Aves: auks - birds of the family Alcidae, namely: Alca torda Linnaeus (razorbill), Cepphus grylle Linnaeus (black guillemot), Fratercula arctica Linnaeus (common puffin), Uria aalge (Pontoppidan) (common guillemot), Uria lomvia Linnaeus (Brünnich’s guillemot) (Filippova 1977).

Occasional hosts include Fratercula cirrhata (Pallas) (tufted puffin) and also various species of gulls and kittiwakes (Laridae): Rissa brevirostris (Bruch) (red-legged kittiwake), R. tridactyla (Linnaeus) (black-legged kittiwake) (Karpovich 1970; Dietrich et al. 2012), as well as fulmars (Procellariidae) – Fulmarus glacialis Linnaeus (northern fulmar) and cormorants (Phalacrocoracidae) – Phalacrocorax capillatus (Temminck and Schlegel) (Japanese cormorant), Urile pelagicus (Pallas) (pelagic cormorant), Urile urile (Gmelin) (red-faced cormorant) (Lvov et al. 1972b, 1975; Filippova 1977; Dietrich et al. 2012; Duron et al. 2014). A single atypical case of parasitism on Motacilla alba Linnaeus (white wagtail) (Motacillidae) was also reported (Karpovich 1970).

(Fig. 1). Murmansk seacoast (Russia): islands and seashores of the White Sea and also the Barents Sea (Karpovich 1971), namely: the Kuvshin Island and Kharlov Island (Karpovich 1970), Podpakhta Bay (Bekleshova et al. 1970), Dvorovaya Bay (Flint and Kostryko 1967), Seven Islands Reserve (Belopolskaya 1952; Karpovich 1973). The Far East (Russia): Mosolova Bay (the northern coast of the Strait of Tartary) (Savitskaya 1975), Bering Island (Lvov et al. 1975; Dietrich et al. 2012), Ptichy Island and Starichkov Island (Dietrich et al. 2012), Iony Island (Lvov et al. 1975), Kuril Islands (Lvov et al. 1975; Dietrich et al. 2012), Tyuleniy Island (Karpovich 1971; Lvov et al. 1972a; Filippova 1977), Sakhalin (Lvov et al. 1972a), Commander Islands (Dietrich et al. 2012).

Figure 1. 

Map of Russia and neighboring countries showing the locations where Ixodes uriae was reported.

Ixodes uriae is the only representative of the subgenus Ceratixodes in the tick fauna of Russia and the northern hemisphere in general. As a nidicolous parasite of seabirds living in colonies, it is a species with a circumpolar distribution, occurring on oceanic coasts and islands of both the northern and southern hemispheres, from the polar regions to the subtropical zone (Wilson 1967; Filippova 1977).

In the northern hemisphere, this tick species is strongly associated with seabirds of the family Alcidae. The high degree of nest conservativity of these birds contributes to supporting a considerable number of ticks in bird colonies, which use the same places for many years (Karpovich 1971). The occasional hosts of I. uriae usually become involved in its life cycle in mixed bird colonies, where nests of typical and atypical hosts are located very close to each other (Violovich 1962b). In absence of auks, it may also use, for example, cormorants as exclusive hosts, as reported on the Kuril Islands (Lvov et al. 1975; Dietrich et al. 2012). In the southern hemisphere it was noted that penguins (Spheniscidae) are more typical hosts; this can be explained by similarities in the habits of these birds to those of puffins and guillemots.

There were noted rare records of adults from Carnivora: Mustelidae, and nymphs from Rodentia: Muridae (Eley 1977; Jaenson and Jensen 2007; Baggs et al. 2011; Guglielmone et al. 2020) and even humans (Karpovich 1971; Keirans and Lacombe 1998; Martyn 1998; Smith et al. 2006; Jaenson and Jensen 2007).

Mammalia: Myotis blythii Tomes (lesser mouse-eared bat), Miniopterus schreibersii (Kuhl) (common bent-wing bat), Nyctalus leisleri (Kuhl) (lesser noctule) (Filippova 1977).

(Fig. 2). Russia: Krasnodar Krai – outskirts of Sochi (Emchuk 1955; Filippova 1972). Ukraine: eastern Carpathians – outskirts of Solotvyn and Rakhiv (Emchuk 1955; Filippova 1972). Azerbaijan: outskirts of Şahbuz and Hadrut (Emchuk 1955; Filippova 1972).

Figure 2. 

Map of Russia and neighboring countries showing the locations where Ixodes simplex was reported.

Ixodes simplex is a tick species specialized for bats as hosts (Filippova 1977). This species is mainly monoxenous and can be found usually on the common bent-wing bat although some other species of the Chiroptera may also act as hosts, especially which share colonies with its main host (Beaucournu 1967). Some rare cases of human infestation are also recorded (Okino et al. 2010; Péter et al. 2021).

Mammalia: Eptesicus serotinus (Schreber) (serotine bat), Myotis blythii (lesser mouse-eared bat) (Filippova 1977), pond bat Myotis dasycneme (Boie) (Starikov et al. 2017b), Daubenton’s bat Myotis daubentonii (Kuhl) (Orlova et al. 2011), Myotis myotis (Borkhausen) (greater mouse-eared bat) (Filippova 1977), Myotis mystacinus (Kuhl) (whiskered bat) (Bobkova 2003), Nyctalus noctula (Schreber) (common noctule), Pipistrellus pipistrellus (Schreber) (common pipistrelle), Rhinolophus ferrumequinum (Schreber) (greater horseshoe bat), Rhinolophus hipposideros (Bechstein) (lesser horseshoe bat), Rhinolophus mehelyi Matschie (Mehely’s horseshoe bat) (Filippova 1977).

(Fig. 3). Russia: Udmurtia (Orlova et al. 2011), Voronezh Oblast (Usmsnskyi pine forest), (Khitsova and Sherstyanykh 2014), Novosibirsk Oblast (resort Lake Karachi) (Fedorov 2016), Khanty-Mansi Autonomous Okrug (outskirts of the urban locality Mortka) (Starikov et al. 2017b), Krasnodar Krai (Sochi National Park) (Romashin 2021), Stavropol Krai (Tsapko 2019). Ukraine: Ivano-Frankivsk Oblast, Chernivtsi Oblast, Ternopil Oblast, Zakarpattia Oblast, Crimea (Bobkova 2003). Moldova: Codru Reserve (Dniester-Prut interfluve) (Uspenskaya 1987). Georgia: Abkhazia (Kerbabaev 2011). Azerbaijan: Shusha, Hadrut (Filippova 1972). Armenia: Meghri (Ogandzhanyan 1949). Kyrgyzstan: Chüy Valley (Fedorova 2012a). Turkmenistan: rural localities Ahcha-Kuima and Mollagara (Dubinin and Bregetova 1952). Tajikistan: northern spurs of the Zarafshan Range (Filippova 1972).

Figure 3. 

Map of Russia and neighboring countries showing the locations where Ixodes vespertilionis was reported.

Ixodes vespertilionis Koch is a species of ixodid ticks associated with bats as typical hosts (Filippova 1977), mostly from the families Rhinolophidae and Vespertilionidae. Usually, I. vespertilionis can be found in caves inhabited by bats. Occasional findings in Central Russia and Siberia are considered to result from accidental transportation.

Mammalia: Apodemus flavicollis Melchior (yellow-necked field mouse), Chionomys gud Satunin (Caucasian snow vole), Chionomys nivalis (Martins) (European snow vole), Microtus daghestanicus (Shidlovsky) (Daghestan pine vole), Nothocricetulus migratorius (Pallas) (grey dwarf hamster), Sorex raddei Satunin (Radde’s shrew) (Filippova and Panova 1989; Filippova and Stekol’nikov 2007).

(Fig. 4). Russia: Dagestan – the valley of the Akhtychay River which is the right tributary of the Samur River near the confluence of these rivers, ~ 1000 m a.s.l. and at the same location near rural locality Khnov, ~ 1700 m a.s.l.; the valley of the Avar Koysu River, ~ 1000 m a.s.l. (Filippova and Panova 1989); Kabardino-Balkaria, Bezengi gorge – 1550–2500 m a.s.l. and Karachay-Cherkessia – 1900–2200 m a.s.l. (Filippova and Stekol’nikov 2007). Georgia: Mtskheta-Mtianeti region, Kazbegi Municipality, outskirts of the hamlet Suatisi, 2200 m a.s.l. (Filippova and Panova 1989).

Figure 4. 

Map of Russia and neighboring countries showing the locations where Ixodes ghilarovi was reported.

Ixodes ghilarovi is the second representative of the subgenus Filippoviella in the Palearctic tick fauna together with I. trianguliceps but known at the current moment exclusively from several locations of the Caucasus (Filippova and Panova 1988). The species was found only in rocky biotopes on the slopes containing xerophilous herbaceous-shrub vegetation consisting of many endemics of Southern Dagestan (Filippova and Panova 1989).

Further investigations of this poorly studied tick species are of undoubted interest. Ixodes ghilarovi has certain common structural features with the African species I. alluaudi, for example the presence of auriculae, especially visible in nymphs of both species (Filippova 2010); molecular analysis is also necessary to obtain more data on interspecific connections of these ticks and inside the subgenus in general. The host-parasite relations of I. ghilarovi and its distribution and habitats are probably wider than it is known today. The seasonality of I. ghilarovi and its role as a vector of tick-borne infections remain unknown.

The type specimens of I. ghilarovi are deposited at the Zoological Institute of the Russian Academy of Sciences and include the holotype: nymph; Russia, 25, Daghestan, Samur Mt. Range, near Akhty Village, River Akhtychay valley, ~ 1000 m a. s. l., Chionomys gud, Sat., 24.5.1980, coll. I.V. Panova; FBM 610a, 610b and the paratypes: 4 nymphs; FBM I610a, I610b. Description – Filippova and Panova 1989: 419–421 (female, larva; male unknown) (Filippova 2008).

Mammalia: Alexandromys oeconomus (Pallas) (tundra vole), Apodemus agrarius (Pallas) (striped field mouse) (dominates as the host in the Udel’ny forest park in St. Petersburg, according to Tretyakov (2009), Apodemus flavicollis (yellow-necked field mouse), Apodemus sylvaticus (Linnaeus) (wood mouse), Apodemus uralensis Pallas (Ural field mouse), Arvicola amphibius (Linnaeus) (European water vole), Chionomys gud (Caucasian snow vole), Chionomys nivalis (European snow vole), Craseomys rufocanus (Sundevall) (grey red-backed vole), Cricetus cricetus (Linnaeus) (European hamster), Crocidura leucodon (Hermann) (bicolored shrew), Crocidura suaveolens (Pallas) (lesser white-toothed shrew), Eutamias sibiricus (Laxmann) (Siberian chipmunk), Lasiopodomys gregalis (Pallas) (narrow-headed vole), Lepus europaeus Pallas (European hare), Lepus timidus Linnaeus (mountain hare), Micromys minutus (Pallas) (harvest mouse), Microtus agrestris (Linnaeus) (short-tailed field vole), Microtus arvalis (Pallas) (common vole), Microtus majori (Thomas) (Major’s pine vole), Microtus socialis (Pallas) (social vole), Microtus subterraneus (de Selys-Longchamps) (European pine vole), Mus musculus Linnaeus (house mouse), Mustela nivalis Linnaeus (least weasel), Myodes glareolus (Schreber) (bank vole), Myodes rutilus (Pallas) (northern red-backed vole), Myopus schisticolor (Lilljeborg) (wood lemming), Neomys anomalus Cabrera (Mediterranean water shrew) (Filippova 1977), Neomys fodiens (Pennant) (Eurasian water shrew) (Lutta 1968), Nyctalus noctula (common noctule), Ochotona alpina (Pallas) (alpine pika), Prometheomys schaposchnikowi Satunin (long-clawed mole vole), Rattus norvegicus (Berkenhout) (brown rat), Sciurus vulgaris Linnaeus (red squirrel), Sicista betulina Pallas (northern birch mouse), Sorex araneus Linnaeus (common shrew), Sorex caecutiens Laxmann (Laxmann’s shrew), Sorex daphaenodon Thomas (Siberian large-toothed shrew) (Filippova 1977), Sorex isodon Turov (taiga shrew) (Sapegina 1980), Sorex minutus Linnaeus (Eurasian pygmy shrew), Sorex minutissimus Zimmermann (Eurasian least shrew) (Filippova 1977), Sorex roboratus Hollister (flat-skulled shrew) (Shtilmark 1963; Sapegina 1980), Spermophilus suslicus (Güldenstädt) (speckled ground squirrel), Vulpes vulpes (Linnaeus) (red fox) (Filippova 1977).

Aves: Anthus trivialis (Linnaeus) (tree pipit), Carduelis carduelis (Linnaeus) (European goldfinch), Dendrocopos major (Linnaeus) (great spotted woodpecker), Emberiza citronella Linnaeus (yellowhammer), Nucifraga caryocatactes (Linnaeus) (Eurasian nutcracker), Strix uralensis Pallas (Ural owl), Turdus viscivorus Linnaeus (mistle thrush) (Filippova 1977).

Reptilia: Zootoca vivipara (viviparous lizard) (Lichtenstein) (Filippova 1977).

(Fig. 5). Russia: North Karelia – Cape Kartesh (Stanyukovich and Fedorov 2022); Karelia (Lutta 1968) including the village Malaya Gomselga (southern Karelia) (Bespyatova and Bugmyrin 2015; Bespyatova et al. 2019), St. Petersburg (Tretyakov 2009), Leningrad Oblast (Sukhomlinova 1977), Novgorod Oblast (Grigoryeva and Tretyakov 1998), Pskov Oblast – the village Gogolevo (own data, unpublished), Kaliningrad Oblast, the Vistula Spit (own data, unpublished); Tver Oblast (Schipanov and Makhanko 2018), Tula Oblast (Kozlova et al. 2014), Perm Oblast (Korenberg et al. 2015), Eastern Upper Volga (Egorov et al. 2016), Krasnodar Krai and the Caucasus (Shatas 1957; Filippova and Stekol’nikov 2007), Kurgan Oblast (Starikov and Starikova 2021), Tyumen Oblast (Bragina et al. 2013), Omsk Oblast (Rar et al. 2014, 2020), Kemerovo Oblast (Kovalevsky et al. 2018), Western Sayan (Shtilmark 1963), Eastern Sayan (Schluger 1961), Khamar-Daban ridge (Vershinina 1988). Belarus (Arzamasov 1963). Ukraine: Crimea (Filippova 2010), Polesia (Podobivskyi and Fedonyuk 2017). Moldova: north and central Moldova (Uspenskaya et al. 2006). Georgia: the village Bakuriani and the Roki Tunnel (Djaparidze 1960). Armenia: the whole territory (Ogandzhanyan 1960). Azerbaijan: the south of the country (Ogandzhanyan 1960).

Figure 5. 

Map of Russia and neighboring countries showing the locations where Ixodes trianguliceps was reported.

Ixodes trianguliceps Birula has a wide geographical distribution in the Palaearctic region, occurring from the coast of Lake Baikal to Western Europe (Filippova 2010; Estrada-Peña et al. 2018). In the north it reaches northern Karelia and the Scandinavian Peninsula (Fedorov and Leonovich 2021). Also, an isolated southern population of this species was found in the Crimean Peninsula (Filippova 2010) although in other parts of Ukraine it is present in forest zones, such as Polesia (Podobivskyi and Fedonyuk 2017).

The population that was supposed to be isolated in the mountain systems of the Caucasus (Filippova 2010) now seems to be more expanded, as proved by the recent finding in Turkey (Bolu and Kars province, the north of Turkey) (Keskin and Selçuk 2021). The Kars province is located near the border with Georgia, where this species was known before (Djaparidze 1960) and, therefore, the ticks reported from there are probably part of the same Caucasian population.

The map of findings of this tick species in Russia clearly illustrates that it lives in a broad range of forest biotopes throughout a vast territory including the zonal and mountain deciduous and mixed forest of the European type and forests of southern and middle-taiga types. Along the southern border of the largest part of the range in Russia, I. trianguliceps occurs in the forest-steppe zone, populating shrubby and forested biotopes. This distinctly correlates with the main habitats of shrews and rodents, because the presence of these small mammals together with well-developed soil litter, plays an important role in the abundance of ticks in the landscape, as it is known that shrews of the genus Sorex are the most preferable host for larvae (Randolph 1975).

Interestingly, I. triangulipeps was also reported from two bat species (Myotis myotis in Poland (Siuda et al. 2009) and Nyctalus noctula in Russia, as well as several bird species and one reptile species (Filippova 1977). These animals are non-typical and occasional hosts for this tick species. The single cases of parasitism on these host species can be a clear indication that I. trianguliceps is predominantly an exophilic species, because it is unlikely that ticks could contact bats and birds in a burrow. Findings of this tick species in micropores of burrow tunnels in Belarus in winter (Arzamasov 1963) demonstrate only the ability of its larvae to remain active even during winter.

Phylogenetic trees inferred from the concatenated nucleotide sequences of 10 protein-coding genes of the mitochondrial genome of I. trianguliceps, together with consideration of its morphology, justified to establish the new subgenus Filippoviella and include there I. trianguliceps together with aforementioned I. ghilarovi (Apanaskevich et al. 2024) both of which used to belong to the subgenus Exopalpiger.

Mammalia: Alexandromys oeconomus (tundra vole), Apodemus agrarius (striped field mouse), Apodemus flavicollis (yellow-necked field mouse), Apodemus sylvaticus (wood mouse), Arvicola amphibius (European water vole), Cricetus cricetus (European hamster), Craseomys rufocanus (grey red-backed vole), Erinaceus europaeus Linnaeus (European hedgehog), Eutamias sibiricus (Siberian chipmunk), Lasiopodomys gregalis (narrow-headed vole), Lepus timidus (mountain hare), Micromys minutus (Eurasian harvest mouse), Microtus arvalis (common vole), Microtus agrestis (Linnaeus) (short-tailed field vole), Mus musculus (house mouse), Mustela nivalis (least weasel), Mustela sibirica Pallas (Siberian weasel), Myodes glareolus (bank vole), Myodes rutilus (northern red-backed vole), Myopus schisticolor (wood lemming), Neomys fodiens (Eurasian water shrew), Nothocricetulus migratorius (grey dwarf hamster), Ondatra zibethicus (Linnaeus) (muskrat), Rattus rattus (Linnaeus) (black rat), Sicista betulina (northern birch mouse), Sorex araneus (common shrew), Sorex caecutiens (Laxmann’s shrew), Sorex daphaenodon (Siberian large-toothed shrew), Sorex isodon (taiga shrew), Sorex minutus (Eurasian pygmy shrew), Sorex roboratus (flat-skulled shrew), Talpa europaea Linnaeus (European mole), Vulpes vulpes (red fox) (Filippova 1977).

Aves: Anas crecca Linnaeus (Eurasian teal) (Adamovich 1968), Gallinula chloropus (Linnaeus) (common moorhen) (Filippova 1977), Motacilla alba (white wagtail), Turdus merula Linnaeus (common blackbird) (Adamovich 1968).

(Fig. 6). Russia: Arkhangelsk Oblast (Olenev 1931a), Karelia (Lutta 1976), Saint-Petersburg (Tretyakov 2009), Leningrad Oblast (Sukhomlinova 1977), Vologda Oblast, Tver Oblast (Filippova 1977; Belova et al. 2008), Moscow Oblast (Mosolov 1961), the whole territory of the Upper-Volga (Egorov et al. 2016), Samara Oblast (Kirillova and Kirillov 2008a), Bryansk Oblast (Adamovich 1968), Voronezh Oblast, Nyzhny Novgorod Oblast (Solovyov 1966), Chuvash Republic (Petrov et al. 1967), Krasnodar Krai (Kalita and Pelipeychenko 1957; Shevchenko et al. 1960), Kabardino-Balkaria (Bittirova et al. 2019), Dagestan (Aliev et al. 2012), Perm Krai, Chelyabinsk Oblast (Filippova 1958a), Ekaterinburg (Chernousova and Tolkachyov 2009), Omsk Oblast (Znamenskiy district and Bolsheukov district) (Sabitova et al. 2023), Khanty-Mansiysk (Popov 1967), Surgut (Petukhov et al. 2018), Novosibirsk Oblast (Novosibirsk and Toguchinsky District) (Mal’kova and Bogdanov 2004), Tyumen Oblast – Nyzhnevartovsk (Starikov et al. 2017a), Kurgan Oblast (Starikov and Starikova 2021), Salekhard (Starikov et al. 2017a), Tomsk Oblast (Chainsky District) (Mal’kova and Bogdanov 2004), Kemerovo Oblast, Altai Krai, Altai Republic (Bogdanov and Yakimenko 2016), Krasnoyarsk Krai – Podkamennaya Tunguska River and the rural locality Bolshoy Kemchug (Voltsyt 1997). Ukraine: Volyn Polesie (Adamovich 1968), outskirts of Kyiv (Akimov and Nebogatkin 2013), Cherkassy Oblast (Nikitchenko 2011), the North-Western seacoast of the Black Sea (Rusev 2009). Belarus: throughout the whole territory (Subbotina and Osmolovsky 2022). Moldova: reedbeds of the lower reaches of the Prut River (Uspenskaya et al. 1984). Kazakhstan: Jambyl Region (Galuzo 1950), Jetisu Region – outskirts of Taldykorgan and Jarkent, Almaty Region – outskirts of Sarkand and Almaty (Golov 1933; Sorokoumov 1937; Ushakova and Fedosenko 1972; Ushakova et al. 1976). Kyrgyastan: outskirts of Bishkek, Tokmak Reserve (Grebenyuk 1966), Chuy Valley (Kharadov et al. 2013).

Figure 6. 

Map of Russia and neighboring countries showing the locations where Ixodes apronophorus was reported.

Ixodes apronophorus has a wide distribution in the Northern Palearctic from the Atlantic coast to Eastern Siberia. Its geographical range generally coincides with the distribution of the water vole, its most frequent host, as both the tick and its common host prefer swampy and humid places for living, especially near water bodies.

Aves: Alectoris chukar (Gray) (chukar partridge), Anthus campestris (Linnaeus) (tawny pipit), Athene noctua (Scopoli) (little owl), Chroicocephalus ridibundus (Linnaeus) (black-headed gull), Coccothraustes coccothraustes (Linnaeus) (hawfinch), Coloeus monedula (Linnaeus) (western jackdaw), Curruca communis (Latham) (common whitethroat), Emberiza bruniceps Brandt (red-headed bunting), Galerida cristata (Linnaeus) (crested lark), Lullula arborea (Linnaeus) (woodlark), Luscinia svecica (Linnaeus) (bluethroat), Melanocorypha bimaculata (Ménétrés) (bimaculated lark), Monticola solitarius (Linnaeus) (blue rock thrush), Oenanthe sp. (wheatear), Passer domesticus (Linnaeus) (house sparrow), Perdix perdix (grey partridge), Petronia petronia (Linnaeus) (rock sparrow), Phoenicurus erythronotus (Eversmann) (Eversmann’s redstart), Phylloscopus griseolus (Blyth) (sulphur-bellied warbler), Pica pica (Linnaeus) (Eurasian magpie), Sitta tephronota Sharpe (Eastern rock nuthatch), Turdus merula (common blackbird) (Filippova 1977).

Mammalia: Crocidura leucodon (bicolored shrew), Meriones persicus (Blanford) (Persian jird), Mus musculus (house mouse), Nesokia indica (Gray) (short-tailed bandicoot rat), grey dwarf hamster Nothocricetulus migratorius (Pallas), Rattus pyctoris (Hodgson) (Turkestan rat), Rhinolophus mehelyi (Mehely’s horseshoe bat) (Filippova 1977).

(Fig. 7). Russia: Dagestan and North Osetia-Alania (Shatas 1957; Filippova 1977). Ukraine: Crimean Peninsula, in particular the Tarkhankut Peninsula and the Kara Dag (Filippova 1974). Georgia: the Shiraki Plain and the Vashlovani Nature Reserve (Djaparidze 1950, 1960). Armenia: Vayots Dzor Province – the rural locality Herher (Ogandzhanyan 1959). Azerbaijan: Karabakh Plateau – Lachin District and Hadrut District, Adzhynokhur Steppe (Ogandzhanyan 1959; Filippova 1977). Kazakhstan: Dzungarian Alatau (Ushakova et al. 1976) and Trans-Ili Alatau (Filippova 1977). Kyrgyzstan: Terskey Ala-too Range (Filippova 1974). Turkmenistan: the Kopet Dagh – the valley of the Chandyr River, Magtymguly, Gökdepe District, outskirts of Ashgabad, Köytendag Range, Bayramaly (Kerbabaev 1960; Kochkareva et al. 1971; Berdyev 1973; Scherbinina 1973). Uzbekistan: Termez (Filippova 1977). Tajikistan: Hisar Range, Varzob gorge, outskirts of Dushanbe (Filippova 1977).

Figure 7. 

Map of Russia and neighboring countries showing the locations where Ixodes eldaricus was reported.

Ixodes eldaricus is a little studied endophilic tick species which is mainly a parasite of ground-feeding birds although nymphs and larvae, besides birds, were also found on small mammals – rodents and shrews. It usually inhabits deciduous mountain forests and shrub thickets in mountain river valleys. The vertical distribution range of its occurrence varies from 300 (Ashgabat) to 1800 m (Terskey Ala-too Range and Hisar Range) a. s. l. (Filippova 1977).

Briefly described by a female from the east of Georgia (type locality: the Shiraki Plain), I. eldaricus was later found in Armenia and Azerbaijan, and the male, nymph descriptions were based on the material from Azerbaijan (Ogandzhanyan 1959). The holotype female described from the grey partridge is stored at the Institute of Zoology of Ilia State University. The above findings from post-Soviet territories are known from the Crimea, as well as the Causasus and Central Asia. The majority of samples are stored at the collection of the Zoological Institute of the Russian Academy of Sciences.

Additionally, it is important to note that in Crimea this tick species is considered disappearing (Nebogatkin 1998) due to anthropogenic pressure followed by destruction of its habitats and decline in its host populations (Uspensky 2021).

Mammalia: Apodemus sylvaticus (wood mouse), Canis familiaris Linnaeus (dog), Ovis aries Linnaeus (sheep) (Filippova 1977).

(Fig. 8). Kyrgyzstan: the Tien Shan – northern and eastern slopes of the Terskey Ala-too range (gorges Ulken-Kokpak and Chon-Dzhargylchak) (Filippova 1969).

Figure 8. 

Map of Russia and neighboring countries showing the locations where Ixodes kashmiricus was reported.

Ixodes kashmiricus is a tick species with a disjunctive relict range limited by the Tien Shan in Kyrgyzstan as well as India (Filippova 1977) and Pakistan (Numan et al. 2022). In Kyrgyzstan the tick was found mainly in the mid-altitude vertical zone of the mountains at the lower border of the forest at the altitude of 2000 and 2500 m a. s. l. Cases of parasitism on humans have been recorded (Hoogstraal 1970).

Phylogenetic analysis of mitochondrial and nuclear genes showed that I. kashmiricus belongs to the I. ricinus group (Kovalev et al. 2018) and clusters with such members of the I. ricinus group as I. apronophorus and I. kazakstani (Numan et al. 2022).

The type specimens are stored at the Zoological Institute of the Russian Academy of Sciences and include the lectotype - female; [India], Kashmir, Vardvan Maru River, northern tributary of Chinab River, 10–13. V.1910, coll. S.P. Trubetskoi; AL I533, as well as the paralectotype - male; AL 533a. Ixodes kashmiricus (see: Filippova 1969: 677). Description – Filippova 1977: 292–296 (female, male, nymph, larva) (Filippova 2008). Originally the tick was named I. persulcatus kaschmiricus (lapsus).

Mammalia: Apodemus sylvaticus (wood mouse) (Filippova 1977), Canis familiaris (domestic dog) (Kovalev et al. 2018), Dryomys nitedula (Pallas) (forest dormouse), Lepus tolai Pallas (tolai hare), Mus musculus (house mouse), Nothocricetulus migratorius (grey dwarf hamster) (Filippova 1977).

Aves: Phasianus colchicus Linnaeus (common pheasant) (Filippova 1977).

(Fig. 9). Kazakhstan: Betpak-Dala – the valley of the Chu River (Ushakova 1961), Tian Shan – the valley of the Ili River (Ushakova 1958; Kovalev et al. 2018), outskirts of Jarkent (Olenev and Sorokoumov 1934; Pomerantsev 1950). Kyrgyzstan: the Issyk-Kul basin (Filippova 1958b; Kovalev et al. 2018), the valley of the Talas River (Olenev and Sorokoumov 1934; Pomerantsev 1950; Grebenyuk 1966; Lyashko 1973; Kovalev et al. 2018).

Figure 9. 

Map of Russia and neighboring countries showing the locations where Ixodes kazakstani was reported.

Ixodes kazakstani is a tick species with a disjunctive relict range limited by Southeastern Kazakhstan and neighboring territories of Kyrgyzstan (Filippova 1977). The patchy arrangement of its range can be explained, above all, by associations of this tick mainly with the animals dwelling in tugai forests which also create humidity conditions in the soil suitable for this tick species (Filippova 1958b). Also, there are some cases of parasitism on livestock and humans (Lyashko 1973; Filippova 1977). On livestock it was found in few numbers among mass parasitism of other tick species.

Phylogenetic analysis of mitochondrial and nuclear genes showed that I. kazakstani belongs to the I. ricinus group (Kovalev et al. 2018) and clusters with such members of the I. ricinus group as I. apronophorus and I. kashmiricus (Numan et al. 2022). Ixodes kazakstani can presumably exemplify links between Nearctic and Palearctic species, so further studies of genetic sequences of I. kazakstani are necessary to understand better evolutionary connections between more tick species in the I. ricinus group.

The type specimens are stored at the Zoological Institute of the Russian Academy of Sciences and include the holotype: female; Kazakhstan, Jarkent, collected from human dress, 20.VI.1932, coll. Kirin; AL I536. Description - Filippova 1977: 283–290 (female, male, nymph, larva) (Filippova 2008).

Mammalia: Allactaga major (Kerr) (great jerboa), Allocricetulus eversmanni (Brandt) (Eversmann’s hamster), Apodemus sylvaticus (wood mouse), Chionomys nivalis (European snow vole), Cricetus cricetus (European hamster), Dryomys nitedula (forest dormouse), Ellobius talpinus (Pallas) (northern mole vole), Erinaceus europaeus (European hedgehog), Glis glis (Linnaeus) (European edible dormouse), Hemiechinus auratus (Gmelin) (long-eared hedgehog), Lagurus lagurus (Pallas) (steppe lemming), Marmota bobak (Müller) (bobak marmot), Martes martes (Linnaeus) (European pine marten), Microtus arvalis (common vole), Microtus socialis (social vole), Meles meles (Linnaeus) (European badger), Meriones meridianus (Pallas) (midday jird), Mesocricetus brandti (Pallas) (Turkish hamster), Mesocricetus raddei (Nehring) (Ciscaucasian hamster), Mus musculus (house mouse), Mustela eversmanii (Lesson) (steppe polecat), Mustela nivalis (least weasel), Nothocricetulus migratorius (grey dwarf hamster), Pygeretmus pumilio (Kerr) (dwarf fat-tailed jerboa), Rattus rattus (black rat), Spalax microphthalmos Gueldenstaedt (greater blind mole-rat), Spermophilus citellus (Linnaeus) (European ground squirrel), Spermophilus fulvus (Lichtenstein) (yellow ground squirrel), Spermophilus pygmaeus (Pallas) (little ground squirrel), Spermophilus suslicus (speckled ground squirrel), Spermophilus xanthoprymnus (Bennett) (Asia Minor ground squirrel), Stylodipus telum (Lichtenstein) (thick-tailed three-toed jerboa), Vormela peregusna (Güldenstädt) (marbled polecat), Vulpes corsac (Linnaeus) (Corsac fox), Vulpes vulpes (red fox) (Filippova 1977).

(Fig. 10). Russia: Samara Oblast (Kirillova and Kirillov 2008b), Rostov Oblast (Stakheev and Panasyuk 2016), Krasnodar Krai (Popov et al. 2019), Stavropol Krai (Tsapko 2019), Volgograd Oblast, Astrakhan Oblast (Nelzina et al. 1955), Kalmyk Republic (Sandzhiev et al. 2006), Chechnya (Baisarova 2021), Dagestan (Musaev et al. 2019) and North Osetia-Alania (Filippova 1977). Ukraine: Kyiv (Omeri and Moysak 2013), Odesa Oblast (Rusev 2008), Kherson Oblast, Chernivtsi Oblast, Ternopil Oblast, Luhansk Oblast, Donetsk Oblast, the Crimean Peninsula, particularly in the Syvash (Filippova 1958a; Emchuk 1960; Sklyar 1970; Andryushchenko et al. 2005; Evstafiev 2017). Moldova: Bălți Steppe, Bugeac Steppe (Filippova 1977) and Tiraspol (Kravchenko 2014). Georgia: Abkhazia (Shaposhnikova and Sakhno 2012), Imereti (Sukhiashvili et al. 2020), Lagodekhi Nature Reserve (Djaparidze 1960). Armenia: Lori Province –Nalband and the valley of the river Hrazdan (Filippova 1977). Azerbaijan: Talysh (Pomerantsev 1950), the Nakhchivan Autonomous Republic – the Zangezur Mountains (Kadatskaya and Shirova 1963; Filippova 1977). Kazakhstan: West Kazakhstan Region (Pomerantsev 1950; Levit 1957; Filippova 1977), Kyzylorda Region (Loseva 1963), Kostanay Region, Akmola Region (Ushakova 1961, 1962). Turkmenistan: the Kopet Dagh (Kerbabaev 1961).

Figure 10. 

Map of Russia and neighboring countries showing the locations where Ixodes laguri was reported.

Ixodes laguri is a tick species which is mainly a nidicolous parasite of rodents and small and medium carnivores, first of all ground squirrels. It is present usually in zonal and mountainous steppes at the altitude of 1500 m a.s.l. This tick species is less common in desert and semi-desert biotopes (Filippova 1977).

Filippova (1977) states that the tick has four subspecies – I. laguri laguri, I. l. armeniacus, I. l. colchicus and I. l. slovacicus. The differential characters of the female and the male of I. l. slovacicus are based on comparison with characters of the other subspecies in Pomerantsev (1950) but some of them, such as the genital aperture and chaetotaxy of the scutum and the hypostome and the coxa 1, are not characterized precisely enough (Filippova 1977).

According to Filippova (1977), I. laguri laguri can be found in Moldova, Ukraine, Kazakhstan, as well as in the south of Russia; I. l. armeniacus is distributed in the Caucasus – North Osetia-Alania, Dagestan, Georgia, Armenia and Azerbaijan; I. l. colchicus is known from the western spurs of the Greater Caucasus, the now abandoned rural locality Babuk-Aul; I. l. slovacicus was described from the south-east of Slovakia.

The type specimens of I. laguri are deposited at the Zoological Institute of the Russian Academy of Sciences and include I. l. armeniacus: the lectotype, female; Armenia, Nalband, from Mesocricetus brandti Nehr., 9.9.1936; AL I558 and the paralectotype, male; AL I556, description – Filippova 1977: 384 (female, male, nymph; larva unknown). (Filippova 2008), as well as I. l. colchicus: the lectotype, male; Western Caucasus, near Babuk-Aul, Glis glis L., 30.9.1935, coll. V. K. Popov, det. В. Pomerantsev: I. l. colchicus, type; AL I554a; paralectotypes: 2 females; AL I554a, description – Filippova 1977: 384 (female, male; nymph and larva unknown) (Filippova 2008).

Mammalia: Apodemus agrarius (striped field mouse), Craseomys rufocanus (grey red-backed vole), Microtus fortis (Büchner) (reed vole), Myodes rutilus (northern red-backed vole) (Filippova 1977).

(Fig. 11). Russia: Primorsky Krai – the Lake Khasan, the Poyma River, the Partizansky District, outskirts of urban localities Posyet, Kraskino, Slavyanka and cities Vladivostok and Nakhodka, near the village Rechitsa (Filippova 1969; Filippova and Belyaev 1970; Allenov et al. 2015).

Figure 11. 

Map of Russia and neighboring countries showing the locations where Ixodes nipponensis was reported.

Ixodes nipponensis is a tick species found in Russia in the south and south-west of the Primorsky Krai and also in the Korean peninsula and Japan (Filippova 1977). In Russia it was reported mainly from murine rodents, although in the Republic of Korea it was also observed on lizards (Kim et al. 2018) and cattle, goats, dogs, horses, and birds in Japan (Kitaoka and Saito 1967; Yamaguti et al. 1971).

Multiple cases of parasitism on humans have been recorded (Nakatsukase and Hatsushika 1985; Paik et al. 1989; Cho et al. 1995; Chu et al. 1997; Ryu et al. 1998; Ko et al. 2002).

Mammalia: Crocidura suaveolens (lesser white-toothed shrew), Diplomesodon pulchellum (Lichtenstein) (piebald shrew), Meriones libycus Lichtenstein (Libyan jird), Meriones meridianus (midday jird), Meriones persicus (Persian jird), Mustela nivalis (least weasel), Nothocricetulus migratorius (grey dwarf hamster), Rhombomys opimus (Lichtenstein) (great gerbil), Spermophilopsis leptodactylus (Lichtenstein) (long-clawed ground squirrel), Vormela peregusna (marbled polecat) (Filippova 1977).

Reptilia: Gloydius halys (Pallas) (Halys pit viper) (Filippova 1977).

(Fig. 12). Kazakhstan: Mangystau Region – the Mangyshlak Peninsula (Kaluzhenkova et al. 1961) and the Ustyurt Plateau; Kyzylorda Region (Filippova 1958a; Loseva 1963; Maslennikova and Ushakova 1971), Jambyl Region – the Moiynkum Desert (Maslennikova and Ushakova 1971), Almaty Region – the foothills of the Dzungarian Alatau: the Sholak and Katutau mountains, the deserts Taukum and Saryesik-Atyrau (Ushakova 1960; Maslennikova et al. 1964; Ushakova et al. 1976). Turkmenistan: distributed everywhere – the southern Ustyurt, the Octumkumy Desert, the Üňüzaňyrsy and Türkmenbaşy Plateau, the Meshed and Saynaksan Desert, the Karakum Desert (Pomerantsev 1950; Kerbabaev 1961; Kochkareva et al. 1971); Hojagala (Berdyev and Annaev 1997). Uzbekistan: the Pistalitau Ridge and the rural locality Tashrabat (Maslennikova and Ushakova 1971).

Figure 12. 

Map of Russia and neighboring countries showing the locations where Ixodes occultus was reported.

Ixodes occultus is a tick species inhabiting deserts. It is mainly a nidicolous parasite of gerbils and jirds (subfamily Gerbillinae), first of all, the great gerbil, as well as of those small mammals which also use long and deep burrows of great gerbils as shelters (Filippova 1977). Some predators which have strong trophic relationships with gerbils and regularly contact with their colonies act as secondary hosts for this tick species.

The type specimen of I. occultus is deposited at the Zoological Institute of the Russian Academy of Sciences and includes the holotype: male; Turkmenia, Repetek, Rhombomys opimus, 5.10.1937, coll. B.I. Pomerantsev, type; AL I550. Description – Filippova 1977: 365–371 (female, male, nymph, larva) (Filippova 2008).

Aves: Acrocephalus dumetorum Blyth (Blyth’s reed warbler), Acrocephalus schoenobaenus (Linnaeus) (sedge warbler), Anas platyrhynchos Linnaeus (mallard), Anthus trivialis (tree pipit), Calliope calliope (Pallas) (Siberian rubythroat), Carduelis carduelis (European goldfinch), Carpodacus erythrinus (Pallas) (common rosefinch), Chloris chloris (Linnaeus) (European greenfinch) Columba livia Gmelin (rock dove), Corvus cornix Linnaeus (hooded crow), Corvus corone Linnaeus (carrion crow), Coturnix coturnix (Linnaeus) (common quail), Crex crex (Linnaeus) (corn crake), Curruca communis (Latham) (common whitethroat), Curruca curruca (Linnaeus) (lesser whitethroat), Cyanopica cyanus Pallas (azure-winged magpie), Emberiza calandra Linnaeus (corn bunting), Emberiza citrinella Linnaeus (yellowhammer), Emberiza leucocephalos Gmelin (pine bunting), Emberiza spodocephala Pallas (black-faced bunting), Ficedula hypoleuca (Pallas) (European pied flycatcher), Fringilla coelebs Linnaeus (Eurasian chaffinch), Fringilla montifringilla Linnaeus (brambling), Lanius collurio Linnaeus (red-backed shrike), Locustella lanceolata (Temminck) (lanceolated warbler), Luscinia luscinia (Linnaeus) (thrush nightingale), Luscinia svecica (Linnaeus) (bluethroat) Parus major Linnaeus (great tit), Passer montanus (Linnaeus) (Eurasian tree sparrow), Pastor roseus (Linnaeus) (rosy starling), Phoenicurus phoenicurus (Linnaeus) (common redstart), Phylloscopus fuscatus (Blyth) (dusky warbler), Phylloscopus trochiloides (Sundevall) (greenish warbler), Pica pica (Eurasian magpie), Sitta europaea Linnaeus (Eurasian nuthatch), Sturnus vulgaris Linnaeus (common starling), Sylvia borin (garden warbler), Tetrao urogallus (western capercaillie), Tetrastes bonasia (hazel grouse), Turdus iliacus Linnaeus (redwing), Turdus philomelos Brehm (song thrush), Turdus pilaris Linnaeus (fieldfare), Turdus ruficollis Pallas (red-throated thrush), Turdus viscivorus (mistle thrush) (Filippova 1977; Moskvitina et al. 2014).

Mammalia: Alexandromys oeconomus (tundra vole), Apodemus agrarius (striped field mouse), Arvicola amphibius (European water vole), Craseomys rufocanus (grey red-backed vole), Cricetus cricetus (European hamster), Eutamias sibiricus (Siberian chipmunk), Lepus timidus (mountain hare), Microtus agrestis (short-tailed field vole), Microtus arvalis (common vole), Mus musculus (house mouse), Myodes glareolus (bank vole), Myodes rutilus (northern red-backed vole), Neomys fodiens (Eurasian water shrew), Nothocricetulus migratorius (grey dwarf hamster), Ochotona alpina (Alpine pika), Sciurus vulgaris (red squirrel), Sicista betulina (northern birch mouse), Sicista subtilis (Pallas) (southern birch mouse), Sorex araneus (common shrew), Sorex minutus (Eurasian pygmy shrew), Sorex roboratus (flat-skulled shrew), Stenocranius gregalis (Pallas) (narrow-headed vole) (Filippova 1977).

(Fig. 13). Russia: Tomsk Oblast (Kovalev et al. 2015), Novosibirsk Oblast, Altai Republic (Tkachev et al. 2017), Altai Krai, Kemerovo Oblast, Krasnoyarsk Krai, Khakassia, northern spurs of the Western Sayan, Amur Oblast, Khabarovsk, Primorsky Krai – the Sikhote-Alin (Filippova 1969; Sapegina and Ravkin 1969; Filippova and Panova 1998), Russky Island (Nikitin et al. 2021). Kazakhstan: East Kazakhstan Region (Tkachev et al. 2017; Perfilyeva et al. 2020), Abai Region, Jetisu Region (Filippova 1977), Tarbagatai Mountains, Dzungarian Alatau, Küngöy Ala-Too Range (Ushakova et al. 1976; Filippova and Panova 1998). Kyrgyzstan: Küngöy Ala-Too Range (Filippova and Panova 1998), Terskey Ala-too (Fedorova 2017).

Figure 13. 

Map of Russia and neighboring countries showing the locations where Ixodes pavlovskyi was reported.

Ixodes pavlovskyi is a tick species distributed in Western Siberia, the Far East, Eastern Kazakhstan, and Kyrgyzstan (Filippova 1977; Fedorova 2017), as well as in China (Guo et el. 2016) and Japan (Nakao et al. 1992; Guglielmone et al. 2023). It more often prefers birds as hosts, as well as small mammals although some cases of human and livestock infestation are also recorded. Its preferred habitats include usually coniferous and deciduous forests, undergrowth, as well as motley grass (Filippova 1977).

Often it can be found in the same biotopes together with I. persulcatus with complete coincidence of the seasons of activity of both species at each ontogenetic stage (Filippova 1999) and where their hybridization can also occur (Kovalev et al. 2015; Rar et al. 2019).

In certain areas of Siberia I. pavlovskyi outnumbers I. persulcatus and also other tick species due to the high abundance of ground-feeding birds, especially in urban landscapes with habitats suitable for ticks like parks and cemeteries. So, for example, in the city of Tomsk in Western Siberia I. pavlovskyi dominates everywhere in the city and its outskirts (Romanenko 2011). Probably eventually over time I. persulcatus was gradually replaced by I. pavlovskyi because it is too difficult for adult I. persulcatus to find its preferred hosts, namely mammals (Romanenko and Leonovich 2015).

Filippova and Panova (1998) recognize two subspecies in Russian populations of this tick, namely I. pavlovskyi pavlovskyi and I. pavlovskyi occidentalis which differentiation is based on morphological features between western and eastern specimens.

The type specimens of I. pavlovskyi are deposited at the Zoological Institute of the Russian Academy of Sciences and include I. pavlovskyi, Pomerantsev (Pomerantsev 1946: 11), the holotype: female; [Russia], DVK [Primorskii Terr.], Imanskii Forestry, hazel, 2.9.1932, type; AL I513. Description – Filippova 1977: 305–312 (female, male, nymph, larva); as well as I. pavlovskyi subsp. occidentalis (Filippova and Panova 1998: 396–411 – female, male, nymph, larva) the holotype: female; Russia, western foothills of Kuznetskii Ala Tau, basin of upper Tom River, environs of Mezhdurechensk, from vegetation, flagging, 24.5.1972, coll. E.D. Chigirik, det. N.A. Filippova; AL I1016 and finally I. pavlovskyi subsp. pavlovskyi (Filippova and Panova 1998: 396–411, female, male, nymph, larva), the holotype (the same as the holotype of the species): see I. pavlovskyi (Filippova 2008).

The spectrum of hosts of I. persulcatus is extremely broad both systematically and ecologically and includes more than 200 species of mammals and 100 species of birds (Shilova and Clabovskii 1968). Rarely it can parasitize reptiles – lizards of the family Lacertidae (Ravkin 1969). Literally almost all mammals and birds inhabiting various types of forests and their derivative biotopes can act as hosts for I. persulcatus. Larvae and nymphs parasitize more often small and medium-sized mammals, such as shrews, hedgehogs, rodents, and lagomorphs, as well as ground-feeding and ground-nesting birds. Adults usually feed on large and medium-sized mammals – ungulates, carnivores, lagomorphs. Humans and domestic animals can also be hosts for this tick species (Filippova 1977).

(Fig. 14). The range of I. persulcatus, like no other Palearctic species, is extended in the latitudinal direction by a continuous strip, covering a significant part of the taiga forest zone in Eurasia between 21°–66° latitude in the northern hemisphere from the Scandinavian Peninsula, the Baltic states, Belarus and Ukraine in the west where it is present sporadically to the east up to the Pacific coast including the Kamchatka Peninsula and the Sakhalin Island and further to the north-east of China, the Korean Peninsula and Japan (Filippova 1977; Wang et al. 2023). This tick belongs to the tick fauna of the next post-Soviet countries: Estonia, Latvia, Lithuania, Belarus, Russia, Ukraine, Kazakhstan, Kyrgyzstan (Guglielmone et al. 2023). The presence of I. persulcatus in Ukraine outside the south-west border of the taiga was mentioned by Filippova (1977), although the possibility of permanent populations existing there was disputed by Nebogatkin (1993). Therefore, this probably exemplifies transportation by migratory birds.

Figure 14. 

Map of Russia and neighboring countries showing the locations where Ixodes persulcatus was reported.

Ixodes persulcatus is an exophilic tick species widely distributed in the northern Palearctic along the forest zone. It may use almost all mammals and birds living in its biotopes; therefore, it is one of the most important vectors of a broad range of tick-borne pathogens. Since it can also transmit tick-borne encephalitis virus, together with I. ricinus it has the greatest medical and veterinary significance among other ticks of the genus Ixodes in the Palearctic. Another important fact is that I. persulcatus is a very aggressive species toward humans (Uspensky 1993) and, therefore, this species represents especially high medical-epidemiological risks.

The most significant part of the range of I. persulcatus stretches across the territory of Russia where we can observe the full spectrum of biotopes where I. persulcatus can be found. There are a lot of published works about its ecology in different regions which depend on the climatic region and biotic-abiotic conditions in it.

This tick prefers various types of forest and forest-steppe biotopes, especially taiga forests and their derivatives, i.e., mixed forests and bushes (both plain and mountainous), up to 2000 m a.s.l., like in the Tian Shan. In other words, it can inhabit any herbaceous forest and forest-steppe biotope with the level of humidity high enough for reproduction and supporting the life cycle, even in urban landscapes (Filippova 1977). In the Dzungarian Alatau there were some observations of occurring in steppe regions bordering forests and parasitizing the unusual host, namely the grey marmot Marmota baibacina Kastschenko (Bibikov et al. 1961). Permanent and stable populations of I. persulcatus exist in some areas adjacent to cities within its range and even inside these cities on condition that the the suitable forest environment together with hosts, such as wild animals of different sizes and stray dogs are present. Examples of such cities are Saint Petersburg, Petrozavodsk, Novosibirsk, Tomsk, Irkutsk, and Vladivostok (Uspensky 2017).

Several studies attest the changing boundaries of the ranges of I. persulcatus. It is assumed that ticks of the I. presulcatus group appeared and evolved in forest biotopes similar to modern relict forests of the Ussuri type and the taiga of the mountains of Southern Primorye, Southern Siberia, and the Korean Peninsula in the Pliocene. The wide ecological niche of I. persulcatus was formed during the formation of the species in the process of its adaptation to various landscape and climatic conditions. This allowed the species to gradually expand its range in the northwestern direction in the Holocene (Filippova 2017). An increase in air temperature by one or several degrees in a particular region near the boundaries of its range was probably the main driver of its expanding distribution. The fact of finding I. presulcatus populations in Sweden (Jaenson et al. 2016) and even in the Magadan Oblast in the north-east of Russia where it was absent before (Yamborko et al. 2015) are good examples of the distribution expansion in several directions and confirm the tendency which continues.

In Russia, high numbers of observations show noticeable changes in the distribution of I. persulcatus in certain regions. In Karelia the range expansion of I. persulcatus to the north is noted in relation to general climate warming (Bugmyrin et al. 2013). A similar observation was also recorded in the Komi Republic (Glushakova et al. 2011). The range expansion of this tick species in Arkhangelsk Oblast and Western and Central Siberia to the north is confirmed both by the results of their records and by the data on tick bites and morbidity in the human population, not only in places which were free from ticks before (Pogodina 2021). Besides that, there are some data about the range expansion of I. persulcatus to the north in the Republic of Sakha (Yakutia). The reasons causing these changes are under evaluation but climate change, anthropogenic pressure in natural landscapes as well as the number of vertebrate animals are among the most influential factors. At the same time, it is also possible that inadvertent dispersal of ticks by timber material transported from tick-infested areas may be in part responsible for this phenomenon (Danchinova et al. 2006). Although other factors are not excluded, it is believed that climate changes have made the greatest contribution to the increase in areas primarily for TBE foci in the northern regions of the country. But despite all this, as a result of the same changes, the southwestern part of the range of I. persulcatus in Belarus and the Baltic countries has decreased (Pogodina 2021).

Often it can be found in the same biotopes together with I. ricinus in Europe and I. pavlovskyi in Siberia with complete or partial coincidence of the seasonal activity of these species at each ontogenetic stage (Ushakova and Filippova 1968; Bolotin et al. 1977; Filippova 1999). In zones of sympatry their hybridization can occur, and their hybrids can also transmit tick-borne encephalitis virus and probably other pathogens (Kovalev et al. 2015; Rar et al. 2019; Belova et al. 2023). Under laboratory conditions, interspecific hybridization between I. ricinus and I. persulcatus was successfully conducted as well. F1 hybrid ticks were completely sterile, as revealed by unsuccessful attempts of their subsequent hybridization with ticks of the parent generation (Balashov et al. 1998). In I. persulcatus and I. ricinus, any morphological barrier to crossing is undoubdetly absent and then sterility of the F1 hybrid generation is probably a quite significant factor limiting the population size of both species in their sympatric areas. Hybrid ticks also have morphological features allowing to differentiate them at preimaginal and imaginal stages (Bugmyrin et al. 2015, 2016). Moreover, some studies were conducted in the Southern Primorye (Filippova 2002) in sympatric zones of I. persulcatus and I. pavlovskyi occidentalis, due to the close cohabitation of both species. These showed that in case of these two species there are distinct morphological barriers which are manifested in the fitting of organs involved in mating, in particular their size proportions. According to the result of the studies, mating and hybridization of different tick species are possible only in the next combination: female I. pavlovskyi and male I. persulcatus. Whereas in case of the reverse combination, the parameters of the genital aperture of the female exceed those of the largest width of the hypostome in the male.

There is an excellent summary on the questing behavior of I. persulcatus in the monograph by Filippova (1985). In brief, the ticks climb onto the vegetation in quest of a host. When the host approaches, the tick spreads its first pair of legs and, upon contact with the host, become attached. From time to time, ticks perform vertical migrations and go even into the soil litter for rehydration. Horizontal movements of ticks towards trails used by potential hosts are also possible, as well as crawling onto a nearby animal. Ticks react to humans by spreading their first legs from distances of ~ 15–20 m. At short distances, ticks also react to a heat source. In general, a similar pattern of questing behavior is used by other exophilic ticks of the genus Ixodes.

In I. persulcatus there is an important signaling mechanism causing a morphogenetic diapause – a developmental delay which is the response of ticks to the duration of the diurnal photoperiod (Belozerov 1976). Moreover, I. persulcatus has a behavioral diapause of non-engorged adult ticks, which is not connected with photoperiodic regulation (Korenberg et al. 2021). But as the studies in the Kirov Oblast and Udmurt Republic showed, in more warmer areas, an increased proportion of engorged larvae and nymphs develop without the diapause and the reason for this is the early activation and, as a result, their mass feeding on hosts in the first half of summer. The factors determining the diapause of engorged larvae and nymphs in the compared regions practically do not differ (Korotkov 2008). The correlation of the tick number varies, depending on the type of biotope, as well as temperature and humidity and also many other abiotic factors. For example, in boreal taiga forests of Karelia mainly I. persulcatus dominates (except the southwestern part where the mass species is I. ricinus) (Bugmyrin et al. 2013). The beginning of adult I. persulcatus activity also differs in different regions depending on the sum of abiotic factors listed above. For example, in the Far East the seasonal peak in the number of larvae is observed in the third decade of May – second decade of July, whereas in the European part of its range in the third decade of July (Belozerov 1976; Filippova 1977; Balashov 1998; Korenberg et al. 2013). In the territory from the Volga River to Primorye the average activity of adult ticks varies from 60 to 140 days (Korenberg et al. 1974). The boundaries of the range of the tick are determined mainly by the combination of photo- and hygrothermal factors. The general indicators of warmth and moisture along the range of this tick species vary widely. The fundamental ecological niche of I. persulcatus with the broad scope of its preferred conditions allows it to adapt to the wide diversity of biotopes in the forest zone.

Some type specimens of I. persulcatus are deposited at the Zoological Institute of the Russian Academy of Sciences and include I. persulcatus subsp. diversipalpis (Schulze 1930: 300), lectotype: male; [Russia, Primorskii Terr.], lower Amur River, 8 km of Vyatskoe Vill., 26.VI.1910, coll. Soldatov, det. N.О. Olenev: I. ricinus ovatus; AL I266, as well as the paralectotypes: 1 female, 1 male; AL I266a. I. persulcatus (see: Filippova 1969: 677). Description – Filippova 1977: 316–327 (female, male, nymph, larva) (Filippova 2008). But Filippova (1969) also states that re-examination of the type material of the above subspecies demonstrated that the specimens used for describing differences of this subspecies are damaged in some morphologically important parts (not noticed before), and the key morphological characters that were previously thought to distinguish the subspecies are not specific enough and can be found in ticks throughout their entire geographical range.

Mammalia: Apodemus agrarius (striped field mouse), Apodemus mystacinus (Danford and Alston) (eastern broad-toothed field mouse), Apodemus uralensis (Ural field mouse), Arvicola amphibius (European water vole), Chionomys nivalis (European snow vole), Chionomys roberti (Thomas) (Robert’s snow vole), Cricetus cricetus (European hamster), Crocidura leucodon (bicolored shrew), Crocidura suaveolens (lesser white-toothed shrew), Dryomys nitedula (forest dormouse), Erinaceus europaeus (European hedgehog), Glis glis (European edible dormouse), Hemiechinus auratus (long-eared hedgehog), Lepus europaeus (European hare), Marmota bobak (bobak marmot), Martes martes (European pine marten), Meles meles (European badger), Meriones libycus (Libyan jird), Meriones meridianus (midday jird), Meriones persicus (Persian jird), Meriones tamariscinus (Pallas) (tamarisk jird), Meriones tristrami Thomas (Tristram’s jird), Mesocricetus auratus Waterhouse (golden hamster), Mesocricetus raddei (Ciscaucasian hamster), Microtus arvalis (common vole), Microtus majori (Major’s pine vole), Microtus socialis (social vole), Mus musculus (house mouse), Mustela eversmanii (steppe polecat), Mustela nivalis (least weasel), Nesokia indica (short-tailed bandicoot rat), Nothocricetulus migratorius (grey dwarf hamster), Rattus norvegicus (brown rat), Rattus pyctoris (Turkestan rat), Rattus rattus (black rat), Rhombomys opimus (great gerbil), Sciurus anomalus Gmelin (Caucasian squirrel), Sciurus vulgaris (red squirrel), Sicista betulina (northern birch mouse), Sicista subtilis (southern birch mouse) Spalax giganteus Nehring (giant blind mole-rat), Spalax microphthalmos Gueldenstaedt (greater blind mole-rat), Spermophilopsis leptodactylus (long-clawed ground squirrel), Spermophilus pygmaeus (little ground squirrel), Sorex araneus (common shrew), Vormela peregusna (marbled polecat) Vulpes vulpes (red fox) (Filippova 1977).

Aves: Alauda arvensis Linnaeus (Eurasian skylark), Alectoris chukar (chukar partridge), Anthus campestris (tawny pipit), Anthus pratensis (Linnaeus) (meadow pipit), Coccothraustes coccothraustes (hawfinch), Columba livia (rock dove), Emberiza calandra (corn bunting), Emberiza schoeniclus (Linnaeus) (common reed bunting), Erithacus rubecula (Linnaeus) (European robin), Galerida cristata (crested lark), Garrulus glandarius (Linnaeus) (Eurasian jay), Lullula arborea (woodlark), Melanocorypha calandra (Linnaeus) (calandra lark), Mergus serrator Linnaeus (red-breasted merganser), Oenanthe hispanica (Linnaeus) (western black-eared wheatear), Oenanthe isabellina (Temminck) (Isabelline wheatear), Oenanthe lugens (Lichtenstein) (mourning wheatear), Oenanthe oenanthe (Linnaeus) (northern wheatear), Oenanthe picata (Blyth) (variable wheatear), Phylloscopus collybita (Vieillot) (common chiffchaff), Phylloscopus fuscatus (dusky warbler), Pica pica (Eurasian magpie), Pterocles orientalis (Linnaeus) (black-bellied sandgrouse), Saxicola torquatus (Linnaeus) (African stonechat), Sturnus vulgaris (common starling), Turdus merula (common blackbird), Turdus philomelos (song thrush), Turdus ruficollis (red-throated thrush) (Filippova 1977).

Reptilia: Darevskia chlorogaster (Boulenger) (greenbelly lizard) (Orlova et al. 2022), Lacerta agilis Linnaeus (sand lizard) (Filippova 1977), Lacerta strigata Eichwald (Caucasus emerald lizard) (Orlova et al. 2023), Pseudopus apodus (Pallas) (Pallas’s glass lizard) (Filippova 1977).

(Fig. 15). Russia: Rostov Oblast (Khametova et al. 2018), Krasnodar Krai, Stavropol Krai, Kalmyk Republic, Chechnya, Dagestan, and North Osetia-Alania (Shatas 1957; Shevchenko et al. 1960; Zaytsev and Popova 1967; Tiflova 1974; Filippova 1977; Abdulmagomedov et al. 2017; Zaytseva et al. 2022). Ukraine: Odesa Oblast (Bugeac Steppe), Kherson Oblast (Black Sea Biosphere Reserve), Poltava Oblast, Chernivtsi Oblast, Dnipropetrovsk Oblast, Luhansk Oblast, Donetsk Oblast, widely distributed in the Crimean Peninsula (Emchuk 1960; Emchuk 1967; Sklyar 1970; Filippova 1977). Moldova: the north of the country (Uspenskaya et al. 2006). Georgia: outskirts of Kutaisi and Tbilisi and the Lagodekhi Nature Reserve, as well as the seacoast of the Black Sea (Kirschenblatt 1936; Djaparidze 1960; Filippova 1977). Armenia: outskirts of Yerevan and most of the rest of the territory (Zilfyan et al. 1960; Tiflova 1974). Azerbaijan: Zagatala State Reserve, Hadrut District, and the Mil plain (Tiflova 1974), outskirts of the Bilasuvar, the Sara Peninsula (Kirschenblatt 1936), Talysh (Pomerantsev 1950), Nakhchivan Autonomous Republic (Kadatskaya and Shirova 1963; Filippova 1977). Kazakhstan: West Kazakhstan Region, Kyzylorda Region, North Kazakhstan Region, Jambyl Region, Turkistan Region, Abai Region (Loseva 1963; Popova and Sokolova 1963). Kyrgyzstan: outskirts of Bishkek, Chüy Valley, Talas Valley, Issyk-Kul Basin, Terskey Ala-too Range (Filippova 1958b; Grebenyuk 1966; Filippova 1977). Turkmenistan: foothills of the Uly Balkan and the Kopet Dagh; the Kugitangtau Range (Kochkareva et al. 1971; Filippova 1977). Uzbekistan: outskirts of Tashkent, foothills of the Chatkal Range, Qurama Mountains, the Hisar Range, the Kugitangtau Range and Karakalpakstan – the Ustyurt Plateau and the lower reaches of the Amu Darya River (Kuklina 1967; Uzakov 1972; Filippova 1977). Tajikistan: Hisar Range - Varzob gorge, outskirts of Dushanbe – the Ramit State Nature Reserve, Vakhsh Range, Peter the First Range (Lotozky 1951; Sosnina 1957; Filippova et al. 1966; Kochkareva et al. 1971; Filippova 1977).

Figure 15. 

Map of Russia and neighboring countries showing the locations where Ixodes redikorzevi was reported.

Ixodes redikorzevi is a tick species which is mainly a parasite of rodents, shrews, and small carnivores, as well as of dendrophilic ground-feeding birds and rarely reptiles (Filippova 1977). According to Tiflova (1974), this species is considered exophilic and can be found in significant numbers on dendrophilic birds. In the absence of mammalian and avian hosts, I. redikorzevi can parasitize lizards in significant numbers (Orlova et al. 2022). It usually inhabits mountain deciduous forests and steppes located nearby.

Beyond the post-Soviet territories considered above, the range of this tick covers also Eastern Europe, Turkey, Israel, as well as Afghanistan (Filippova 1977) and China (Yin et al. 2010).

At the current moment it is still questionable whether I. redikorzevi is a synonym of I. acuminatus or not. Kolonin (2009) considers this species a synonym of I. acuminatus, but Guglielmone et al. (2010) regard it as provisionally valid. As it was fairly noted by Guglielmone et al. (2014) this question can be solved by comparison of the type specimens of both species. Moreover, Pomerantsev (1950) described by females two subspecies: I. redikorzevi redikorzevi and I. redikorzevi emberizae. Later the other subspecies I. redikorzevi theodori was described although Filippova comments (1977) that the authors had quite little material during descriptions but the differences in size and shape of some characters are visible and it is necessary to compare more specimens from more locations of its large area of distribution.

Ixodes redikorzevi redikorzevi occurs in Ukraine, the Transcaucasus and Tajikistan according to Pomerantsev (1950); and I. redikorzevi emberizae can be found in Lankaran and the Hisar Range in Tajikistan. Later the other subspecies, I. redikorzevi theodori was described from the Middle East (Warburton 1927).

The type specimens of I. redikorzevi are deposited at the Zoological Institute of the Russian Academy of Sciences and include the holotype: female; [former] Tavricheskaya Province (Crimea), Yaman-Kala, near Baidar, 25.10.1924, coll. V. Shnitnikov, AL I338 and the paralectotype of I. redikorzevi emberizae female; AL I522. Description – Pomerantsev 1950: 63 (female; male unknown); Filippova 1977: nymph, larva (Filippova 2008).

The host spectrum of I. ricinus is extremely broad both systematically and ecologically, including literally almost all mammals and birds of its geographical range, rarely even reptiles inhabiting the same biotopes with the tick. The fact of mass parasitism of immature stages on lizards of the Lacertidae family, in particular species of the genus Darevskia in the Caucasus (Kidov et al. 2013; Orlova et al. 2022) in habitats where they outnumber small mammals probably brightly demonstrates that I. ricinus is a generalist tick capable to use almost any available terrestrial vertebrates as hosts. Overall, the list of hosts consists of more than 300 species of mammals, birds and reptiles which have been recorded (Gern et al. 2002). Humans and domestic animals can also be hosts for the tick (Filippova 1977).

(Fig. 16). The distribution of I. ricinus in Russia includes almost the whole territory of its European part excluding subpolar tundra areas (see the map) (Filippova 1977; Kahl and Gray 2023) and due to climate changes, the distribution of this tick species becomes wider (Gray et al. 2009; Yasyukevich et al. 2009). Ixodes ricinus is part of the tick fauna of the following post-Soviet countries: Estonia, Latvia, Lithuania, Belarus, Russia, Ukraine, Moldova, Georgia, Azerbaijan, Armenia, Turkmenistan, and Kazakhstan (Guglielmone et al. 2023). In Kazakhstan a little number of specimens were found in the northern part of West Kazakhstan Oblast (Maikanov 2012). In Turkmenistan the tick was also recorded in few numbers in the western foothills of the Kopet-Dag (Kerbabaev 1960) which probably could be transported there by migratory birds.

Figure 16. 

Map of Russia and neighboring countries showing the locations where Ixodes ricinus was reported: A before 1975 B from 1976.

Ixodes ricinus is an exophilic tick species widely distributed in Europe, mostly inhabiting deciduous and mixed forest zones in both plain and mountainous areas, as well as forest-steppes bordering them. It also occurs in city parks and gardens (Gray 1998). In addition, it can be found in North Africa (Arthur 1965). In Ukraine I. ricinus colonized and reached a high abundance in artificial forest plantations of the Askania-Nova Nature Reserve surrounded from all sides by steppes for a period of less than 80 years (Emchuk 1972). In urban areas with conditions able to support tick populations, for example, Minsk or Kyiv, I. ricinus usually dominates among other tick species, especially among members of the genus Ixodes (Uspensky 2017). This tick species uses almost all forest vertebrate animals as hosts and, together with I. persulcatus, it is one of the most important vectors of a broad spectrum of tick-borne pathogens, first of all, tick-borne encephalitis virus (Filippova 1977).

Often it can be found in the same biotope with I. persulcatus, often exhibiting complete or partial coincidence of seasonal activity at each ontogenetic stage (Filippova 1999). In zones of sympatry their hybridization can occur, and although hybrid offspring are incapable of reproduction (Bugmyrin et al. 2015), they can still transmit tick-borne encephalitis virus and probably other pathogens (Kovalev et al. 2016; Belova et al. 2023). The absence of any morphological barrier for copulation was discovered in geographical points of probably the secondary sympatric zone (Filippova 2002) of I. persulcatus and I. ricinus in the north-west of the East European Plain (Balashov et al. 1998). However, in some areas of this sympatric zone, for example, in southern Karelia, its slight shrinking has recently been noted due to the withdrawal of I. ricinus from territories where it used to live (Bespyatova and Bugmyrin 2021).

Due to the high epidemiological significance and wide distribution of I. ricinus and its regular contacts with humans and domestic animals, its biology and life cycle were more extensively studied than in case of any other species of its genus inhabiting the same territories. As a species, I. ricinus probably appeared approximately 8–12 thousand years ago when deciduous and mixed forests formed in the southeast of Europe and the Mediterranean, as well as in the northern and northeastern slopes of the Greater Caucasus, when current environmental conditions of these territories have begun to shape. And the climate there was also milder than in Siberian taiga forests where I. persulcatus evolved (Filippova 2017).

It was revealed that in a certain region the duration of tick activity period and the number of adult ticks depend on spring and summer temperatures and air humidity (Korotkov et al. 2015; Korenberg et al. 2021). Females and larvae usually attach to hosts when the air near the soil warms up from +2 to +30 °С, and in the case of nymphs from +2 to +22 °С. The relative humidity of the surrounding air has to be higher than 60% for an extended period of time (Sirotkin and Korenberg 2018). It is absolutely important for ticks to receive the necessary amount of warmth to complete their metamorphosis at each stage within a strictly defined period of time (Korenberg et al. 2013). As a consequence, the seasonal activity of all stages of I. ricinus is more extended than in the case of I. persulcatus, and engorged ticks begin oviposition or metamorphosis without strict dependance on the photoperiod. Therefore, in the southern range of distribution (the Mediterranean, Central Europe, the Caucasus) ticks initiate activity in the end of March – the beginning of April (Korenberg et al. 2021), whereas in Eastern European regions – in April (Medvedev et al. 2016; Korenberg et al. 2021). Ixodes ricinus also uses a diapause as a biological mechanism, although due to warmer conditions in the majority of its distribution range, no more than 10–20% of ticks at each stage undergo such an interruption of development (Korenberg and Kovalevsky 1977; Korenberg et al. 2016).

Mammalia: Lepus timidus (mountain hare) (Filippova 1977).

(Fig. 17). Russia: the Sakhalin Island, Sachalin Oblast, the rural locality Khomutovo (Filippova 1971).

Figure 17. 

Map of Russia and neighboring countries showing the locations where Ixodes sachalinensis was reported.

Ixodes sachalinensis is a tick species known only by the single finding from Sakhalin. It was collected from a mountain hare together with 79 females, 15 males and 7 nymphs of I. persulcatus (Filippova 1971).

Kolonin (2009) and Camicas et al. (1998) consider I. sachalinensis a synonym of I. persulcatus, but Barker and Murrell (2004) and Guglielmone et al. (2009, 2010) recognize this species as valid.

The type specimen is deposited at the Zoological Institute of the Russian Academy of Sciences and includes the holotype: female; [Russia], Sakhalin, near Khomutovo Vill., Lepus timidus, 27.5.1950, [coll.: unknown]; AL I729 (Filippova 2008).

Mammalia: Alexandromys oeconomus (tundra vole), Craseomys rufocanus (grey red-backed vole), Eutamias sibiricus (Siberian chipmunk), Mus musculus (house mouse), Myodes rutilus (northern red-backed vole), Ochotona alpina (alpine pika), Rattus norvegicus (brown rat), Sicista caudata Thomas (long-tailed birch mouse), Sorex araneus (common shrew), Sorex minutus (Eurasian pygmy shrew) (Filippova 1977).

(Fig. 18). Russia: outskirts of Magadan and the lower reaches of the Kukhtui River, Okhotsky district – the northernmost points of record of I. angustus in the Palearctic (Belyaev 1963); Kamchatka Peninsula – outskirts of the villages Tigil and Ust-Khayryuzovo (Pomerantsev 1950), the valley of the Kamchatka River to Ust-Kamchatsk (Serdjukova 1956), the eastern coast of the Kamchatka peninsula to Petropavlovsk-Kamchatsky (Speranskaya 1958), the valley of the rivers Avacha and Pinachevskaya (Paramonov et al. 1966); Middle Outer Manchuria (Filippova 1977); Sovetsko-Gavansky district (Emelyanova and Koshkin 1962); Sikhote-Alin (Belyaev and Filippova 1976); Sakhalin – Novoaleksandrovka (former Konuma), the valley of the Lyutoga River (Pomerantsev 1950) and the Cape Patience (Skrynnik 1950; Asanuma 1951; Violovich 1958, 1960; Savitsky and Okuntsova 1967; Timofeeva and Kon’kova 1971); Kuril Islands – Simushir (Pomerantsev 1950; Violovich 1958, 1960; Timofeeva and Kon’kova 1971).

Figure 18. 

Map of Russia and neighboring countries showing the locations where Ixodes angustus was reported.

Ixodes angustus occurs in the Palearctic predominantly on the East Asian coast and also in the Nearctic – Canada and the USA (Filippova 1977). In the Russian Far East in Outer Manchuria, the islands and along the main ridges of the Sikhote-Alin it inhabits a wide range of biotopes: various types of mixed and broad-leaved forests in mountains and valleys, as well as tundra and rocks, stone outcrops, coastal biotopes, meadow and river valleys (Speranskaya 1958; Violovich 1958; Emelyanova and Koshkin 1962; Belyaev 1963; Paramonov et al. 1966; Savitsky and Okuntsova 1967; Belyaev and Filippova 1976).

Ixodes angustus is considered a nidicolous ectoparasite of rodents and shrews because it was found not only on hosts but also in their burrows (Filippova 1977), although there are documented cases on this species biting humans without contacts with burrows (Cooley 1946). As a parasite which is connected with rodents, and, like other rodent ticks, I. angustus plays a role in supporting natural foci of tick-borne infections such as anaplasmosis (Yamborko and Eremeeva 2014) and the Lyme disease (Peavey et al. 2000).

Although hyperparasitism is not common in Ixodes ticks, I. angustus belongs to a small number of species of the genus, in which this phenomenon was recorded (Durden et al. 2018), when a male was feeding from a female attached to a red squirrel Tamiasciurus hudsonicus. The other Ixodes species in which males have been recorded to attach and feed on engorging conspecific females include I. holocyclus in Australia and I. pilosus in South Africa (Oliver et al. 1986).

Mammalia: Apodemus agrarius (striped field mouse), Craseomys rufocanus (grey red-backed vole), Erinaceus amurensis Schrenk (Amur hedgehog), Eutamias sibiricus (Siberian chipmunk), Microtus fortis (reed vole), Myodes rutilus (northern red-backed vole), Sorex araneus (common shrew) (Filippova 1977), Sorex caecutiens (Laxmann’s shrew), Sorex unguiculatus Dobson (long-clawed shrew), (individual specimens ((Okulova et al. 1986), Rattus norvegicus (brown rat), Tscherskia triton (De Winton) (greater long-tailed hamster) (Filippova 1977).

(Fig. 19). Russia: Sikhote-Alin – outskirts of Dal’ny Kut (the northernmost point of finding (Filippova 1977), valley of the Dorozhnaya River, Dalnegorsk, Ussurisky (former Komarovskii) Nature Reserve; coast of the Sea of Japan – outskirts of the villages Terney, Dukhovo, Kamenka, Lazovsky nature reserve, Fokino (former Promyslovka), the bays Razboynik and Linda; the coast of the Peter the Great Gulf: Kedrovaya Pad Nature Reserve, the rural localities Barabash and Posyet (Serdjukova 1941; Pomerantsev 1950; Slonov 1961; Khudyakov 1963; Belyaev and Filippova 1976).

Figure 19. 

Map of Russia and neighboring countries showing the locations where Ixodes pomerantzevi was reported.

Ixodes pomerantzevi is a relict species occurring on the East Asian coast (Filippova 1977) and in Russia its distribution is limited to a few locations in Outer Manchuria (a.k.a. Primorsky Krai) in the Russian Far East (Tsapko 2020). It is also known to occur in Korea (Kim et al. 2009a, 2010, 2011) and China (Guo et al. 2016). Predominantly it can be found in coniferous and broad-leaf forests, or secondary forests and bush thickets, as well as rock and stone outcrops among trees in the Sikhote-Alin and on the coast of the Sea of Japan (Belyaev and Filippova 1976).

Luh and Woo (1950) supposed that I. pomerantzevi is possibly a synonym of Ixodes angustus; Filippova (1977) considered it as valid and in the last list of valid tick species names, it is also considered valid (Guglielmone et al. 2020).

Ixodes pomerantzevi is a nidicolous tick species, an ectoparasite of rodents, hedgehogs, and shrews (Filippova 1977).

The type specimen of I. pomerantzevi is deposited at the Zoological Institute of the Russian Academy of Sciences and include the holotype: female; [Russia], DVK [Primorskii Terr.], Suputinskii [Komarovskii or Ussurisky] Nature Reserve, from Myodes rufocanus, 9–13.VI.1939, coll. B.I. Pomerantsev; AL I502. Description – Filippova 1977: 128–132 (female, male - unknown, nymph, larva) (Filippova 2008).

Mammalia: Alticola argentatus (Severtzov) (silver mountain vole), Apodemus agrarius (striped field mouse), Craseomys rufocanus (grey red-backed vole), Crocidura sp. (shrew), Lasiopodomys gregalis (narrow-headed vole), Microtus arvalis (common vole), Mustela sp. (weasel), Myodes centralis (Miller) (Tien Shan red-backed vole), Nothocricetulus migratorius (grey dwarf hamster), Ochotona macrotis (Günther) (large-eared pika), Rattus pyctoris (Turkestan rat) (Filippova 1977).

(Fig. 20). Russia: Western Sayan (Arumova and Dineva 1973). Kazakhstan: Tarbagatai Mountains (Afanas’eva 1959), Dzungarian Alatau (Ushakova and Fedosenko 1963; Ushakova et al. 1976), Trans-Ili Alatau (Ushakova and Fedosenko 1963). Kyrgyzstan: Kyrgyz Ala-Too Range (Fedorova 2012b), Terskey Alatau (Fedorova 2012b), Chuy Valley – found in 1966 (Grebenyuk 1966), was not found in the same territories in 2018 (Fedorova 2021). Tajikistan: Peter the First Range (Filippova 1977), Varzob gorge (Sosnina 1954 – here I. stromi was incorrectly identified as I. trianguliceps because the new species was described by Filippova in 1957b).

Figure 20. 

Map of Russia and neighboring countries showing the locations where Ixodes stromi was reported.

Ixodes stromi is a tick species only indigenous to southern Siberia in Russia (Tsapko 2020). The main part of its distribution spans in Kazakhstan and Middle Asia. In all territories of its range, it is confined to the forest-meadow and forest-steppe belt of medium-altitude mountains, to stony and rocky habitats, which are insolated and, therefore, have a warmer microclimate (Filippova 1967).

This species is nidicolous and uses rodents, shrews, and small carnivores as hosts at all stages. It is considered a rare species reaching small individual number (Filippova 1977).

The type specimens of I. stromi are known from Kyrgyzstan and stored at the Zoological Institute of the Russian Academy of Sciences: the lectotype: the nymph; Kyrgyzstan, Tien Shan, Kungei Ala Tau Mt. Range, Ch-Aksu Canyon, talus, from Clethrionomys frater (synonym of Myodes centralis), 11.VIII.1953, coll. N.А. Filippova; AL I78. The paralectotypes: 6 larvae; FBM I586, I876; 6 larvae; FBM I873, I875. Description – Filippova 1977: 122–127 (female, nymph, larva; male unknown) (Filippova 2008).

To date hosts of this tick species are unknown (Guglielmone et al. 2014).

(Fig. 21). Russia: Khabarovsk Krai – Khekhtsir Range and the rural locality Vyatskoye (Emelyanova and Kozlovskaya 1967); Krasnoyarsk Krai – Kozulsky District, the village Bolshoy Kemchug (Voltsyt 1997).

Figure 21. 

Map of Russia and neighboring countries showing the locations where Ixodes maslovi was reported.

Ixodes maslovi is an almost unstudied tick known and described from two findings of its male and female (Emelyanova and Kozlovskaya 1967), as well as the nymph (Voltsyt 1997).

Camicas et al. (1998) and Kolonin (2009) regard I. maslovi as an abnormal form of I. persulcatus although Filippova (1977) and Guglielmone et al. (2020) consider I. maslovi a valid taxon.

The type specimens are deposited at the Zoological Institute of the Russian Academy of Sciences – the holotype: male; [Russia], environs of Khabarovsk, Khekhtsir Mt. Range, 12.VI.1964, collected from vegetation by O.L. Kozlovskaya; FBM I1412; the paratype: female; FBM I1413. Description – Filippova 1977: 248–251 (female, male); Voltsit 1997: 265–268 (nymph; larva unknown) (Filippova 2008).

Aves: Accipiter gentilis (Linnaeus) (northern goshawk), Acrocephalus scirpaceus (Hermann) (Eurasian reed warbler), Aegithalos caudatus (Linnaeus) (long-tailed tit), Aegolius funereus (Linnaeus) (boreal owl), Athene noctua (little owl), Certhia brachydactyla Brehm (short-toed treecreeper), Certhia familiaris Linnaeus (Eurasian treecreeper), Chloris chloris (European greenfinch), Coloeus monedula (Linnaeus) (western jackdaw), Columba palumbus Linnaeus (common wood pigeon), Coracias garrulus Linnaeus (European roller), Corvus frugilegus Linnaeus (rook), Curruca communis (common whitethroat), Cyanistes caeruleus (Linnaeus) (Eurasian blue tit), Dendrocopos major (great spotted woodpecker), Emberiza citrinella (yellowhammer), Erithacus rubecula (European robin), Falco peregrinus Tunstall (peregrine falcon), Falco tinnunculus Linnaeus (common kestrel), Ficedula albicollis (Temminck) (collared flycatcher), Ficedula hypoleuca (European pied flycatcher), Garrulus glandarius (Eurasian jay), Glaucidium passerinum (Linnaeus) (Eurasian pygmy owl), Hirundo rustica Linnaeus (barn swallow), Lophophanes cristatus (Linnaeus) (crested tit), Motacilla alba (white wagtail), Muscicapa striata (Pallas) (spotted flycatcher), Parus major (great tit), Passer domesticus (house sparrow), Passer montanus (Eurasian tree sparrow), Periparus ater (Linnaeus) (coal tit), Phoenicurus ochruros (Gmelin) (black redstart), Phoenicurus phoenicurus (common redstart), Phylloscopus trochilus (Linnaeus) (willow warbler), Picus canus Gmelin (grey-headed woodpecker), Poecile montanus (Conrad von Baldenstein) (willow tit), Poecile palustris (Linnaeus) (marsh tit), Pyrrhula pyrrhula (Linnaeus) (Eurasian bullfinch), Remiz pendulinus (Linnaeus) (Eurasian penduline tit), Riparia riparia (Linnaeus) (sand martin), Serinus serinus (Linnaeus) (European serin), Sitta europaea (Eurasian nuthatch) (Filippova 1977; Keve et al. 2022), western rock nuthatch Sitta neumayer Michahelles (Ogandzhanyan 1984), Spinus spinus (Linnaeus) (Eurasian siskin), Strix aluco Linnaeus (tawny owl), Sturnus vulgaris (common starling), Troglodytes troglodytes (Linnaeus) (Eurasian wren), Turdus merula (common blackbird), Turdus philomelos (song thrush), Tyto alba (Scopoli) (barn owl), Upupa epops Linnaeus (Eurasian hoopoe) (Filippova 1977; Keve et al. 2022).

(Fig. 22). Russia: Southern Primorskyi Krai (Pogranichny District, Vladivostok, Nakhodka, Putyatin Island) (Emelyanova and Gordeeva 1969; Emelyanova 1972; Bolotin 2000). Ukraine: outskirts of Kyiv (Nebogatkin 2014), Dnipropetrovsk Oblast (the rural locality Andriivka), Crimea (Olenivka and Alushta) (Filippova 1977). Belarus: Białowieża Forest, Minsk, Gomel Oblast (the village Markovskoye) (Gembetsky 1966, 1972). Armenia: Syunik Province (former Goris Province) (Ogandzhanyan 1984). Azerbaijan: Karabakh (Kirschenblatt 1936). Georgia: Lagodekhi Nature Reserve (Djaparidze 1960). Kyrgyzstan (Fedorova 2012a).

Figure 22. 

Map of Russia and neighboring countries showing the locations where Ixodes arboricola was reported.

Ixodes arboricola is an endophilic parasite, mainly of birds from ecological groups nesting in tree holes and nest boxes and also even in ground burrows (Filippova 1977). Also, certain cases of this species infesting bats in tree holes have been recorded (Arthur 1963).

The interesting feature of its distribution is the disjunctivity, which is confirmed by the discovery of this species in the areas quite distant from each other – western and central Europe, North Africa, Transcaucasia, western Asia, and the Far East in Russia (Estrada-Peña et al. 2018) and China (Chen et al. 2010).

Mammalia: Mustela erminea Linnaeus (stoat) (Filippova 1977).

(Fig. 23). Tajikistan: Peter the First Range, the source of the Divansu River, close to the Oshanin glacier (Filippova 1977).

Figure 23. 

Map of Russia and neighboring countries showing the locations where Ixodes cornutus was reported.

Ixodes cornutus is a species described from two identical females (Lotozky 1956) that were found in Tajikistan, in the eastern part of Peter the First Range, by the source of the Divansu River (the basin of the Surkhob River), near the Oshanin glacier, on a stoat.

The type specimen of I. cornutus is deposited at the Zoological Institute of the Russian Academy of Sciences (Lotozky 1956: 27). Lectotype: female; 38 [Tajikistan, the Peter the First Mt. Range], the source of the Divansu River, the ancient moraine of the Oshanin glacier, Mustela erminea, ad.; male; 4.VII.1954; AL I845. Description – Filippova 1977: 178 (female; male, nymph, larva unknown) (Filippova 2008).

Tick names are used sensu Guglielmone et al. (2014) in this review. Thus, this species is not synonymous with I. canisuga Johnston as suggested by Filippova (1977) based on their morphological similarities and because the latter is not known to occur in Russia. Ixodes crenulatus was erroneously synonymized with I. kaiseri Arthur (Sonenshine et al. 1969), as clarified later (Filippova and Uspenskaya 1973).

Mammalia: Allactaga major (Kerr) (great jerboa), Allactaga sibirica (Forster) (Mongolian five-toed jerboa), Allocricetulus eversmanni (Brandt) (Eversmann’s hamster), Apodemus sylvaticus (wood mouse), Canis aureus Linnaeus (golden jackal), Canis familiaris (domestic dog), Canis lupus Linnaeus (gray wolf), Cricetulus barabensis (Pallas) (Chinese striped hamster), Ellobius talpinus (northern mole vole), Erinaceus europaeus (European hedgehog), Felis catus Linnaeus (domestic cat), Felis lybica Forster (African wildcat), Hemiechinus auratus (long-eared hedgehog), Homo sapiens Linnaeus (human), Lasiopodomys gregalis (narrow-headed vole), Lepus tolai (tolai hare), Marmota baibacina (gray marmot), Marmota bobak (bobak marmot), Marmota caudata (Geoffroy) (long-tailed marmot), Marmota kastschenkoi Stroganov and Yudin (forest-steppe marmot), Marmota menzbieri (Kashkarov) (Menzbier’s marmot), Marmota sibirica (Tarbagan marmot), Meles meles (Eurasian badger), Microtus arvalis (common vole), Mustela eversmanii (steppe polecat), Mustela nivalis (least weasel), Myodes glareolus (bank vole), Myospalax myospalax (Siberian zokor), Nothocricetulus migratorius (grey dwarf hamster), Nyctereutes procyonoides (Gray) (common raccoon dog), Ochotona dauurica (Pallas) (Daurian pika), Ochotona pallasi (Gray) (Pallas’s pika), Otocolobus manul (Pallas) (Pallas’s cat), Ovis aries (domestic sheep), Phodopus sungorus (Pallas) (winter white dwarf hamster), Procyon lotor (Linnaeus) (raccoon), Spermophilus dauricus Brandt (Daurian ground squirrel), Spermophilus pygmaeus (little ground squirrel), Spermophilus relictus (Kashkarov) (relict ground squirrel), Spermophilus suslicus (speckled ground squirrel), Vulpes corsac (corsac fox), Vulpes vulpes (red fox) (Filippova 1977; Litvinov and Sapegina 2003; Kalyagin et al. 2005, 2008).

Aves: Emberiza cia Linnaeus (rock bunting), Oenanthe isabellina (isabelline wheatear) (Filippova 1977).

(Fig. 24). Russia: Tula Oblast (Myasnikov and Katelina 1964), Kursk Oblast (Lgovsky District), Voronezh Oblast (Kamennaya Steppe Nature reserve), Rostov Oblast (Aksay), Republic of Kalmykia (Derbetovsky District, Sarpinsky District), Volgograd Oblast (Gorodishchensky, Derbetovsky and Sarpinsky District) (Denisov 2010, 2019), Kabardino-Balkaria (tract Khaimasha) (Bittirova et al. 2019), Dagestan (Aliev et al. 2007), Astrakhan Oblast, Stavropol Krai (Filippova 1977), Saratov Oblast (Turtseva 2007; Denisov 2010, 2019; Porshakov et al. 2020), Yekaterinburg (Milintsevich et al. 2016), Tyumen Oblast (Glazunov and Zotova 2014), Kurgan Oblast (Starikov and Starikova 2021), Novosibirsk Oblast (Suzunsky, Karasuksky and Maslyaninsky District) (Davydova and Lukin 1969), Omsk Oblast (Tarasevich et al. 1971), Kemerovo Oblast (Kalyagin et al. 2005, 2008; Kovalevsky et al. 2018); Altai Krai (Oberth et al. 2015) (Sovetsky District, the village Kokshi) (Filippova 1977), Altai Republic (Shebalinsky District, the village Cherga) (Litvinov and Sapegina 2003), Tuva (Glazunov and Zotova 2014; Filippova 1977), Transbaikal (villages Borgoy, Kyakhta, Selenge and Borzinsky District) (Filippova 1977); Amur Oblast (village Krasny Vostok), Southern Outer Manchuria (Khankaysky District (Kolonin 1986; Bolotin 2000). Ukraine: outskirts of Kyiv (Akimov and Nebogatkin 2016), Zakarpattia Oblast and Western Ukraine in general (Podobivskyi and Fedonyuk 2017), Cherkasy Oblast, Dnipropetrovsk Oblast, Askania-Nova Nature Reserve, Striltsivskyi Steppe Nature Reserve, Kharkiv Oblast (Tokarsky and Zorya 2007), Lugansk Oblast (Kuznetsov and Bondarev 2007) (including Khomutovs’kyi Steppe) (Filippova 1977), the north-western sea coast of the Black Sea (Rusev 2009), Crimea (Evstafiev 2017) – plain and mountainous lands (Filippova 1977). Belarus: Viciebsk Voblasts (Subbotina and Osmolovsky 2022), Białowieża Forest (Filippova 1977), considered rare (Bychkova et al. 2015). Moldova: Lozova, Ivancea, Leova, reedbeds of the low Dniester and Pruth (Filippova 1977; Uspenskaya et al. 2006). Georgia: Samegrelo-Zemo Svaneti, Imereti (Sukhiashvili et al. 2020). Armenia: Aragats mountain range (Dilbaryan and Poghosyan 2018). Kazakhstan: through the whole territory of Kazakhstan (Filippova 1977) and plus recent findings in the next regions: West Kazakhstan Region (Tanitovsky and Maikanov 2018), Almaty Region (Bibikov and Bibikova 2010), Pavlodar Region (Amirova et al. 1989), the north of Betpak-Dala (Rapoport et al. 2017), Jambyl Region (Kyrgyz Ala-Too Range, Talas Alatau) (Sarsenbaeva et al. 2016). Kyrgyzstan: Tian Shan in general (Abdikarimov et al. 2018) and its certain ranges and valleys including Kyrgyz Ala-Too Range (Akyshova et al. 2022) and Terskey Ala-too Range (Fedorova 2012b); Chuy Valley (Fedorova 2021). Turkmenistan: Krasnovodsk Peninsula, Daşoguz, the foothills of The Köpet Dag, Badhyz State Nature Reserve, Karakum Desert (Kochkareva et al. 1971), Serhetabat (former Kushka) (Filippova 1977). Uzbekistan: Tashkent Region (Muratbekov 1954). Tajikistan: outskirts of the rural locality Jilikul (Filippova 1977), Tigrovaya Balka (Manilova and Shakhmatov 2008).

Figure 24. 

Map of Russia and neighboring countries showing the locations where Ixodes crenulatus was reported.

Ixodes crenulatus is among the tick species that have the most extensive ranges comparing to other representatives of its family within Russia (Tsapko 2020).

It is a typical nidicolous parasite of mammals and in the Asian part of its range as the main hosts it uses species of marmots of the genus Marmota (with a predominance of gray marmot) and such representatives of predatory mammals as badgers, steppe polecats, red and corsac foxes. The composition of the main host spectrum from different orders (rodents and predatory mammals) finds an explanation in close connections of topical and trophic relationships of marmots and predators. All of them have burrows of medium diameter, complex design, with a nesting chamber, remote from the entrance, which provides the stability of the microclimate, where ticks find suitable conditions. The above species of carnivores often use the burrows of their prey, marmots, and small carnivores, facilitating the exchange of ticks not only between individual burrows, but also between remote host settlements (Filippova 2011).

This tick species is considered rare, for example, only few findings were mentioned in the Astrakhan Oblast (Zimina et al. 1965; Zimina et al. 1996) and Saratov Oblast (Denisov 2019). In most of the recognized range, I. crenulatus coexists with the closely related I. kaiseri. These species not only inhabit the same territory and the same biotopes but can also parasitize one host individual at the same time (Tsapko 2017). Therefore, it is necessary to consider that accurate identification of these species is required and there is always a chance of their misidentification.

According to some suggestions (Emelyanova 1979), I. crenulatus is probably a species group, or at least has remarkable intraspecific variations involving morphotypes (Filippova and Panova 2000).

Mammalia: Bos taurus Linnaeus (cattle), Canis familiaris (domestic dog), Erinaceus europaeus (European hedgehog), Felis catus (domestic cat), Lutra lutra (Linnaeus) (Eurasian otter), Meles meles (Eurasian badger), Mustela erminea (stoat), Mustela nivalis (least weasel), Mustela putorius Linnaeus (European polecat), Oryctolagus cuniculus (Linnaeus) (European rabbit), Ovis aries (sheep), Rattus norvegicus (brown rat), Vulpes vulpes (red fox) (Filippova 1977).

Aves: Turdus merula (common blackbird) (Filippova 1977).

(Fig. 25). Ukraine: outskirts of Kyiv, Khmelnytskyi Oblast (Levytska et al. 2021), the North-Western seacoast of the Black Sea (Rusev 2009), Zakarpattia Oblast (the rural locality Malyi Berezny) (Filippova 1961).

Figure 25. 

Map of Russia and neighboring countries showing the locations where Ixodes hexagonus was reported.

Ixodes hexagonus is a typical nidicolous parasite of carnivores and hedgehogs. It has certain morphological similarities to I. crenulatus and I. kaiseri and has common sympatric zones with this species along its range (Filippova 1999). Ukraine is the only country of the former Soviet Union, on the territory of which this European species is present in the tick fauna (Filippova 1977). In general I. hexagonus was detected quite rarely in Ukraine, and almost always in the west of Ukraine and mainly from hedgehogs (Kolonin 2009). Akimov and Nebogatkin (2016) assumed that it can be found in the vicinity of Kyiv, and eventually it was confirmed by Levytska et al. (2021). Rare occasional human bites have been recorded (Rosický and Weiser 1952; Arthur 1963; Bursali et al. 2012).

Mammalia: Canis familiaris (domestic dog), Erinaceus concolor Martin (southern white-breasted hedgehog), Erinaceus europaeus (European hedgehog), Erinaceus roumanicus Barrett-Hamilton (northern white-breasted hedgehog), Felis chaus Schreber (jungle cat), Felis lybica (African wildcat), Hyaena hyaena (Linnaeus) (striped hyena), Hystrix indica Kerr (Indian crested porcupine), Lepus europaeus (brown hare), Mustela eversmanii (steppe polecat), Meles meles (Eurasian badger), Nyctereutes procyonoides (common raccoon dog), Vulpes corsac (corsac fox), Vulpes vulpes (red fox) (Filippova 1977; Tsapko 2017).

(Fig. 26). Russia: southwestern peripheries of the Central Russian Upland and also Rostov Oblast (Khametova et al. 2018) and Stavropol Krai (Tsapko 2017) and the North Caucasus – the outskirts of Grozny (Chechnya) (Filippova 1977, 1999; Tsapko 2017, 2020) and Nogaysky District (Dagestan) (Tsapko 2017). Ukraine: outskirts of Kyiv (Akimov and Nebogatkin 2016) and the south of Ukraine, in particular Askania-Nova Nature Reserve (Filippova 1977), the North-Western seacoast of the Black Sea (Matyukhin 2017), Crimea (Filippova 1977). Moldova: Lozova, Ivancea, Doibani, Leova, Etulia, reedbeds of the low Dniester and Pruth (Filippova 1977; Uspenskaya et al. 2006). Georgia: the outskirts of Tbilisi, Lagodekhi Nature Reserve (Filippova 1977), Eldari Steppe (Tsapko 2017). Armenia: Gegharkunik Province (rural locality Geghamashen) (Tsapko 2017), Aragats mountain range (Dilbaryan and Poghosyan 2018). Azerbaijan: Mil plain (Filippova and Uspenskaya 1973), Beylagan District, Zangilan District, Aghjabadi District, Martuni Province, Shaki District (Tsapko 2017). Kazakhstan: West Kazakhstan Region, Dzungarian Alatau – outskirts of the rural locality Topolëvka and the Koksu district (Ushakova et al. 1976; Filippova 1999). Kyrgyzstan (Fedorova 2012b).

Figure 26. 

Map of Russia and neighboring countries showing the locations where Ixodes kaiseri was reported.

Ixodes kaiseri is a typical nidicolous parasite of carnivores and also hedgehogs and porcupines which is morphologically very similar to I. crenulatus and has common sympatric zones with this species along its range (Filippova 1999). Its range itself is patchy and disjunctive areas of sympatry for both of these species are found in southeastern Europe - Romania, Moldova, Ukraine, including the Crimean Peninsula as well as in Russia - the southwestern extremities of the Central Russian Upland and the Northern Caucasus; then after a long gap - in Western Kazakhstan and again after a big gap - in the Dzungarian Alatau (Filippova 1999). Judging by literature, it is also known from Egypt and Israel (Arthur 1957, 1960, 1965). As Filippova and Uspenskaya (1973) assumed, its distribution in the Middle East and also other parts of Asia can be wider than it is known at present, which was already confirmed by the findings of this tick species in Turkey (Orkun and Karaer 2018) and Xinjiang in China (Zhao et al. 2019) near the border with Kazakhstan and the Dzungarian Alatau and hence, it is possible that the sympatry of these two species is more widespread. This is also supported by literature data, because until the 1970s in the territory of the former USSR I. kaiseri was not differentiated from I. hexagonus and I. crenulatus (Emelyanova 1979; Filippova 1999).

These species not only inhabit the same territory and inhabit the same biotopes in some places but can also parasitize one host individual at the same time (Tsapko 2017). It is also important to note that according to Filippova’s opinion (1999), the territorial signs of the ranges of these two species, their biotope and host-parasite relationships indicate that the range of I. crenulatus (which is predominantly connected with marmots and their burrows in steppe and forest-steppe zones) over the most part of its distribution has a Central Asian origin, while the range of I. kaiseri (mainly the parasite of carnivores and occurring in their burrows) is supposedly of European origin.

Aves: Alauda arvensis (Eurasian skylark), Delichon urbicum (Linnaeus) (common house martin), Merops apiaster Linnaeus (European bee-eater), Passer domesticus (house sparrow), Passer montanus (Eurasian tree sparrow), Riparia diluta (Sharpe & Wyatt) (pale martin), Riparia riparia (sand martin) (Filippova 1977; Tagiltsev et al. 1984; Rusev 2009; Bolshakova et al. 2019; Kovalevsky et al. 2019).

Mammalia: Mus musculus (house mouse) (Filippova 1977).

(Fig. 27). Russia: Curonian Spit (Kaliningrad Oblast) (Filippova 1961), Karelia (Bobrovskikh 1979), Leningrad Oblast, (Zolotov and Buker 1976), Moscow Oblast (Glashchinskaya-Babenko 1956), Ryazan Oblast (Filippova 1977), Ivanovo Oblast (Mayorova 2004), Saratov Oblast (Korneev et al. 2018), Kuybyshev Reservoir (Republic of Tatartstan) (Lvov et al. 2014), Voronezh Oblast (Gaponov and Tewelde 2021), Tymen Oblast (Starikov et al. 2017b), Kurgan Oblast (Starikov and Starikova 2021), Omsk Oblast (Tagiltsev et al. 1984; Yakimenko et al. 1991), Tomsk Oblast, Kemerovo Oblast (Kovalevsky et al. 2019), Novosibirsk Oblast (Yakimenko et al. 2013), Irkutsk Oblast (Danchinova et al. 2007), Ikatsky Ridge (Republic of Buryatia), Zabaykalsky Krai (Emelyanova et al. 1963), Republic of Tuva (Kholodilov et al. 2019), Sakha (Yakutia) (Shadrina et al. 2011), Khabarovsk Krai (Volkov and Chernykh 1977). Ukraine: Zakarpattia Oblast, Kaniv Nature Reserve (Cherkasy Oblast), (Emchuk 1960), Danube Biosphere Nature Reserve (Odesa Oblast) (Emchuk 1960; Didyk 2013), Kyiv (Akimov and Nebogatkin 2002), delta of the Dniestr River (Rusev 2009). Belarus: Gomel Region, Minsk Region (Gembetsky 1972). Moldova: banks of the Dniestr River (Movila et al. 2008). Kazakhstan: Atyrau Region (Pomerantsev 1950; Levit 1957), Kostanay Region (Makhmetov 1961), Pavlodar Region (Amirova et al. 1989), Akmola Region (Filippova 1961; Ushakova 1962).

Figure 27. 

Map of Russia and neighboring countries showing the locations where Ixodes lividus was reported.

Ixodes lividus is a specific nidicolous ectoparasite of the sand martin, Riparia riparia. Also, it has been collected from birds and house mice which occasionally could visit sand martin’s nests such as house sparrows and common house martins.

Due to the wide distribution of its main host, this tick species also occurs in a vast geographical range and can be characterized by having a trans-Palearctic distribution. The locations of findings in Russia and the neighboring countries listed above reflect the general pattern of its distribution on a map so we can suppose that this tick can be found in the north of the Palearctic almost everywhere in habitats of the sand martin.

Mammalia: Daurian hedgehog Mesechinus dauuricus (Sundevall) (Emelyanova 1979).

(Fig. 28). Russia: Transbaikal (Emelyanova 1979).

Figure 28. 

Map of Russia and neighboring countries showing the locations where Ixodes prokopjevi was reported.

Ixodes prokopjevi is an extremely poorly studied tick species initially described based on the male holotype from steppes of north-eastern Mongolia; its paratypes, larvae and nymphs, are noted as originating from the outskirts of the lake Baruun Shavart Nuur in Eastern Mongolia, as well as females and nymphs from the south-eastern Transbaikal without any indications of certain points of findings (Emelyanova 1979).

Kolonin (2009) states that this species should be considered a synonym of I. crenulatus but Guglielmone et al. (2010, 2014) recognize it as a valid species.

The Daurian hedgehog was recorded as a host, but we can assume that carnivores, lagomorphs, and rodents are also hosts of this tick species, as in case of I. crenulatus, another representative of the subgenus Pholeoixodes and the most similar species to this tick. The distribution area and ecology of I. prokopjevi, as well host-parasite relationships and their role in transmission of vector-borne infections remain unknown.

Aves: Athene noctua (little owl), Carduelis carduelis (European goldfinch), Coracias garrulus (European roller), Coturnix coturnix (common quail), Galerida cristata (crested lark), Falco naumanni Fleischer (lesser kestrel), Oenanthe hispanica (western black-eared wheatear), Oenanthe isabellina (isabelline wheatear), Oenanthe oenanthe (northern wheatear), Parus major (great tit), Passer ammodendri Gould (saxaul sparrow), Passer domesticus (house sparrow), Passer montanus (Eurasian tree sparrow), Pastor roseus (rosy starling), Petronia petronia (rock sparrow), Pica pica (Eurasian magpie), Sturnus vulgaris (common starling), Turdus ruficollis (red-throated thrush) (Filippova 1977).