Hard ticks (Acari: Ixodidae) associated with birds in Europe: Review of literature data

Abstract

Hard ticks (Acari: Ixodidae) are considered the most important transmitters of pathogens in the temperate zone that covers most of Europe. In the era of climate change tick-borne diseases are predicted to undergo geographical range expansion toward the north through regions that are connected to southern areas of the continent by bird migration. This alone would justify the importance of synthesized knowledge on the association of tick species with avian hosts, yet birds also represent the most taxonomically and ecologically diverse part of urban vertebrate fauna. Birds frequently occur in gardens and near animal keeping facilities, thus playing a significant role in the dispersal of ticks and tick-borne pathogens in synanthropic environments. The primary aim of this review is to provide a comprehensive reference source (baseline data) for future studies, particularly in the context of discovering new tick-host associations after comparison with already published data. The records on the ixodid tick infestations of birds were assessed from nearly 200 papers published since 1952. In this period, 37 hard tick species were reported from 16 orders of avian hosts in Europe. Here we compile a list of these tick species, followed by the English and Latin name of all reported infested bird species, as well as the tick developmental stage and country of origin whenever this information was available. These data allowed a first-hand analysis of general trends regarding how and at which developmental stage of ticks tend to infest avian hosts. Five tick species that were frequently reported from birds and show a broad geographical distribution in the Western Palearctic (Ixodes arboricola, I. frontalis, I. ricinus, Haemaphysalis concinna and Hyalomma marginatum) were also selected for statistical comparisons. Differences were demonstrated between these tick species regarding their association with bird species that typically feed from the ground and those that rarely occur at the soil level. The ecology of these five bird-infesting tick species is also illustrated here according to avian orders, taking into account the ecology (habitat type) and activity (circadian rhythm and feeding level) of most bird species that represent a certain order.

Background

Hard ticks (Acari: Ixodidae) are considered the most important vectors (transmitters) of pathogens under the temperate zone (1) including most of Europe. In the era of climate change (global warming), vector-borne diseases in general, and tick-borne disease in particular are predicted to undergo dramatic changes in geographical distribution (2, 3) mostly implying range expansion of tick species toward more northern latitudes. In such scenarios birds may play the most important role in tick dispersal over both short and long distances due to their migratory habits (4). Passeriform birds can carry significant numbers of ticks [in a large scale study this was up to 20 ticks per bird: (5)]. These ticks are mostly larvae and nymphs of generalist (non-ornithophilic) tick species. These ticks may remain on their avian host for a period raging from a few days (in case of larvae of three-host tick species exemplified by Ixodes ricinus) to even 5 weeks or longer [in case of two-host ticks species, e.g., Hyalomma rufipes: (6)]. While most studies focus on tick species carried by birds during their spring migration from the southern wintering grounds toward the north in Europe, tick “transportation” by birds may equally be important in western, eastern, and north-to-southern directions (5).

If thermophilic ticks carried by birds from south to north detach after engorgement, and find themselves in a suitable (warm enough) environment, they can molt to the next developmental stage. These ticks may then survive through mild winters, potentially establishing new, northern populations (7). This may contribute to the increased risks and evident changes of tick-borne pathogen transmission due to climate warming and tick dispersal by migratory birds, even in northern Europe (8).

On the other hand, several bird-specialist (ornithophilic) Ixodes species occur in the Western Palearctic, most of which associate with sea birds in Western Europe (e.g., I. uriae, I. caledonicus, I. rothschildi). These ticks occur in and near avian nests, or are frequently endophilic and live in tree holes or burrows inhabited by nesting birds, particularly in the subgenus Pholeoixodes (e.g., I. arboricola, I. lividus) (9). Since the avian hosts of tick species are expected to be less affected by climate change in terms of their geographical distribution, ornithophilic ticks will likely undergo less significant changes in their geographical occurrence than generalist tick species carried by birds. In addition, ornithophilic ticks seldom infest humans or domestic mammals [e.g., I. frontalis: (10)] and thus may have a less important epidemiological role from a veterinary-medical point of view.

This review focuses on hard ticks reported from birds in Northern, Western, Central, and Southern Europe, in the hope that the checklist will provide useful baseline information for future scientific research in the topics of ticks and tick-borne pathogens. Nearly 700 scientific papers published since 1952 were checked for this review. Soft ticks (Acari: Argasidae) were not included in this review, because in the majority of cases these are reported from the environment and thus data on their association with avian host species are not representative.

Methods

The primary corpus of publications used in this review was collated with database search using the following keywords: “ticks” OR “Ixodidae” OR “Ixodes” OR “Hyalomma” OR “Haemaphysalis” AND “birds” AND “Europe,” followed by a search with names of bird-specialist tick species (I. arboricola, I. caledonicus, I. eldaricus, I. festai, I. frontalis, I. lividus, I. rothschildi, I. unicavatus and I. uriae). The following databases were used: Web of Science, Zoological Record, and Google Scholar. These records were imported into an Excel file, followed by screening the publications and references cited within. After exclusion of duplicates, we extracted each individual bird host-tick record from these references, noting the location (country), host and parasite species and developmental stage of ticks.

Within Prostriata and Metastriata, tick species names are arranged alphabetically and are used sensu Guglielmone et al. (10). Ixodes redikorzevi is considered as a synonym of I. acuminatus. Data were only included in the checklist if the tick collected from a bird was reported as identified to the species level. E.g., in Norway a rare finding was mentioned, the occurrence of a Dermacentor sp. on a willow warbler (Phylloscopus trochilus) (11), but this was excluded from the main text.

Bird species as hosts of a certain tick species are listed alphabetically according to their international English name (https://bou.org.uk/british-list/bird-names/). When higher avian taxa are mentioned, these follow the phylogenetic order based on Kuhl et al. (12), starting with Passeriformes. Genus names are not abbreviated in the headings, owing to the high number and mixed usage of host and tick Latin names, also taking into account that scientists from a broad range of biology-related fields may use this checklist. Whenever a bird species was mentioned by its binominal name in an article, this was used for identification, even if the English name was also written in the text. English and Latin host species names are followed by tick developmental stages if this information was available (L: larva, N: nymph, M: male, F: female, A: adult [where there was no information about the sex of the adult tick)]. The abbreviation “NA” is used to indicate that no data were available about the sex and developmental stage of ticks collected from birds. If a tick species was reported from bird(s) in a country without mentioning avian host species, an exclamation mark (!) follows the country name in the list. If tick species infesting a certain bird species are mentioned in a reference but only some of the data inform about the tick developmental stage, only these were incorporated into the text. It is also noteworthy that in some reports blood meal analysis allowed the identification of previous tick hosts. In addition, data on ticks reported from bird nests are included and marked in the text as “in nest”: these ticks probably also originate from or can associate with birds.

The geographical area covered by this review is in the Western Palearctic, excluding North Africa, the Middle East, Belarus and Russia but including Ukraine. Cyprus however, is also considered. Not just because it is partially European territory, but due to the fact that the island has a high epidemiological significance concerning the aim of this manuscript. Regarding geographical names, old references often refer to Czechoslovakia, which no longer exists. In cases when it was unambiguous whether the samples came from the Czech Republic or Slovakia, the country is mentioned as such. To maintain the user friendliness of this checklist the broader Palearctic distribution, general ecology, and vector role of tick species are not mentioned. This is because this information is available in other sources [e.g., (9)].

Review of literature data

Prostriata

Ixodes acuminatus

Overview: Ixodes acuminatus is distributed in temperate and Mediterranean Europe (8). Accordingly, this tick species is occasionally found on birds in South-European countries. Based on literature data it is mainly a parasite of passeriform birds (from this order 15 species have been shown as hosts) but has also been found twice on galliform birds.

Passeriformes: 15; Galliformes: 2

Hosts:

• - Bearded reedling – Panurus biarmicus [N: (13)]

• - Common blackbird – Turdus merula [F: (14–16); NA: (17, 18)]

• - Common chaffinch – Fringilla coelebs [L: (14); N: (14)]

• - Common pheasant – Phasianus colchicus [A: (19)]

• - Common redstart – Phoenicurus phoenicurus [L: (14)]

• - Eurasian blue tit- Cyanistes caeruleus [N: (14)]

• - Eurasian magpie – Pica pica [L: (14); N: (14)]

• - European goldfinch – Carduelis carduelis [F: (14)]

• - European robin – Erithacus rubecula [N: (13)]

• - Fieldfare -Turdus pilaris [NA: (14)]

• - Garden warbler – Sylvia borin [N: (20)]

• - Great tit – Parus major [L: (14); N: (13, 14)]

• - House sparrow – Passer domesticus [NA: (14)]

• - Red-legged partridge – Alectoris rufa [A: (19)]

• - Redwing – Turdus iliacus [N: (21)]

• - Song thrush – Turdus philomelos [F: (16)]

• - Winter wren – Troglodytes troglodytes [L: (21); N: (21)]

Distribution of reported cases: Greece (20), Italy (19, 22!), Romania (13, 14, 23!), Cyprus (16), Portugal (15, 17, 21), France (18).

Ixodes arboricola

Overview: Ixodes arboricola is widespread throughout Europe. As its English name suggests, the Tree-hole tick is primarily a parasite of hole-nesting birds (8). Literature data usually support this theory, as I. arboricola was mostly found on passeriform (35 species), strigiform (5 species), piciform (1 species), and columbiform (1 species) birds. The fact that other, non-hole-nesting predators (2 falconiform and 1 accipitriform species) have been described as hosts, does not contradict the former statement, as these species can become infected with ticks by getting in contact with their prey.

Passeriformes: 34; Strigiformes: 5; Falconiformes: 2; Columbiformes: 1; Piciformes: 1; Accipitriformes: 1

Hosts:

• - Barn owl – Tyto alba [L: (24); NA: (25, 26)]

• - Barn swallow – Hirundo rustica [NA: (26)]

• - Boreal owl – Aegolius funereus [N: (27 in nest)]

• - Coal tit – Periparus ater [N: (28); NA: (26, 29, 30)]

• - Collared flycatcher – Ficedula albicollis [N: (31, 32); M: (31, 32 in nest); F: (32^a); NA: (33)]

• - Common blackbird – Turdus merula [L: (15); N: (13, 14, 34); NA: (30, 35)]

• - Common kestrel- Falco tinnunculus [F: (34)]

• - Common redstart – Phoenicurus phoenicurus [L: (14, 31); N: (14); NA: (33)]

• - Common starling – Sturnus vulgaris [L: (31, 34, 36, 37); N: (31, 34, 36, 37); M: (37); F: (31, 34, 36, 37); NA: (14, 26, 30, 33, 38^a in nest)]

• - Common wood pigeon – Columba palumbus [L: (34)]

• - Eurasian blue tit- Cyanistes caeruleus [L: (14, 28, 39–42); N: (13, 15, 28, 31, 40–45); M: (32 in nest); F: (31, 32, 40, 42, 43); NA: (17, 25, 29, 33, 38, 46)]

• - Eurasian bullfinch – Pyrrhula pyrrhula [NA: (30)]

• - Eurasian jay – Garrulus glandarius [A: (47)]

• - Eurasian nuthatch – Sitta europaea [L: (21, 46); N: (21, 28, 45–47); A: (46); M: (32 in nest); NA: (26, 29, 33, 48)]

• - Eurasian penduline tit – Remiz pendulinus [L: (49 in nest)]

• - Eurasian pygmy owl – Glaucidium passerinum [L: (31)]

• - Eurasian reed warbler – Acrocephalus scirpaceus [NA: (30)]

• - Eurasian siskin – Carduelis spinus [N: (41)]

• - Eurasian tree sparrow – Passer montanus [NA: (26, 30, 38)]

• - Eurasian treecreeper – Certhia familiaris [L: (33), NA: (29)]

• - European greenfinch – Carduelis chloris [L: (42)]

• - European pied flycatcher – Ficedula hypoleuca [N: (31, 45); NA: (30, 38)]

• - European robin – Erithacus rubecula [L: (13, 14); N: (11, 13, 14, 28, 31); NA: (30)]

• - European serin – Serinus serinus [N: (21); F: (21)]

• - Great spotted woodpecker – Dendrocopos major [NA: (47)]

• - Great tit – Parus major [L: (14, 28, 31, 34, 36, 40–42, 44, 50, 51); N: (13– 15, 17, 21, 24, 28, 31, 32, 34, 36, 40–43, 45, 50, 51); M: (32^a in nest); F: (24, 32^a, 40, 41, 43, 52, 53); NA: (17, 25, 29, 33, 38, 46)]

• - House sparrow – Passer domesticus [NA: (26, 30)]

• - Little owl – Athene noctua [N: (34); F: (34); NA: (30)]

• - Long-tailed tit – Aegithalos caudatus [L: (21)]

• - Marsh tit – Poecile palustris [L: (28, 42); N: (42); NA: (29, 30)]

• - Northern goshawk – Accipiter gentilis [L: (34)]

• - Peregrine falcon – Falco peregrinus [L: (54); N: (54); F: (54)]

• - Rook – Corvus frugilegus [L: (55)]

• - Sand martin – Riparia riparia [N: (24 in nest); F: (24 in nest); NA: (30)]

• - Short-toed treecreeper – Certhia brachydactyla [L: (15); NA: (17)]

• - Song thrush – Turdus philomelos [L: (13); N: (11, 13)]

• - Spotless starling – Sturnus unicolor [L: (17); NA: (17)]

• - Spotted flycatcher – Muscicapa striata [N: (14); F: (31)]

• - Tawny owl -Strix aluco [L: (31); NA: (26)]

• - Western jackdaw – Corvus monedula [L: (55); N: (31); F: (31); NA: (30, 48)]

• - Willow tit – Poecile montana [N: (41); NA: (30)]

• - Willow warbler – Phylloscopus trochilus [NA: (30)]

• - Winter wren – Troglodytes troglodytes [N: (41); NA: (30)]

• - Yellowhammer – Emberiza citrinella [NA: (25)]

Distribution of reported cases: Sweden (31, 36, 56!), United Kingdom (24, 25, 30, 46, 53), Czech Republic (28, 29, 32, 57!), Slovakia (26, 32^a, 33, 49), Netherlands (43), Denmark (37), Norway (11), Belgium (40, 45, 48, 50), Romania (13, 14, 23!, 55), Ukraine (52), Portugal (15, 17, 21), Poland (26, 41, 51), Spain (42, 44), Switzerland (34), Germany (38, 54), Hungary (47), Croatia (35), Belarus (38^a).

Ixodes berlesei

Overview: According to literature data, I. berlesei was reported from birds in France. Unfortunately, the source article (18) is not accurate about the host species, but we do know that they belong to the Columbiformes order.

Columbiformes:1

Distribution of reported cases: France (18!).

Ixodes caledonicus

Overview: Ixodes caledonicus is an ornithophilic tick species (10). It is primarily a parasite of the northern European bird fauna. Despite the limited data available, the host range appears to be broad: it has been reported from 6 passeriform, 3 falconiform, 1 caprimulgiform, and one procellariiform bird species.

Passeriformes: 6; Falconiformes: 3; Caprimulgiformes: 1; Procellariiformes: 1; Columbiformes: 1

Hosts:

• - Common kestrel- Falco tinnunculus [NA: (30)]

• - Common redstart – Phoenicurus phoenicurus [N: (24)]

• - Common starling – Sturnus vulgaris [NA: (30)]

• - Common swift – Apus apus [NA: (26)]

• - Gyrfalcon – Falco rusticolus [L: (58); N: (58); A: (58); NA: (30)]

• - Hooded crow – Corvus cornix [NA: (30)]

• - Northern fulmar – Fulmarus glacialis [F: (59); NA: (30)]

• - Northern raven – Corvus corax [NA: (30)]

• - Peregrine falcon – Falco peregrinus [N: (31)]

• - Red crossbill – Loxia curvirostra [NA: (30)]

• - Rock dove – Columba livia [NA: (59^a)]

• - Western jackdaw – Corvus monedula [NA: (30)]

Distribution of reported cases: United Kingdom (24, 30), Faroe Islands (59), Sweden (31), Poland (26), Norway (30!), Germany (30!), Iceland (58), NA (59^a).

Ixodes canisuga

Overview: Ixodes canisuga is primarily a parasite of mammals (8) and it is relatively rare on birds. So far, it has been found on 5 passeriform and on 2 strigiform hosts.

Passeriformes: 5; Strigiformes: 2

Hosts:

• - Common starling – Sturnus vulgaris [NA: (30)]

• - Eurasian blue tit- Cyanistes caeruleus [NA: (30)]

• - Eurasian eagle-owl – Bubo bubo [N: (60)]

• - Eurasian tree sparrow – Passer montanus [NA: (30)]

• - Great tit – Parus major [NA: (30)]

• - Little owl – Athene noctua [L: (24); N: (24); F: (24)]

• - Sand martin – Riparia riparia [M: (24 in nest); F: (24 in nest)]

Distribution of reported cases: United Kingdom (24, 30), Germany (61!), Portugal (60).

Ixodes eldaricus

Overview: Ixodes eldaricus is a rare, poorly known tick species. The only available data about this parasite feeding on European birds are from Poland and Cyprus, where this tick species was reported from four passeriform bird species in total

Passeriformes: 4

Hosts:

• - Common blackbird – Turdus merula [N: (16)]

• - Dunnock – Prunella modularis [F: (62)]

• - European robin – Erithacus rubecula [M: (62); F: (62)]

• - Tree pipit – Anthus trivialis [N: (16)]

Distribution of reported cases: Poland (62), Cyprus (16).

Ixodes festai

Overview: Ixodes festai is a poorly known species that was reported from 5 passeriform birds, mostly in Central European countries.

Passeriformes: 5

Hosts:

• - Common blackbird – Turdus merula [N: (34); F: (34, 51, 63); NA: (64)]

• - Common chaffinch – Fringilla coelebs [F: (34)]

• - Dunnock – Prunella modularis [N: (34); F: (65)]

• - European greenfinch – Carduelis chloris [F: (65)]

• - Song thrush – Turdus philomelos [M: (64); F: (63, 64)]

Distribution of reported cases: Italy (63, 64), Hungary (65), Switzerland (34), Poland (51).

Ixodes frontalis

Overview: Ixodes frontalis has a pan-European distribution. However, it appears to be more frequent in warmer regions. It is an ornithophilic tick species that rarely feeds on other, exceptional hosts (10). As such, it is commonly found on birds, mostly on Passeriformes, but the host range appears to be broad.

Passeriformes: 56; Charadriiformes: 1; Accipitriformes: 4; Galliformes: 4; Falconiformes: 1; Coraciiformes: 1; Gruiformes: 3; Columbiformes: 2; Strigiformes: 2

Hosts:

• - Barn owl – Tyto alba [F: (25)]

• - Black redstart – Phoenicurus ochruros [L: (66); NA: (35)]

• - Black-headed gull – Chroicocephalus ridibundus [F: (67)]

• - Bohemian waxwing – Bombycilla garrulus [NA: (35)]

• - Booted eagle – Hieraaetus pennatus [A: (60)]

• - Carrion crow – Corvus corone [NA: (68)]

• - Cetti's warbler – Cettia cetti [L: (21)]

• - Chicken – Gallus gallus domesticus [NA: (46)]

• - Coal tit – Periparus ater [NA: (30, 68)]

• - Common blackbird – Turdus merula [L: (15, 17, 20, 21, 34, 39, 47, 66, 67, 69–74); N: (15–17, 21, 24, 39, 43, 47, 60, 66–76); M: (71, 75); F: (21, 25, 44, 66, 67, 69–74, 77); A: (78); NA: (48, 68)]

• - Common buzzard – Buteo buteo [N: (60); A: (60)]

• - Common chaffinch – Fringilla coelebs [L: (21, 24, 66, 69); N: (15, 21, 69); F: (20, 21, 66, 70, 79), (79); NA: (30, 68)]

• - Common chiffchaff – Phylloscopus collybita [L: (16, 21, 66, 71); N: (21, 80)]

• - Common firecrest – Regulus ignicapilla [N: (15, 21)]

• - Common kestrel – Falco tinnunculus [F: (74); NA: (46, 68)]

• - Common kingfisher – Alcedo atthis [F: (67)]

• - Common linnet – Carduelis cannabina [NA: (30)]

• - Common moorhen – Gallinula chloropus [F: (81)]

• - Common nightingale – Luscinia megarhynchos [N: (71, 82); F: (21)]

• - Common pheasant – Phasianus colchicus [NA: (68)]

• - Common redstart – Phoenicurus phoenicurus [N: (67, 80)]

• - Common reed bunting – Emberiza schoeniclus [L: (66)]

• - Common starling – Sturnus vulgaris [NA: (48, 68)]

• - Common whitethroat – Sylvia communis [N: (25, 53, 71, 75, 76, 80); F: (25, 53)]

• - Common wood pigeon – Columba palumbus [F: (25); NA: (68)]

• - Corn crake – Crex crex [NA: (68)]

• - Dunnock – Prunella modularis [L: (66); F: (25, 53, 66)]

• - Eurasian blackcap – Sylvia atricapilla [L: (16, 21, 66, 69, 71); N: (11, 21, 60, 65, 71, 80); F: (25, 53, 71); A: (47); NA: (48)]

• - Eurasian blue tit – Cyanistes caeruleus [L: (15, 21, 25, 40, 53, 71); N: (40); F: (21, 40, 70, 71, 77); NA: (68)]

• - Eurasian bullfinch – Pyrrhula pyrrhula [F: (25, 53)]

• - Eurasian collared dove – Streptopelia decaocto [F: (25, 67, 81, 83)]

• - Eurasian jay – Garrulus glandarius [L: (47); F: (21); A: (47); NA: (30, 68)]

• - Eurasian magpie – Pica pica [NA: (68)]

• - Eurasian nuthatch – Sitta europaea [NA: (68)]

• - Eurasian reed warbler – Acrocephalus scirpaceus [L: (21, 67); N: (21, 70, 80); F: (21)]

• - Eurasian sparrowhawk – Accipiter nisus [NA: (68)]

• - Eurasian stonechat – Saxicola torquatus [NA: (68)]

• - Eurasian tree sparrow – Passer montanus [F: (25); NA: (68)]

• - Eurasian treecreeper – Certhia familiaris [NA: (30)]

• - European goldfinch – Carduelis carduelis[(F: (74)]

• - European greenfinch – Carduelis chloris [L: (15, 21, 67); F: (16, 21, 65, 67); NA: (68)]

• - European pied flycatcher – Ficedula hypoleuca [NA: (30)]

• - European robin – Erithacus rubecula [L: (15–17, 20, 21, 34, 39, 65– 67, 69, 71–73); N: (15, 17, 21, 34, 39, 41, 65, 66, 71–73, 84); M: (21); F: (16, 21, 25, 65, 73, 82)]

• - Fieldfare -Turdus pilaris [F: (74); NA: (68)]

• - Goldcrest – Regulus regulus [N: (41); NA: (30)]

• - Great gray shrike – Lanius excubitor [NA: (68)]

• - Great tit – Parus major [L: (15, 17, 21, 34, 40, 65, 66, 71); N: (15, 17, 21, 40); F: (21, 25, 40, 65); NA: (68)]

• - Gray partridge – Perdix perdix [NA: (68)]

• - Gray wagtail – Motacilla cinerea [N: (67)]

• - Harris's hawk – Parabuteo unicinctus [F: (74)]

• - House sparrow – Passer domesticus [L: (24); N: (25); F: (21, 24, 25, 67); NA: (68)]

• - Lesser redpoll – Carduelis cabaret [F: (25, 70)]

• - Long-eared owl – Asio otus [L: (67); N: (85); F: (85); NA: (68)]

• - Long-tailed tit – Aegithalos caudatus [N: (21, 71); F: (25, 53, 71, 72)]

• - Marsh tit – Poecile palustris [NA: (30)]

• - Melodious warbler – Hippolais polyglotta [N: (21)]

• - Mistle trush – Turdus viscivorus [NA: (48, 68)]

• - Red-legged partridge – Alectoris rufa [M: (67); A: (86); NA: (68, 87)]

• - Redwing – Turdus iliacus [L: (21, 70); N: (21, 67, 70, 73); A: (60); NA: (30, 68)]

• - Ring ouzel – Turdus torquatus [F: (16); NA: (30)]

• - Rook – Corvus frugilegus [NA: (30)]

• - Sardinian warbler – Sylvia melanocephala [L: (15); N: (71); F: (67)]

• - Scarlet-headed blackbird – Amblyramphus holosericeus [F: (25)]

• - Sedge warbler – Acrocephalus schoenobaenus [F: (21)]

• - Short-toed treecreeper – Certhia brachydactyla [L: (15, 21); N: (15, 21); F: (66)]

• - Song thrush – Turdus philomelos [L: (15, 16, 21, 41, 65–67, 70–72, 80); N: (15, 16, 21, 34, 39, 41, 65, 66, 70–73, 80); F: (16, 21, 25, 44, 53, 70, 71, 77); NA: (68)]

• - Subalpine warbler – Sylvia cantillans [N: (84)]

• - Tree pipit – Anthus trivialis [NA: (30)]

• - Water rail – Rallus aquaticus [F: (67); NA: (30)]

• - Western yellow wagtail – Motacilla flava [L: (21); N: (16)]

• - Whinchat – Saxicola rubetra [N: (84)]

• - Willow tit – Poecile montana [NA: (30, 48)]

• - Willow warbler – Phylloscopus trochilus [L: (16, 24, 73, 80); N: (80); F: (80)]

• - Winter wren – Troglodytes troglodytes [L: (15, 17, 21, 66, 67, 73); N: (15, 17, 21, 31, 66); F: (25, 53); NA: (68)]

Distribution of reported cases: United Kingdom (24, 25, 30, 46, 53, 75–77, 80), France (67, 68, 81, 83), Spain (44, 66, 71, 72, 86, 87), Poland (39, 41), Greece (20), Germany (74), Netherlands (43, 70), Norway (11), Belgium (40, 48), Hungary (47, 65), Sweden (56!, 73), Portugal (Azores) (69), Italy (22!, 78, 84), Switzerland (79, 82), Moldova (88!), Portugal (15, 17, 21, 60, 85), Croatia (35), Cyprus (16).

Ixodes gibbosus

Overview: Ixodes gibbosus is a Mediterranean tick species, rarely reported from birds (8).

Passeriformes: 1

Hosts:

• - Common blackbird – Turdus merula (L/N: (89)]

Distribution of reported cases: Greece (89).

Ixodes hexagonus

Overview: Ixodes hexagonus is a common parasite of European foxes and hedgehogs (8). In the United Kingdom, this tick was reported from 5 passeriform, 1 galliform, 1 falconiform, 1 columbiform and, 1 strigiform birds. Whereas, in Spain it was reported from 1 galliform bird. This tick species was also found on a bird in Germany but no information is available on the host species.

Passeriformes: 3; Galliformes: 2; Falconiformes: 1; Columbiformes: 1

Hosts:

• - Chicken – Gallus domesticus [F: (25)]

• - Common kestrel- Falco tinnunculus [NA: (30)]

• - Common starling – Sturnus vulgaris [NA: (30)]

• - Common wood pigeon – Columba palumbus [F: (46)]

• - Eurasian blue tit- Cyanistes caeruleus [NA: (30)]

• - Red-legged partridge – Alectoris rufa [NA: (90)]

• - Winter wren – Troglodytes troglodytes [NA: (30)]

Distribution of reported cases: United Kingdom (25, 30, 46), Germany (61!), Spain (90).

Ixodes lividus

Overview: Ixodes lividus has a pan-European distribution, and it is the host-specific parasite of the sand martin (Riparia riparia). In Poland, it has been found on a Barn swallow (Hirundo rustica) as well.

Passeriformes: 2

Hosts:

• - Barn swallow – Hirundo rustica [dNA: (26)]

• - Sand martin – Riparia riparia [L: (31, 34, 53, 91 in nest, 92 in nest, 93–95 in nest, 96); N: (31, 34, 52, 53, 81, 91 in nest, 92 in nest, 93–95 in nest, 96); M: (31, 34, 91 in nest, 92 in nest, 95 in nest, 96 in nest); F: (31, 34, 48, 53, 91 in nest, 92 in nest, 95 in nest, 96 in nest, 97); A: (94, 96); NA: (25, 98–102)]

Distribution of reported cases: United Kingdom (25, 53, 97, 98), France (81), Portugal (99), Finland (96), Sweden (31, 103!) Germany (61!, 94), Lithuania (91, 92), Moldova (93), Ukraine (52), Czech Republic (95), Schwitzerland (34), Hungary (100, 101), Poland (26, 102), Belgium (48).

Ixodes persulcatus

Overview: Ixodes persulcatus is widely distributed in the Northern European region (Russia, Scandinavia, and the Baltic region) (8). According to literature data however, it should be considered as a rare parasite of birds in Europe. It was reported only from 2 passeriform bird species.

Passeriformes: 2

Hosts:

• - Sedge warbler – Acrocephalus schoenobaenus [N: (104)]

• - Willow warbler – Phylloscopus trochilus [N: (31)]

Distribution of reported cases: Estonia (104), Sweden (31).

Ixodes ricinus

Overview: Ixodes ricinus has a pan-European distribution, including countries of Southern, Western, Central, Eastern and Northern Europe. As outlined below, this tick species was reported from 99 passeriform bird species, and from species of further 12 avian orders. According to available data, the immature stages of this tick species appear to be the most frequent hard ticks feeding on European birds.

Passeriformes: 99; Galliformes: 8; Accipitriformes: 6; Ciconiiformes: 1; Anseriformes: 3; Gruiformes: 3; Cuculiformes: 1; Charadriiformes: 7; Falconiformes: 1; Columbiformes: 2; Strigiformes: 3; Piciformes: 4; Bucerotiformes: 1

Hosts:

• - Barn swallow – Hirundo rustica [N: (31)]

• - Barred warbler – Sylvia nisoria [L: (105); N: (36, 105, 106); NA: (47)]

• - Bearded reedling – Panurus biarmicus [N: (13)]

• - Black grouse – Tetrao tetrix [L: (107); N: (31, 107)]

• - Black kite – Milvus migrans [NA: (47)]

• - Black redstart – Phoenicurus ochruros [N: (14, 41, 73, 79)]

• - Black stork – Ciconia nigra [N: (108)]

• - Bluethroat – Luscinia svecica [L: (31, 34, 43, 109); N: (31, 70, 73, 105, 106, 110–113)]

• - Blyth's reed warbler – Acrocephalus dumetorum [N: (104)]

• - Bohemian waxwing – Bombycilla garrulus [NA: (14)

• - Brambling – Fringilla montifringilla [L: (34, 73, 79, 114); N: (34, 36, 79, 114); NA: (115)]

• - Canada goose – Branta canadensis [L: (107); N: (107)]

• - Carrion crow – Corvus corone [N: (34); NA: (30)]

• - Cetti's warbler – Cettia cetti [L: (15); N: (15); NA: (35)]

• - Chicken – Gallus gallus domesticus [M: (34)]

• - Coal tit – Periparus ater [L: (34, 116, 117); N: (117, 118); NA: (26, 29, 57, 115)]

• - Collared flycatcher – Ficedula albicollis [N: (119); NA: (33)]

• - Common blackbird – Turdus merula [L: (11, 13–15, 17, 21, 28, 31, 34, 36, 39, 41, 43, 44, 51, 57, 65, 66, 70, 73–76, 78, 79, 93, 105, 106, 109, 112–140); N: (11, 13–15, 17, 21, 24, 25, 28, 31, 34, 36, 39, 41, 43, 44, 51, 52, 57, 65, 66, 70–79, 93, 104–106, 108, 109, 111–142); M: (31); F: (13–15, 21, 52, 73, 76); A: (119); NA: (29, 33, 35, 46, 48, 143–145)]

• - Common buzzard – Buteo buteo [N: (34); NA: (26, 33, 35, 46, 47)]

• - Common chaffinch – Fringilla coelebs [L: (11, 14, 15, 17, 25, 34, 36, 39, 41, 43, 66, 70, 73, 76, 79, 93, 105, 109, 112–121, 124, 129, 133, 135– 140); N: (11, 14, 15, 34, 39, 41, 43, 52, 70, 71, 73, 78, 79, 93, 105, 109, 112^a, 113, 114, 116, 117, 120, 121, 126, 129, 133, 135–140, 146); F: (17, 21, 79); NA: (33, 46–48, 57, 115, 123, 130, 144, 145, 147)]

• - Common chiffchaff – Phylloscopus collybita [L: (34, 39, 41, 65, 71, 112, 113, 129, 139, 142); N: (13, 21, 28, 31, 39, 41, 65, 70, 73, 76, 80, 104, 109, 120, 128, 129, 133, 138, 140, 141); NA: (57, 143); NA: (59)]

• - Common coot – Fulica atra [NA: (26)]

• - Common crane – Grus grus [NA: (30)]

• - Common cuckoo – Cuculus canorus [NA: (26)]

• - Common firecrest – Regulus ignicapilla [N: (39); NA: (123)]

• - Common grasshopper warbler – Locustella naevia [L: (122); N: (39, 104, 122); NA: (30)]

• - Common house martin – Delichon urbicum [F: (34); NA: (30)]

• - Common kestrel- Falco tinnunculus [L: (34); N: (34); NA: (14)]

• - Common linnet – Carduelis cannabina [L: (76); N: (43, 70, 105, 106, 109, 112, 113); NA: (47)]

• - Common nightingale – Luscinia megarhynchos [L: (34, 43, 65, 78, 128, 140, 148); N: (28, 34, 65, 71, 78, 128, 140, 148, 149); A: (140); NA: (14, 26, 33, 35)]

• - Common pheasant – Phasianus colchicus [L: (24, 31, 150); N: (24, 31, 34, 43, 150); M: (34); NA: (14, 33, 46, 47)]

• - Common quail – Coturnix coturnix [N: (75)]

• - Common redpoll – Carduelis flammea [L: (105); N: (73); NA: (30)]

• - Common redstart – Phoenicurus phoenicurus [L: (11, 31, 36, 39, 41, 43, 70, 76, 79, 105, 106, 119, 124, 130, 132, 133, 139); N: (11, 31, 36, 41, 43, 70, 73, 76, 79, 104–106, 110–113, 119, 124, 130, 132, 133, 141); NA: (33, 144)]

• - Common reed bunting – Emberiza schoeniclus [L: (13, 65, 105); N: (28, 43, 70, 73, 104)]

• - Common rosefinch – Carpodacus erythrinus [L: (28); N: (28, 104, 105, 109, 112, 113); NA: (26)]

• - Common starling – Sturnus vulgaris [L: (14, 34, 36, 78, 105, 106, 109, 112, 113, 121, 133, 135, 137, 139); N: (14, 34, 36, 73, 78, 105, 106, 112, 113, 121, 133, 135, 137, 139, 146); A: (119); NA: (30, 33, 47, 48, 147)]

• - Common whitethroat – Sylvia communis [L: (24, 31, 36, 43, 65, 70, 73, 75, 76, 105, 106, 109, 112, 113, 119, 122, 128, 132, 138); N: (28, 31, 34, 36, 41, 43, 53, 65, 70, 73, 75, 76, 79, 80, 104–106, 109, 110, 112^a, 113, 122, 124, 128, 132); F: (31); NA: (33, 144)]

• - Corn bunting – Emberiza calandra [NA: (33)]

• - Corn crake – Crex crex [L: (36); N: (14, 36); NA: (47)]

• - Dunnock – Prunella modularis [L: (11, 28, 39, 41, 43, 65, 70, 73, 76, 79, 105, 106, 109, 112, 113, 117, 120, 122, 128, 129, 133, 138, 142); N: (11, 28, 31, 34, 36, 39, 41, 43, 65, 70, 73, 76, 79, 82, 105, 106, 109, 111–113, 117, 119, 120, 122, 128, 129, 132, 133, 138, 142); NA: (57, 115, 145)]

• - Eurasian blackcap – Sylvia atricapilla [L: (15, 17, 31, 34, 36, 39, 41, 44, 65, 70, 71, 73, 78, 93, 105, 112, 113, 116, 117, 119–122, 126, 128, 129, 131–133, 136, 138); N: (15, 28, 31, 34, 39, 41, 44, 65, 70, 73, 75, 80, 93, 105, 109, 111, 112^a, 113, 116, 118, 119, 122, 126, 128, 129, 131–133, 138, 142); A: (118); NA: (33, 57, 115, 123, 143)]

• - Eurasian blue tit- Cyanistes caeruleus [L: (13, 28, 34, 40, 105, 106, 116, 117, 120, 121, 129, 135, 137); N: (28, 34, 39, 40, 43, 73, 104–106, 117, 120, 126, 133); NA: (29, 33, 115, 123, 143, 145)]

• - Eurasian bullfinch – Pyrrhula pyrrhula [L: (34, 39, 105, 117); N: (34, 36, 39, 105, 119, 131, 132, 138); NA: (115, 143)]

• - Eurasian collared dove – Streptopelia decaocto [NA: (33)]

• - Eurasian curlew – Numenius arquata [L: (107, 151); N: (107)]

• - Eurasian eagle-owl – Bubo bubo (L: (152); N: (152, 153)]

• - Eurasian golden oriole – Oriolus oriolus [L: (33); N: (33)]

• - Eurasian hoopoe – Upupa epops [L: (93); N: (93, 108)]

• - Eurasian jay – Garrulus glandarius [L: (15, 34, 44, 93, 116, 117, 129, 133); N: (14, 15, 33, 34, 43, 44, 70, 93, 116, 118, 129, 133); A: (118); NA: (26, 47, 115)]

• - Eurasian magpie – Pica pica [L: (14); N: (14, 34); F: (14, 34); NA: (26, 30, 33, 35, 47)]

• - Eurasian nuthatch – Sitta europaea [L:(15, 28, 34, 39, 116, 121, 129, 137); N: (34, 116, 121, 129, 133, 135); F: (52); NA: (33, 47, 48, 115, 123)]

• - Eurasian oystercatcher – Haematopus ostralegus [L: (31); N: (31)]

• - Eurasian reed warbler – Acrocephalus scirpaceus [L: (21, 28, 31, 39, 65, 112^a, (128)]; N: (21, 25, 28, 31, 34, 43, 53, 65, 70, 73, 78, 104, 105, 109, 112^a, 119, 128, 132, 142)]

• - Eurasian siskin – Carduelis spinus [L: (24, 34, 39, 117); N: (34, 39, 70, 105, 117)]

• - Eurasian sparrowhawk – Accipiter nisus [L: (24); N: (24, 31, 73, 105, 110, 140); A: (140)]

• - Eurasian stonechat – Saxicola torquatus [NA: (30)]

• - Eurasian tree sparrow – Passer montanus [L: (34, 70, 120); N: (34, 43, 70, 132); NA: (26, 33, 47)]

• - Eurasian treecreeper – Certhia familiaris [L: (73, 117, 121, 122); N: (79, 104, 105, 122, 130)]

• - Eurasian woodcock – Scolopax rusticola [N: (39); NA: (30, 33)]

• - Eurasian wryneck – Jynx torquilla [NA: (33)]

• - European crested tit – Lophophanes cristatus [L: (41); N: (41, 130)]

• - European golden plover – Pluvialis apricaria [NA: (154)]

• - European goldfinch – Carduelis carduelis [N: (109); NA: (33, 57)]

• - European green woodpecker – Picus viridis [N: (133)]

• - European greenfinch – Carduelis chloris [L: (15, 17, 105, 106, 112, 113, 117); N: (21, 33, 36, 39, 105, 106, 112, 113, 117, 120, 123, 142); NA: (46, 48, 57)]

• - European herring gull – Larus argentatus [N: (43)]

• - European honey buzzard – Pernis apivorus [NA: (155)]

• - European pied flycatcher – Ficedula hypoleuca [L: (13, 21, 34, 36, 41); N: (34, 36, 70, 119); NA: (30, 48)]

• - European robin – Erithacus rubecula [L: (11, 13–17, 28, 31, 34, 36, 39, 41, 43, 44, 65, 66, 70, 71, 73, 75, 76, 78, 79, 93, 105, 106, 109, 112^a, 113, 116, 117, 119–122, 124, 126, 128–133, 135, 136, 138, 139, 141, 142, 156); N: (11, 13–17, 21, 28, 31, 34, 36, 39, 41, 43, 65, 66, 70, 71, 73, 78, 79, 82, 93, 104–106, 109–112^a, 113, 116–122, 124, 127–133, 135, 136, 138–142, 149); F: (13, 39); A: (118); NA: (33, 48, 57, 73, 115, 123, 126, 143–145, 155)]

• - European serin – Serinus serinus [L: (79); N: (17, 34, 79); NA: (33)]

• - Fieldfare -Turdus pilaris [L: (14, 34, 75, 105, 112, 113); N: (14, 31, 34, 36, 74, 75, 105, 112^a, 113); NA: (26, 47, 48, 57, 115)]

• - Garden warbler – Sylvia borin [L: (36, 114, 119); N: (31, 34, 36, 70, 104, 105, 109, 112–114, 116, 119, 131); NA: (26, 33)]

• - Goldcrest – Regulus regulus [L: (13, 31, 39, 41, 73); N: (13, 31, 39, 41, 73, 104, 105, 119, 141); NA: (145)]

• - Golden eagle – Aquila chrysaetos [F: (34)]

• - Great reed warbler – Acrocephalus arundinaceus [N: (104, 109, 112^a, 140, 142)]

• - Great spotted woodpecker – Dendrocopos major [L: (105, 106); N: (105, 106); NA: (33)]

• - Great tit – Parus major [L: (13–15, 17, 21, 28, 31, 34, 36, 39–41, 43, 50, 51, 65, 70, 76, 78, 79, 105, 106, 116, 117, 120, 122, 124, 126, 129, 130, 132, 133, 135–137, 139, 140); N: (13–15, 17, 21, 25, 28, 34, 36, 39–41, 43, 50–52, 57, 65, 70, 73, 78, 79, 104–106, 112^a, (116, 117, 120, 122, 124, 126, 129, 131, 133, 135–142, 146); M: (52); F: (52, 70); A: (124); NA: (29, 33, 115, 123, 143–145, 147, 157)]

• - Greenish warbler – Phylloscopus trochiloides [L: (119); N: (36, 111)]

• - Gray partridge – Perdix perdix [L: (31); N: (31, 108); NA: (14, 47)]

• - Gray wagtail – Motacilla cinerea [L: (122); N: (57, 122)]

• - Hawfinch – Coccothraustes coccothraustes [L: (33, 79, 121, 129, 135, 137, 139); N: (28, 33, 34, 65, 70, 79, 109, 112, 113, 121, 128, 129, 135– 137, 139, 140); A: (140); NA: (57)]

• - Hazel grouse – Tetrastes bonasia [NA: (47)]

• - Hooded crow – Corvus cornix [NA: (33, 47)]

• - House sparrow – Passer domesticus [L: (70, 126, 151); N: (34, 70, 73); NA: (33, 47, 123)]

• - Iberian chiffchaff – Phylloscopus ibericus [L: (15)]

• - Icterine warbler – Hippolais icterina [L: (73, 105, 112, 113, 119); N: (34, 36, 70, 73, 112, 113, 132); NA: (48)]

• - Lesser redpoll – Carduelis cabaret [L: (109); N: (70, 112, 113)]

• - Lesser whitethroat – Sylvia curruca [L: (41, 73, 105, 106, 132); N: (31, 36, 41, 70, 73, 105, 106, 110–113, 132, 140)]

• - Long-eared owl – Asio otus [N: (34); NA: (26, 30)]

• - Long-tailed tit – Aegithalos caudatus [L: (78)]

• - Mallard – Anas platyrhynchos [NA: (26)]

• - Marsh tit – Poecile palustris [L: (116, 120, 129); N: (28, 36, 39, 116, 126, 129, 138); NA: (29, 33)]

• - Marsh warbler – Acrocephalus palustris [L: (28, 31, 34, 132); N: (28, 31, 34, 70, 73, 104, 105, 109, 112^a, 124, 131, 132, 140); NA: (143, 144)]

• - Meadow pipit – Anthus pratensis [L: (24, 28, 31, 76, 107); N: (24, 31, 43, 70, 75, 107, 112, 113); M: (110); NA: (48)]

• - Melodious warbler – Hippolais polyglotta [L: (71); N: (78)]

• - Mew gull – Larus canus [L: (31); N: (31, 151)]

• - Middle spotted woodpecker – Dendrocoptes medius (NA: (26, 33)]

• - Mistle trush – Turdus viscivorus [L: (70, 79, 123, 133); N: (34, 79, 123, 133); NA: (30, 33, 47)]

• - Northern goshawk – Accipiter gentilis [N: (108)]

• - Northern lapwing – Vanellus vanellus [F: (43); NA: (30)]

• - Oriental turtle dove – Streptopelia orientalis [N: (52)]

• - Pechora pipit – Anthus gustavi [NA: (30)]

• - Pied wheatear – Oenanthe pleschanka [NA: (30)]

• - Red crossbill – Loxia curvirostra [N: (36); NA: (30)]

• - Red kite – Milvus milvus [N: (34)]

• - Red-backed shrike – Lanius collurio [L: (105); N: (73, 104, 105, 112^a, 129); NA: (26, 33, 47)]

• - Redwing – Turdus iliacus [L: (31, 36, 41, 65, 70, 73, 79, 105, 106, 109, 124, 136, 141); N: (31, 34, 36, 39, 41, 65, 70, 73, 76, 79, 104–106, 109, 111–114, 124, 128, 136, 141, 142); NA: (14, 48, 115, 144)]

• - Ring ouzel – Turdus torquatus [N: (34, 138); NA: (30)]

• - River warbler – Locustella fluviatilis [N: (129)]

• - Rock bunting – Emberiza cia [N: (120)]

• - Rook – Corvus frugilegus [L: (55); N: (52); M: (52); F: (52); NA: (26)]

• - Rustic bunting – Emberiza rustica [NA: (26)]

• - Sand martin – Riparia riparia [L: (95 in nest); N: (95 in nest)]

• - Sardinian warbler – Sylvia melanocephala [L: (15); N: (21)]

• - Savi's warbler – Locustella luscinioides [L: (65); N: (65, 142)]

• - Sedge warbler – Acrocephalus schoenobaenus [L: (65, 128); N: (28, 65, 70, 104, 110, 112^a, 128); NA: (30)]

• - Short-toed treecreeper – Certhia brachydactyla [L: (15, 17, 34); N: (15, 17)]

• - Skylark – Alauda arvensis [L: (105); N: (33, 70, 79); NA: (30)]

• - Song thrush – Turdus philomelos [L: (11, 13–15, 24, 31, 34, 36, 39, 41, 44, 57, 65, 70, 73, 76, 78, 79, 93, 105, 106, 109, 112, 114, 116, 117, 119–122, 124, 126, 128, 130, 131, 133, 135–139, 141); N: (11, 13, 14, 16, 21, 24, 31, 34, 36, 39, 41, 44, 57, 65, 70, 73, 75, 76, 78, 79, 93, 104–106, 108, 109, 111–114, 116, 117, 119–122, 124, 126, 128, 130–133, 135–139, 141, 142, 146); NA: (33, 35, 115, 143, 144, 147)]

• - Spotted flycatcher – Muscicapa striata [L: (110, 120, 132); N: (31, 34, 110)]

• - Spotted nutcracker – Nucifraga caryocatactes (N: (79); NA: (26)]

• - Tawny owl -Strix aluco [NA: (47)]

• - Thrush nightingale – Luscinia luscinia [L: (16, 31, 36, 41, 65, 73, 105, 106, 112^a, 119, 124, 128, 132, 138); N: (31, 36, 41, 65, 73, 105, 106, 112^a, 119, 124, 128, 138, 140); A: (140); NA: (144)]

• - Tree pipit – Anthus trivialis [L: (21, 31, 34, 36, 41, 43, 73, 79, 105, 106, 110, 121, 128, 132, 136); N: (24, 31, 34, 36, 41, 43, 73, 79, 104– 106, 110, 121, 128, 132, 136, 139); NA: (33, 35, 48)]

• - Turkey – Meleagris gallopavo – [NA: (14)]

• - Western jackdaw – Corvus monedula [L: (55); N: (55); NA: (33)]

• - Western yellow wagtail – Motacilla flava [L: (34, 36); N: (31, 34, 110)]

• - Wheatear – Oenanthe oenanthe [N: (31, 76, 79, 112, 113); NA: (59)]

• - Whinchat – Saxicola rubetra [N: (112, 113, 138); NA: (30)]

• - White wagtail – Motacilla alba [L: (73); N: (24, 52); M: (52)]

• - Willow ptarmigan – Lagopus lagopus [L: (107, 158, 159); N: (107, 158, 159)]

• - Willow tit – Poecile montana [L: (31, 39, 70, 120, 126); N: (31, 34, 39, 70, 112^a, 120, 126, 130); NA: (26, 57)]

• - Willow warbler – Phylloscopus trochilus [L: (11, 21, 31, 34, 36, 41, 43, 70, 73, 75, 79, 105, 106, 112, 113, 117, 119, 132, 138); N: (11, 28, 31, 34, 36, 41, 43, 70, 73, 75, 78–80, 104–106, 110–112^a, 24, 113, 114, 138); NA: (33)]

• - Winter wren – Troglodytes troglodytes [L: (15, 17, 21, 31, 34, 39, 41, 44, 65, 66, 70, 73, 78, 79, 105, 106, 114, 116, 117, 122, 129, 132, 133, 139, 141, 142); N: (15, 17, 21, 31, 34, 36, 39, 65, 66, 70, 73, 79, 104–106, 114, 116, 122–124, 129, 130, 133, 138–141); F: (25); NA: (33, 57, 73, 115, 143–145)]

• - Wood warbler – Phylloscopus sibilatrix [L: (36, 41, 105, 106); N: (36, 41); NA: (33)]

• - Woodlark – Lullula arborea [L: (112, 113); N: (79, 112, 113)]

• - Yellow-browed warbler – Phylloscopus inornatus [NA: (48)]

• - Yellowhammer – Emberiza citrinella [L: (33, 93); N: (33, 39, 93, 129); NA: (14, 26, 57)]

Distribution of reported cases: United Kingdom (24, 25, 30, 46, 53, 75, 76, 80, 107, 117, 150, 154, 158, 159), Sweden (31, 36, 56!, 73, 102!, 105, 106, 110, 119), Slovakia (33, 120, 126, 138), Poland (26, 39, 41, 51, 108, 121, 125, 135, 137), Switzerland (34, 79, 82, 115, 116, 118, 157), Latvia (130, 141), Germany (61!, 74, 124, 127, 131, 134, 143, 144), Czech Republic (28, 29, 57, 95!, 122, 129, 148, 149), Belgium (40, 48, 50, 152), Italy (22!, 78, 123, 136, 155, 156), Netherlands (43, 70, 151), Estonia (104), Norway (11, 109, 111–113), Hungary (47, 65, 128, 142), Faroe Islands (59), Denmark (132), Romania (13, 14, 23!, 55), France (133), Moldova (88!, 93), Russia (Kaliningrad, 114, 139, 147, 146), Ukraine (52), Portugal (15, 17, 21), Spain (44, 66, 71), Lithuania (112^a, 145), Croatia (35), Bulgaria (140, 153), Cyprus (16).

Ixodes rothschildi

Overview: Ixodes rothschildi has so far been reported from birds only in the United Kingdom and France. This tick species is a parasite of seabirds, collected from 4 charadriiform, 1 suliform, and 1 procellariform species.

Charadriiformes: 4; Suliformes: 1; Procellariiformes: 1

Hosts:

• - Atlantic puffin – Fratercula arctica [L: (76, 160); N: (76, 160); F: (76); NA: (18)]

• - Common murre – Uria aalge [NA: (30)]

• - European herring gull – Larus argentatus [NA: (30)]

• - European shag – Phalacrocorax aristotelis [F: (75)]

• - Manx shearwater – Puffinus puffinus [NA: (30)]

• - Razorbill – Alca torda [N: (161); F: (161)in burrows]

Distribution of reported cases: United Kingdom (30, 75, 76, 160, 161), France (18).

Ixodes unicavatus

Overview: Ixodes unicavatus is distributed in the coastal area of the UK, Sweden and France. The hosts are birds that usually nest on rocky cliffs. Six of such bird species have been described as hosts of this parasite.

Charadriiformes: 1; Suliformes: 3; Passeriformes: 2; Falconiformes: 1

Hosts:

• - Atlantic puffin – Fratercula arctica [NA: (30)]

• - Common shag – Gulosus aristotelis [NA: (30)]

• - European rock pipit – Anthus petrosus [NA: (30)]

• - Great cormorant – Phalacrocorax carbo [L: (31); N: (31); F: (31); NA: (30)]

• - Gyrfalcon – Falco rusticolus [NA: (30)]

• - Water pipit – Anthus spinoletta [L: (24)]

Distribution of reported cases: United Kingdom (24, 30), Sweden (31), France (18!).

Ixodes uriae

Overview: Ixodes uriae is a common parasite of North European seabirds, as it was reported from 8 charadriiform, 1 anseriform, 3 suliform, and 1 procellariform bird species. According to the data it was also recorded from 2 passeriform bird species.

Charadriiformes: 8; Anseriformes: 1; Suliformes: 3; Procellariiformes: 1; Passeriformes: 3

Hosts:

• - Atlantic puffin – Fratercula arctica [N: (24, 76, 162 in nest); M: (162 in nest); F: (24, 162 in nest); NA: (59, 163^a, 163^b); (164^a, 164^b, 165^a, 166)]

• - Black guillemot – Cepphus grylle [NA: (30)]

• - Black-legged kittiwake – Rissa tridactyla (L: (167); N: (162 in nest, 167–170); M: (162 in nest, 167); F: (161, 162 in nest, 167, 169); NA: (163^b, 164^a, 164^b, 165^a, 170, 171^a, 171^b, 172, 173)]

• - Common eider – Somateria mollissima [NA: (174)]

• - Common murre – Uria aalge [N: (31, 162 in nest, 175–177); M: (162 in nest, 175); F: (31, 75, 161, 162 in nest, 175–177); NA: (59, 163^b, 164^a, 164^b, 165^a)]

• - Common starling – Sturnus vulgaris [NA: (30)]

• - Eurasian curlew – Numenius arquata [NA: (30)]

• - European herring gull – Larus argentatus [N: (170 in nest); F: (170 in nest); NA: (30)]

• - European shag – Phalacrocorax aristotelis [N: (76)]

• - Northern fulmar – Fulmarus glacialis [NA: (30)]

• - Northern gannet – Morus bassanus [N: (76); F: (76)]

• - Razorbill – Alca torda [N: (177); F: (177); NA: (30, 163^b, 164^a, 164^b)]

• - Red crossbill – Loxia curvirostra [NA: (30)]

• - Red-faced cormorant – Urile urile [NA: (163^b)]

• - Thick-billed murre – Uria lomvia [N: (175^a); M: (175^a); F: (175); NA: (178–180)]

• - Wheatear – Oenanthe oenanthe [NA: (30)]

Distribution of reported cases: United Kingdom (24, 30, 76, 161, 163, 164, 167–169, 171, 172, 176), France (170, 171^a), Norway (Svalbard) (175^a, 178, 179), Norway (162, 163^a, 164^a, 165, 171^b, 173, 177, 180), Norway (Jan Mayen) (175), Iceland (163^b, 164^b, 165, 174), Faeroe Islands (59, 166), Sweden (31).

Ixodes ventalloi

Overview: Ixodes ventalloi, commonly known as the rabbit tick as it mostly feeds on the European rabbit (Oryctolagus cuniculus), is a rare finding on birds (8). This tick species has been recorded from 11 bird species. Four species belong to the order Strigiformes, which are often in contact with small mammals.

Strigiformes: 4; Passeriformes: 3; Galliformes: 3, Gruiformes: 1

Hosts:

• - Barn owl – Tyto alba [N: (60)]

• - Black redstart – Phoenicurus ochruros [N: (84, 181)]

• - Chukar partridge- Alectoris chukar [NA: (182)]

• - Common blackbird – Turdus merula [N: (84); F: (21); NA: (25)]

• - Common pheasant – Phasianus colchicus [A: (19)]

• - European robin – Erithacus rubecula [N: (181)]

• - Little owl – Athene noctua [F: (183)]

• - Long-eared owl – Asio otus [N: (76); F: (76)]

• - Red-legged partridge – Alectoris rufa [A: (19); NA: (90)]

• - Short-eared owl – Asio flammeus [N: (85); M: (85); F: (85)]

• - Water rail – Rallus aquaticus [A: (19)]

Distribution of reported cases: Cyprus (182), Spain (90, 183), Portugal (21, 60, 85), Italy (19, 84, 181), United Kingdom (25, 76).

Metastriata

Amblyomma lepidum

Overview: While the immature stages of Amlyomma lepidum are relatively common bird parasites in Africa (10), this tick species has been reported only once in European territory, from a passeriform bird.

Passeriformes: 1

Hosts:

• - Common blackbird – Turdus merula [N: (16)]

Distribution of reported cases: Cyprus (16).

Amblyomma marmoreum

Overview: Amblyomma marmoreum is an African tick species, where it was reported from several bird species (10). In Europe, it has been found on a Tree pipit in Italy.

Passeriformes: 1

Hosts:

• - Tree pipit – Anthus trivialis [N: (84)]

Distribution of reported cases: Italy (84).

Amblyomma nuttalli

Overview: Amlyomma nuttali is distributed in the Afrotropical region, where they may infest several bird species (10). So far, we have only one report in Europe, where it was found on a Thrush nightingale (Passeriformes).

Passeriformes: 1

Hosts:

• - Thrush nightingale – Luscinia luscinia [N: (16)]

Distribution of reported cases: Cyprus (16).

Amblyomma variegatum

Overview: Amblyomma variegatum is an African tick species (10) and as such, it does not have European distribution. However, it was reported from 2 passeriform birds in Italy, and from one passeriform in Cyprus.

Passeriformes: 2

Hosts:

• - Icterine warbler – Hyppolais icterina [N: (184)]

• - Tree pipit – Anthus trivialis [N: (16, 181)]

Distribution of reported cases: Italy (181, 184), Cyprus (16).

Dermacentor reticulatus

Overview: Dermacentor reticulatus has a pan European distribution excepting Scandinavia (8), however its occurrence on birds is extremely rare. The only reports are from the United Kingdom and from Poland.

Passeriformes: 2

Hosts:

• - European robin – Erithacus rubecula [L: (39)]

• - Meadow pipit – Anthus pratensis [N: (75)]

Distribution of reported cases: Poland (39), United Kingdom (75).

Dermacentor marginatus

Overview: Dermacentor marginatus is a common parasite throughout Europe, but not in Scandinavia (8). Despite this fact, it is rare to find these ticks on birds. D. marginatus has been reported from 1 passeriform and 1 galliform bird species, as well as from the nest of a passeriform bird.

Passeriformes: 2; Galliformes: 1

Hosts:

• - Bearded reedling – Panurus biarmicus [L: (185 in nest); N: (185 in nest)]

• - Turkey – Meleagris gallopavo – [A: (60)]

• - Yellowhammer – Emberiza citrinella [NA: (14, 47)]

Distribution of reported cases: Austria (185), Hungary (47), Portugal (60), Romania (14).

Haemaphysalis concinna

Overview:Haemaphysalis concinna has a pan-European distribution. Interestingly, H. concinna seems to be extremely common on birds in Hungary, compared to other European countries. This parasite was reported from 36 different bird species so far. Thirty-three of these belong to the order Passeriformes.

Passeriformes: 33; Accipitriformes: 1; Galliformes: 1; Charadriiformes: 1

Hosts:

• - Barred warbler – Sylvia nisoria [NA: (186)]

• - Black kite – Milvus migrans [NA: (47)]

• - Common blackbird – Turdus merula [L: (65); N: (52, 65, 129, 140); NA: (33, 186)]

• - Common chaffinch – Fringilla coelebs [NA: (33)]

• - Common grasshopper warbler – Locustella naevia [L: (65); N: (65)]

• - Common nightingale – Luscinia megarhynchos [L: (65); N: (65); NA: (33, 186)]

• - Common pheasant – Phasianus colchicus [NA: (33)]

• - Common reed bunting – Emberiza schoeniclus (L: (65); N: (65)]

• - Common starling – Sturnus vulgaris [NA: (33)]

• - Dunnock – Prunella modularis [L: (65, 128, 129); N: (65); NA: (186)]

• - Eurasian blackcap – Sylvia atricapilla [L: (17, 65, 128); N: (65, 128); NA: (186)]

• - Eurasian blue tit- Cyanistes caeruleus [NA: (33)]

• - Eurasian golden oriole – Oriolus oriolus (N: (33)]

• - Eurasian magpie – Pica pica [NA: (33)]

• - Eurasian nuthatch – Sitta europaea [NA: (33)]

• - Eurasian reed warbler – Acrocephalus scirpaceus [L: (65, 128); N: (65, 128); NA: (186)]

• - Eurasian tree sparrow – Passer montanus [NA: (33)]

• - Eurasian woodcock – Scolopax rusticola [NA: (33)]

• - European greenfinch – Carduelis chloris [L: (17); N: (65); NA: (186)]

• - European robin – Erithacus rubecula [L: (65, 129); N: (65, 142); NA: (33, 186)]

• - Great reed warbler – Acrocephalus arundinaceus [NA: (186)]

• - Great tit – Parus major [L: (65); N: (52, 65); NA: (33, 186)]

• - Hawfinch – Coccothraustes coccothraustes [L: (65); N: (65); NA: (14, 33, 186)]

• - House sparrow – Passer domesticus [NA: (33)]

• - Lesser whitethroat – Sylvia curruca [NA: (186)]

• - Marsh warbler – Acrocephalus palustris [L: (65); N: (65); NA: (186)]

• - Red-backed shrike – Lanius collurio [NA: (33, 186)]

• - River warbler – Locustella fluviatilis [N: (129); NA: (186)]

• - Rook – Corvus frugilegus [L: (55)]

• - Savi's warbler – Locustella luscinioides [L: (65, 128); N: (65); NA: (186)]

• - Sedge warbler – Acrocephalus schoenobaenus [L: (65); N: (65); NA: (186)]

• - Song thrush – Turdus philomelos [L: (65); N: (65); NA: (33, 186)]

• - Tree pipit – Anthus trivialis [NA: (33, 47)]

• - Willow warbler – Phylloscopus trochilus [L: (16)]

• - Winter wren – Troglodytes troglodytes [L: (129)]

• - Yellowhammer – Emberiza citrinella [L: (33, 65, 129); N: (33, 65, 129); NA: (186)]

Distribution of reported cases: Czech Republic (57!, 129), Slovakia (33), Hungary (47, 65, 128, 142, 186), Romania (23!, 55), Ukraine (52), Bulgaria (140), Cyprus (16).

Haemaphyalis erinacei

Overview: Haemaphysalis erinacei (as its name suggests) is mainly a parasite of hedgehogs and small mammals (8). Finding this tick on birds is a rare event. So far, it has been reported from the South-Eastern European region, where H. erinacei was found on on 4 different bird species. Interestingly, these 4 birds belonged to 4 different orders.

Caprimulgiformes: 1; Strigiformes: 1, Gruiformes: 1, Coraciiformes: 1

Hosts:

• - Alpine swift – Apus melba [N: (187^a)]

• - Common Crane – Grus grus [M: (187)]

• - Eurasian eagle-owl – Bubo bubo [F: (153)]

• - European Bee-eater – Merops apiaster [M: (187)]

Distribution of reported cases: Bulgaria (153), Croatia (187^a), Bosnia and Herzegovina (187).

Haemaphyalis parva

Overview: Haemaphysalis parva is a rare tick species on European birds. In Romania, it was found on 2 passeriforms, and on 1 galliform bird.

Passeriformes: 2; Galliformes: 1

Hosts:

• - Common snipe – Gallinago gallinago [NA: (14)]

• - Rook – Corvus frugilegus [L: (55); N: (55)]

• - Western jackdaw – Corvus monedula [L: (55)]

Distribution of reported cases: Romania (14, 55).

Haemaphysalis punctata

Overview: Haemaphysalis punctata has a pan-European distribution, and its immature stages are fairly common parasites on birds, especially on Passeriformes.

Passeriformes: 40; Charadriiformes: 7; Falconiformes: 1; Strigiformes: 1; Galliformes: 2

Hosts:

• - Black-headed gull – Chroicocephalus ridibundus [F: (31); NA: (30)]

• - Cirl bunting – Emberiza cirlus [L: (21, 66, 71); N: (21, 85)]

• - Common blackbird – Turdus merula [L: (15, 17, 21, 66, 71, 72); N: (13, 15, 21, 66, 69, 71, 85, 188); NA: (30, 33, 46, 47)]

• - Common chaffinch – Fringilla coelebs [L: (21, 66, 71); N: (69, 71, 188)]

• - Common kestrel- Falco tinnunculus [NA: (30)]

• - Common linnet – Carduelis cannabina [L: (73)]

• - Common nightingale – Luscinia megarhynchos [N: (71)]

• - Common redstart – Phoenicurus phoenicurus [L: (151); F: (31, 110); NA: (26)]

• - Common reed bunting – Emberiza schoeniclus [NA: (30)]

• - Common starling – Sturnus vulgaris [L: (73); NA: (30)]

• - Common tern – Sterna hirundo [NA: (30)]

• - Common whitethroat – Sylvia communis [L: (73); N: (31); NA: (30)]

• - Dunnock – Prunella modularis [L: (66); N: (39, 66); NA: (30)]

• - Eurasian blackcap – Sylvia atricapilla [N: (69)]

• - Eurasian blue tit – Cyanistes caeruleus [L: (73)]

• - Eurasian curlew – Numenius arquata [N: (151)]

• - Eurasian eagle-owl – Bubo bubo [F: (153)]

• - Eurasian jay – Garrulus glandarius [L: (15); NA: (17, 47)]

• - Eurasian magpie – Pica pica [NA: (47)]

• - Eurasian oystercatcher – Haematopus ostralegus [L: (31); N: (31)]

• - Eurasian reed warbler – Acrocephalus scirpaceus [N: (85)]

• - Eurasian stonechat – Saxicola torquatus [NA: (30)]

• - Eurasian woodcock – Scolopax rusticola [NA: (33)]

• - European greenfinch – Carduelis chloris [L: (71)]

• - European herring gull – Larus argentatus [NA: (30)]

• - European robin – Erithacus rubecula [L: (21, 66, 156); N: (66, 69); F: (140); NA: (30)]

• - European serin – Serinus serinus [L: (71)]

• - Great black-backed gull – Larus marinus [NA: (26)]

• - Great tit – Parus major [L: (21, 66); N: (188)]

• - Hooded crow – Corvus cornix [NA: (47)]

• - House sparrow – Passer domesticus [L: (71, 151); N: (71, 151, 188); NA: (30)]

• - Meadow pipit – Anthus pratensis [L: (151); NA: (30)]

• - Melodious warbler – Hippolais polyglotta [NA: (30)]

• - Mistle trush – Turdus viscivorus [NA: (30)]

• - Red-billed chough – Pyrrhocorax pyrrhocorax (NA: (30)]

• - Red-legged partridge – Alectoris rufa [NA: (87)]

• - Ring ouzel – Turdus torquatus [NA: (47)]

• - Rook – Corvus frugilegus [L: (55, 188); N: (55, 188); M: (55); F: (55); A: (188)]

• - Skylark – Alauda arvensis [NA: (30)]

• - Song thrush – Turdus philomelos [L: (93); N: (66, 93); NA: (30)]

• - Spotless starling – Sturnus unicolor [L: (71); N: (71)]

• - Spotted flycatcher – Muscicapa striata [L: (16)]

• - Tree pipit – Anthus trivialis [L: (21)]

• - Turkey – Meleagris gallopavo – [N: (60); A: (60); NA: (14)]

• - Western jackdaw – Corvus monedula [N: (55); M: (55); NA: (30)]

• - Western yellow wagtail – Motacilla flava [NA: (35)]

• - Wheatear – Oenanthe oenanthe [L: (73); NA: (30)]

• - Whinchat – Saxicola rubetra [L: (151)]

• - White wagtail – Motacilla alba [L: (73); N: (31, 110); NA: (30)]

• - Willow warbler – Phylloscopus trochilus [L: (16); NA: (30)]

• - Yellowhammer – Emberiza citrinella [NA: (30, 33)]

Distribution of reported cases: Ukraine (188), Spain (66, 71, 87), Poland (26, 39), United Kingdom (30, 46), Slovakia (33), Netherlands (151), Sweden (31, 73, 110), Portugal (Azores, 69), Moldova (93), Portugal (15, 17, 21, 60, 85), Romania (13, 14, 55), Italy (156), Hungary (47), Croatia (35), Bulgaria (140, 153), Cyprus (16).

Haemaphyalis sulcata

Overview: Haemaphysalis sulcata is a relatively rare parasite on birds. According to literature data, it has been recorded from 5 passeriform bird species so far.

Passeriformes: 5

Hosts:

• - Common blackbird – Turdus merula [N: (140)]

• - Great reed warbler – Acrocephalus arundinaceus [N: (140)]

• - Savi's warbler – Locustella luscinioides [N: (140)]

• - Skylark – Alauda arvensis [N: (24)]

• - Thrush nightingale – Luscinia luscinia [N: (140)]

Distribution of reported cases: United Kingdom (24), Bulgaria (140).

Hyalomma aegyptium

Hosts:

Overview: Although Hyalomma aegyptium is a common tick in the Mediterranean region, it is primarily a parasite of tortoises and humans (8). Therefore, finding these ticks on European birds is an uncommon event. The only European report of this tick feeding on birds so far is from Greece, where this parasite has been reported from 2 passeriform species.

Passeriformes: 2

• - Common blackbird – Turdus merula [L: (20)]

• - Common nightingale – Luscinia megarhynchos [L: (20)]

Distribution of reported cases: Greece (20).

Hyalomma lusitanicum

Overview: Hyalomma lusitanicum is a common parasite of mammals in Spain and Portugal (8, 10). Despite this fact, finding this tick species on bird is occasional. So far, it has been recorded from 2 galliform, 1 passeriform, 1 strigiform, 1 struthioniform and 1 columbiform birds from the Iberian Peninsula.

Passeriformes: 1; Galliformes: 2; Strigiformes: 1; Columbiformes: 1, Struthioniformes: 1

Hosts:

• - Chicken – Gallus gallus domesticus [N: (189)]

• - Common blackbird – Turdus merula [L: (15); N (15)]

• - Common ostrich – Struthio camelus [A: (60)]

• - Red-legged partridge – Alectoris rufa [N: (60); NA: (87)]

• - Eurasian eagle-owl – Bubo bubo [N: (60)]

• - Common wood pigeon – Columba palumbus [A: (60)]

Distribution of reported cases: Spain (87), Portugal (15, 60, 189).

Hyalomma marginatum (s.l.)

Overview: Hyalomma marginatum is a two-host tick species (10) that has a Palearctic distribution. While it is more common in the Mediterranean region, this parasite is also reported occasionally from Northern European countries. The most common bird hosts of H. marginatum are from the order Passeriformes.

Passeriformes: 52; Accipitriformes: 5; Galliformes: 1; Strigiformes: 5; Falconiformes: 2; Coraciiformes: 1; Bucerotiformes: 1; Caprimulgiformes: 1, Columbiformes: 1

Hosts:

• - Barn owl – Tyto alba [L: (85); N: (85)]

• - Barn swallow – Hirundo rustica [N: (85)]

• - Black kite – Milvus migrans [N: (85)]

• - Black redstart – Phoenicurus ochruros [N: (16)]

• - Bluethroat – Luscinia svecica [N: (190, 191); NA: (191^b)]

• - Booted eagle – Hieraaetus pennatus [N: (60)]

• - Chicken – Gallus gallus domesticus [NA: (14)]

• - Collared flycatcher – Ficedula albicollis [L: (31)]

• - Common blackbird – Turdus merula [L: (15, 71); N: (16, 71, 85, 192)]

• - Common buzzard – Buteo buteo [N: (60)]

• - Common chaffinch – Fringilla coelebs [L: (16, 71); N: (16); F: (73)]

• - Common chiffchaff – Phylloscopus collybita [L: (16)]

• - Common kestrel – Falco tinnunculus [L: (60); N: (60); NA: (191^d)]

• - Common kingfisher – Alcedo atthis [N: (60, 85)]

• - Common nightingale – Luscinia megarhynchos [L: (71, 193^a); N: (71, 140, 191^e, 193^a)]

• - Common redstart – Phoenicurus phoenicurus [N: (16, 80, 82, 184, 190, 191^c); NA: (30, 191^b, 191^d)]

• - Common whitethroat – Sylvia communis [N: (80, 140, 191^c, 194); NA: (30)]

• - Corn bunting – Emberiza calandra [L: (16)]

• - Crested lark – Galerida cristata [L: (16); N: (16)]

• - Dunnock – Prunella modularis [N: (195)]

• - Eastern olivaceous warbler – Iduna pallida [N: (20)]

• - Eastern woodchat shrike – Lanius senator niloticus [N: (196)]

• - Eurasian blackcap – Sylvia atricapilla [L: (31); N: (16, 191^c)]

• - Eurasian blue tit- Cyanistes caeruleus [N: (71, 85)]

• - Eurasian eagle-owl – Bubo bubo [L: (153); N: (60, 85, 153); M: (153)]

• - Eurasian hoopoe – Upupa epops [NA: (191^d)]

• - Eurasian jay – Garrulus glandarius [N: (15)]

• - Eurasian reed warbler – Acrocephalus scirpaceus [L: (193); N: (60, 85); M: (190); F: (190, 193^a); NA: (38)]

• - Eurasian scops owl – Otus scops [L: (16); N: (16)]

• - Eurasian stonechat – Saxicola torquatus [L: (16); N: (16, 60, 85)]

• - Eurasian tree sparrow – Passer montanus [N: (85)]

• - European greenfinch – Carduelis chloris [L: (16); N: (16, 71, 85)]

• - European nightjar – Caprimulgus europaeus [N: (184)]

• - European pied flycatcher – Ficedula hypoleuca [L: (71)]

• - European robin – Erithacus rubecula [L: (73, 142); N: (16, 82, 142, 191^a, 195)]

• - European turtle dove – Streptopelia turtur [L: (16)]

• - Finsch's wheatear – Oenanthe finschii [L: (16); N: (16)]

• - Great reed warbler – Acrocephalus arundinaceus [L: (16); N: (193^a)]

• - Great tit – Parus major [L: (71); N: (71, 85)]

• - House sparrow – Passer domesticus [L: (71); N: (71)]

• - Iberian gray shrike – Lanius meridionalis [N: (85)]

• - Lesser kestrel – Falco naumanni [N: (60)]

• - Lesser whitethroat – Sylvia curruca (L: (16)]

• - Little owl – Athene noctua [N: (85); NA: (14)]

• - Marsh warbler – Acrocephalus palustris [L: (191^a); N: (16, 140, 191^a, 191^c); NA: (193^a)]

• - Mistle trush – Turdus viscivorus [N: (190, 192)]

• - Northern goshawk – Accipiter gentilis [N: (60)]

• - Ortolan bunting – Emberiza hortulana [N: (191^b)]

• - Red kite – Milvus milvus [L: (85)]

• - Rook – Corvus frugilegus [N: (192)]

• - Sardinian warbler – Sylvia melanocephala [L: (16); N: (16)]

• - Savi's warbler – Locustella luscinioides [N: (193)]

• - Sedge warbler – Acrocephalus schoenobaenus [N: (31, 80, 193, 194, 196); F: (193); NA: (30, 102)]

• - Song thrush – Turdus philomelos [L: (16); N: (16, 197)]

• - Spanish Sparrow – Passer hispaniolensis [L: (16)]

• - Spotted flycatcher – Muscicapa striata [L: (16); N: (14)]

• - Tawny owl -Strix aluco [N: (60)]

• - Thrush nightingale – Luscinia luscinia [L: (16)]

• - Tree pipit – Anthus trivialis [L: (16); N: (16, 36, 191^b)]

• - Water pipit – Anthus spinoletta [L: (16)]

• - Western jackdaw – Corvus monedula [N: (55)]

• - Western olivaceous warbler – Iduna opaca [N: (194)]

• - Western yellow wagtail – Motacilla flava [L: (16, 196); N: (16, 184); NA: (102)]

• - Wheatear – Oenanthe oenanthe [N: (80); NA: (30)]

• - Whinchat – Saxicola rubetra [N: (16); NA: (30)]

• - White wagtail – Motacilla alba [L: (16); N: (16, 31, 190)]

• - Willow warbler – Phylloscopus trochilus [L: (16, 194); N: (73)]

• - Woodchat shrike – Lanius senator [L: (198); N: (198)]

• - Yellowhammer – Emberiza citrinella [NA: (14)]

Distribution of reported cases: Switzerland (82, 190), Sweden (31, 36, 56!, 73), Slovakia (191^a, 193), Czech Republic (191, 193^a), Italy (184, 191^e, 199!), Greece (20, 196, 198), United Kingdom (30, 80), Hungary (142), Romania (14, 55, 197), Portugal (15, 60, 85), Spain (71, 194), France (18!, 195), Poland (102), Germany (38, 191^c), Finland (191^b), Slovenia (191^d), Ukraine (192), Bulgaria (140, 153), Cyprus (16).

Hyalomma rufipes

Overview: Hyalomma rufipes is a two-host tick species (8). It is a widely distributed, common bird parasite in the Mediterranean region. It isc occasionally reported on migrating birds from Northern European countries as well. Similarly to H. marginatum [in the past, H rufipes was considered as a subspecies of H. marginatum (10)] the most common bird hosts belong to Passeriformes.

Passeriformes: 46; Cuculiformes: 1; Piciformes: 1; Columbiformes: 1, Flaconiformes: 1, Bucerotiformes: 1, Strigiformes: 1

Hosts:

• - Barred warbler – Sylvia nisoria [N: (16)]

• - Barn swallow – Hirundo rustica [M: (200)]

• - Black redstart – Phoenicurus ochruros [N: (181); NA: (84)]

• - Black-eared Wheatear – Oenanthe hispanica [L: (16); N: (16)

• - Blue rock thrush – Monticola solitarius [L: (16)]

• - Crested lark – Galerida cristata [L: (16); N: (16)]

• - Cyprus warbler – Sylvia melanothorax [N: (16)]

• - Collared flycatcher – Ficedula albicollis [N: (16, 181, 184); NA: (84)]

• - Common blackbird – Turdus merula [L: (16); N: (16)]

• - Common chaffinch – Fringilla coelebs [N: (16)]

• - Common chiffchaff – Phylloscopus collybita [L: (16)]

• - Common cuckoo – Cuculus canorus [N: (16, 184)]

• - Common kestrel- Falco tinnunculus [L: (16); N: (16)]

• - Common nightingale – Luscinia megarhynchos [L: (184); N: (16, 184, 201)]

• - Common redstart – Phoenicurus phoenicurus [L: (16, 184); N: (11, 16, 181, 184); NA: (84)]

• - Common whitethroat – Sylvia communis [L: (184); N: (11, 16, 65, 181, 184); NA: (84)]

• - Corn bunting – Emberiza calandra [L: (16)]

• - Cretzschmar's bunting – Emberiza caesia [L: (16)]

• - Eastern olivaceous warbler – Iduna pallida [L: (16)]

• - Eastern subalpine warbler – Sylvia cantillans [L: (184)]

• - Eurasian blackcap – Sylvia atricapilla [L: (16); N: (16, 195)]

• - Eurasian golden oriole – Oriolus oriolus [N: (181, 184)]

• - Eurasian hoopoe – Upupa epops [N: (16)]

• - Eurasian reed warbler – Acrocephalus scirpaceus [L: (184); N: (11)]

• - Eurasian scops owl – Otus scops [L: (16); N: (16)]

• - Eurasian stonechat – Saxicola torquatus [L: (16); N: (16)]

• - Eurasian wryneck – Jynx torquilla [L: (184)]

• - European goldfinch – Carduelis carduelis [N: (16)]

• - European greenfinch – Carduelis chloris [N: (16)]

• - European pied flycatcher – Ficedula hypoleuca [L: (184), N: (184)]

• - European robin – Erithacus rubecula [N: (16); NA: (84)]

• - European turtle dove – Streptopelia turtur [L: (184)]

• - Garden warbler – Sylvia borin [L: (184); N: (11)]

• - Great reed warbler – Acrocephalus arundinaceus [L: (184); N: (184)]

• - Icterine warbler – Hyppolais icterina [L: (184); N: (184)]

• - Isabelline wheatear – Oenanthe isabellina [N: (16)]

• - Lesser whitethroat – Sylvia curruca [L: (16); N: (16)]

• - Pied wheatear – Oenanthe pleschanka [L: (16); N: (16)]

• - Red-backed shrike – Lanius collurio [L: (16); N: (16)]

• - Red-throated Pipit – Anthus cervinus [L: (16); N: (16)]

• - Rock thrush – Monticola saxatilis [L: (16); N: (16)]

• - Sedge warbler – Acrocephalus schoenobaenus [N: (184, 200); M: (200); F: (200)]

• - Song thrush – Turdus philomelos [L: (16); N: (16); NA: (84)]

• - Spotted flycatcher – Muscicapa striata [L: (184); NA: (84)]

• - Thrush nightingale – Luscinia luscinia [N: (11)]

• - Tree pipit – Anthus trivialis [L: (16, 181); N: (16, 184)]

• - Orphean Warbler – Sylvia hortensis [L: (16)]

• - Western yellow wagtail – Motacilla flava [L: (184, 191^f)]

• - Wheatear – Oenanthe oenanthe [L: (191^f, 16), N: (11, 16, 181, 184); NA: (84)]

• - Whinchat – Saxicola rubetra [L: (184); N: (181, 184, 202); NA: (84)]

• - Willow warbler – Phylloscopus trochilus [L: (184); N: (16)]

• - Wood warbler – Phylloscopus sibilatrix [L: (184); N: (184); NA: (84)]

Distribution of reported cases: Norway (11), Hungary (65), Italy (84, 181, 184, 202), France (Corsica, 200), France (195, 201), Finland (191^f), Cyprus (16).

Hyalomma truncatum

Overview: There is very limited information about Hyalomma truncatum as a bird parasite. According to the data, it has been found on a passeriform bird in Italy.

Passeriformes: 1

Hosts:

• - Whinchat – Saxicola rubetra [L: (184)]

Distribution of reported cases: Italy (184).

Rhipicephalus annulatus

Overview: Despite the fact that Rhipicephalus annulatus is distributed in Southern-Europe (8) it is amazingly rare for this parasite to feed on birds. So far, only one such case was documented in Portugal.

Galliformes: 1

Hosts:

• - Turkey – Meleagris gallopavo – [N: (60); A: (60)]

Distribution of reported cases: Portugal (60).

Rhipicephalus pusillus

Overview: Rhipicephalus pusillus is found in the Palaearctic region, mainly in Mediterranean climatic areas where birds are considered to be exceptional hosts (8). So far, this was recorded only once.

Strigiformes: 1

Hosts:

• - Eurasian eagle-owl – Bubo bubo [A: (60)]

Distribution of reported cases: Portugal (60).

Rhipicephalus sanguineus

Overview: Rhipicephalus sanguineus s.l. is the most common tick species found on dogs in urban areas around the world (8). However, finding them on birds is rare. So far, it has been recorded from a total of 6 bird species, belonging to 4 different orders.

Struthioniformes: 1; Strigiformes: 1; Passeriformes: 2; Accipitriformes: 2

Hosts:

• - Common ostrich – Struthio camelus [A: (60); NA: (30)]

• - Eurasian eagle-owl – Bubo bubo [A: (60)]

• - Great reed warbler – Acrocephalus arundinaceus [F: (14)]

• - House sparrow – Passer domesticus [F: (201) Rhipicephalus sanguineus sensu stricto (based on 99.7% 12S rRNA gene sequence identity between AY559843 and MH630345).

• - Northern goshawk – Accipiter gentilis [A: (60)]

• - Short-toed snake eagle – Circaetus gallicus [A: (60)]

Distribution of reported cases: Romania (14), France (201), Portugal (60), United Kingdom (30).

Rhipicephalus turanicus

Overview: The taxonomic status of Rhipicephalus turanicus is still under discussion (8). According to literature data, this tick species has been found on three bird species (all birds of prey) in Portugal and in Bulgaria. However, the Portuguese findings (85) were referred to in a later article by the same author (60) as R. sanguineus.

Accipitriformes: 2; Strigiformes: 1

Hosts:

• - Common buzzard – Buteo buteo [F: (85)] [In a later article (60), the same author referred to this finding as R. sanguineus]

• - Eurasian eagle-owl – Bubo bubo [M: (153); F: (153)]

• - Steppe eagle – Aquila nipalensis [F: (85)] [In a later article (60), the same author referred to this finding as R. sanguineus]

Distribution of reported cases: Portugal (85), Bulgaria (153).

Conclusions

In this review data on the ixodid tick infestation of birds were collected from nearly 200 papers published since 1952. In this period, 37 hard tick species (17 from Prostriata and 20 from Metastriata) were reported from 16 orders of avian hosts in Europe. These include endophilic tick species that are ornithophilic (I. arboricola, I. caledonicus) or those that prefer mammalian hosts, such as rodents (I. acuminatus), carnivores, or insectivores (I. canisuga, I. hexagonus). Some of the tick species have a clear host preference for sea birds and predominate in Western-Northern Europe (I. rothschildi, I. unicavatus, I. uriae). A significant number of ixodid species are rarely found in association with birds, as exemplified by D. reticulatus, D. marginatus, H. erinacei, H. parva, H. sulcata, R. annulatus, R. turanicus, R. sanguineus, and H. aegyptium. The latter is a tortoise-associated tick species, whereas R. turanicus and R. sanguineus are mostly reported from birds of prey, most likely transferring from rodent and other prey items to these birds. In addition, exotic (non-indigenous) tick species transported by birds from Africa to Europe include H. truncatum, A. lepidum, A. marmoreum, A. nuttalli and A. variegatum.

The primary aim of this checklist is to provide a comprehensive reference source (baseline data) for future studies, particularly in the context of discovering new tick-host associations after comparison with already published data. Nevertheless, these data also allow a first-hand analysis of general trends regarding how and which developmental stage of ticks tend to infest avian hosts in general. Therefore, five tick species that were frequently reported from birds and show a broad geographical distribution on this continent were selected for statistical comparison (I. arboricola, I. frontalis, I. ricinus, H. concinna and H. marginatum) (Supplementary Figure 1; Supplementary Table 1).

Considering developmental stages, larvae and nymphs predominate on birds in the case of I. ricinus, H. concinna, and H. marginatum (n = 1,667 vs. 37 adults). This is not the case for ornithophilic tick species (I. arboricola and I. frontalis) from which adult ticks were collected significantly more frequently (n = 301 vs. 115 adults) (P < 0.0001). Interestingly, males of generalist tick species occurred significantly more frequently on birds (n = 7 vs. 30 females) than males of ornithophilic tick species (n = 5 vs. 110 females) (P = 0.009).

Based on data that did not specify the tick developmental stage, the tick species I. arboricola was significantly (P < 0.0001) more frequently collected from bird species that typically feed above the ground (n = 134) than from those feeding at the ground level (n = 60). This was not the case for I. ricinus (679 vs. 1,064), I. frontalis (121 vs. 216) and H. concinna (36 vs. 63) (Supplementary Table 1). This general tendency on a continental level is in contrast to what was reported from the southern part of Central Europe where H. concinna is mostly associated with bird species typically feeding above the ground level (65, 127). The chances for finding H. marginatum on bird species characterized by either ground-feeding or arboreal feeding were more equilibrated (82 vs. 84). This is in line with its active host seeking (hunting) strategy (8).

In addition, considering data from references that included the number of tick developmental stages collected from various bird species, comparison of tick infestations according to five habitat categories of avian hosts (Supplementary Table 1) did not reveal statistically significant associations (data not shown). However, by assigning bird species into two groups (i.e., typically feeding from the ground vs. rarely occurring at the soil level: Supplementary Table 1), the following differences were found: Larvae, nymphs and females of I. arboricola significantly more frequently occur on bird species feeding above the ground (P < 0.0001-0.045), than corresponding stages of I. frontalis, I. ricinus and H. concinna. Females of I. ricinus are more likely to infest ground-feeding than arboreal bird species (18 vs. 4) compared to nymphs of this species (507 vs. 336) (P = 0.046). Interestingly, females of H. marginatum significantly more frequently associate with bird species feeding above the ground level than with those collecting food on the soil surface (4 vs. 1) compared to those of I. ricinus (4 vs. 18) (P = 0.017) and I. frontalis (27 vs. 61) (P = 0.041).

The ecology of bird-infesting tick species (8) is also illustrated here according to avian orders (Supplementary Figure 1), taking into account the ecology (habitat type) and activity (circadian rhythm and feeding level) of most bird species that belong to a certain order (203) (though this has limitations due to difficulties in assigning general traits at this taxonomic level). In this context, I. arboricola was only reported from species of avian orders whose members typically use forested habitats and (with the exception of Columbiformes) typically feed above the ground (Supplementary Figure 1). This tick species was also reported from a high number of nocturnal bird species (Strigiformes), although (as an endophilic tick, with preference for tree holes) it is known to detach from diurnal passerine hosts during the night (50). Except for two bird species from two orders (Falconiformes, Coraciiformes), I. frontalis was always reported from avian orders whose members typically (also) feed from the ground level, in both open and forested habitats (Supplementary Figure 1). This is in line with previous data on I. frontalis from Central Europe (65). By contrast, I. ricinus was reported from 13 out of 16 avian orders, including several examples with preference for forest or open habitats, and which typically feed at the ground level or higher (Supplementary Figure 1). Similarly, the majority of avian hosts that were reported to be infested with H. concinna represent orders that share both open habitat- and forest-dwelling bird species, as well as ground level and arboreal feeders (Supplementary Figure 1). Last but not least, H. marginatum was almost exclusively reported from bird species that belong to orders including a significant number of forest dweller bird species and was also collected from a high number of bird species from orders with nocturnal activity (both Caprimulgiformes and Strigiformes: Supplementary Figure 1). This is in agreement with the reported evening activity and occurrence of this tick species in forested habitats (204).

Funding

This study was funded by Project no. TKP2020-NKA-01 implemented with the support provided from the National Research, Development and Innovation Fund of Hungary, financed under the Tématerületi Kiválósági Program 2020 (2020-4.1.1-TKP2020) funding scheme. SH and AS were supported by OTKA K-132794 of the National Research, Development and Innovation Office. The Project is supported by the European Union and co-financed by the European Social Fund (grant agreement no. EFOP-3.6.3-VEKOP-16-2017-00005, project title: Strengthening the scientific replacement by supporting the academic workshops and programs of students, developing a mentoring process).

Publisher's note

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