), Common Chiffchaff

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Dermanyssus carpathicus, Ornithonyssus sylviarum
Phylloscopus sibilatrix (Bechstein, 1793), Wood Warbler
Ornithonyssus sylviarum
Pica pica (Linnaeus, 1758), Eurasian Magpie
Dermanyssus carpathicus, Dermanyssus hirundinis, Ornithonyssus sylviarum
Prunella modularis (Linnaeus, 1758), Dunnock
Dermanyssus carpathicus, Ornithonyssus sylviarum
Regulus sp., Kinglet
Dermanyssus sp.
Remiz pendulinus (Linnaeus, 1758), Eurasian Penduline Tit
Dermanyssus carpathicus, Dermanyssus gallinae, Ornithonyssus sylviarum
Riparia riparia (Linnaeus, 1758), Sand Martin
Dermanyssus carpathicus, Dermanyssus chelidonis, Dermanyssus gallinae, Dermanyssus hirundinis, Ornithonyssus
sylviarum
Saxicola rubetra (Linnaeus, 1758), Whinchat
Dermanyssus carpathicus
Serinus serinus (Linnaeus, 1766), European Serin
Dermanyssus carpathicus, Ornithonyssus sylviarum
Sitta europaea Linnaeus, 1758, Eurasian Nuthatch
Dermanyssus hirundinis, Ornithonyssus pipistrelli, Ornithonyssus sylviarum
Sturnus vulgaris Linnaeus, 1758, Common Starling
Dermanyssus gallinae, Dermanyssus hirundinis
Sylvia atricapilla (Linnaeus, 1758), Eurasian Blackcap
Dermanyssus carpathicus, Dermanyssus gallinae, Dermanyssus hirundinis, Ornithonyssus sylviarum
Sylvia nisoria (Bechstein, 1792), Barred Warbler
Ornithonyssus sylviarum
Sylvia sp., Warbler
Dermanyssus gallinae, Ornithonyssus sylviarum
Taeniopygia castanotis (Gould, 1837), Chestnut-eared Finch
Dermanyssus carpathicus, Ornithonyssus bacoti,
Turdus merula Linnaeus, 1758, Common Blackbird
Dermanyssus carpathicus, Dermanyssus gallinae, Dermanyssus hirundinis, Ornithonyssus sylviarum
Turdus philomelos C. L. Brehm, 1831, Song Thrush
Dermanyssus carpathicus, Dermanyssus gallinae, Dermanyssus hirundinis, Ornithonyssus sylviarum
Turdus pilaris Linnaeus, 1758, Fieldfare
Dermanyssus carpathicus, Ornithonyssus sylviarum
Turdus sp., Thrush
Dermanyssus carpathicus, Dermanyssus gallinae, Ornithonyssus pipistrelli
Turdus torquatus Linnaeus, 1758, Ring Ouzel
Dermanyssus carpathicus

Order: Piciformes Meyer & Wolf, 1810


Dendrocopos major (Linnaeus, 1758), Great Spotted Woodpecker
Dermanyssus carpathicus, Ornithonyssus sylviarum
Dendrocopos sp., Woodpecker
Dermanyssus gallinae

Order: Podicipediformes Fürbringer, 1888


Podiceps cristatus (Linnaeus, 1758), Great Crested Grebe
Ornithonyssus sylviarum
Podiceps grisegena (Boddaert, 1783), Red-necked Grebe
Dermanyssus carpathicus

Order: Strigiformes Wagler, 1830


Asio otus (Linnaeus, 1758), Long-eared Owl
Dermanyssus gallinae, Dermanyssus hirundinis
Bubo bubo (Linnaeus, 1758), Eurasian Eagle-Owl
Ornithonyssus sylviarum
Glaucidium passerinum (Linnaeus, 1758), Eurasian Pygmy Owl

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Ornithonyssus sylviarum
Strix aluco Linnaeus, 1758, Tawny Owl
Dermanyssus carpathicus

Class: Mammalia
Order: Carnivora
Canis lupus familiaris Linnaeus, 1758, Domestic Dog
Dermanysssus carpathicus, Ornithonyssus pipistrelli
Mustela putorius (Linnaeus, 1758), European Polecat
Dermanyssus gallinae

Order: Rodentia
Apodemus flavicollis (Melchior, 1834), Yellow-necked Mouse
Dermanyssus carpathicus, Ornithonyssus pipistrelli, Ornithonyssus sylviarum
Apodemus sp., Mouse
Dermanyssus gallinae, Dermanyssus hirundinis
Glis glis (Linnaeus, 1766), Edible Dormouse
Dermanyssus carpathicus, Dermanyssus hirundinis, Ornithonyssus pipistrelli, Ornithonyssus sylviarum
Micromys minutus (Pallas, 1771), Harvest Mouse
Ornithonyssus pipistrelli
Mus musculus Linnaeus, 1766, House Mouse
Dermanyssus gallinae, Dermanyssus hirundinis, Ornithonyssus bacoti, Ornithonyssus sylviarum
Muscardinus avellanarius (Linnaeus, 1758), Hazel Dormouse
Dermanyssus gallinae, Ornithonyssus pipistrelli
Rattus norvegicus (Berkenhout, 1769), Brown Rat
Dermanyssus gallinae

Order: Soricomorpha
Talpa europaea Linnaeus, 1758, European Mole
Dermanyssus carpathicus

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