A comprehensive survey of Rhinonyssid mites (Mesostigmata: Rhinonyssidae) in Northwest Russia: New mite-host associations and prevalence data

Biodiversity Data Journal 8: e49535 doi: 10.3897/BDJ.8.e49535 Data Paper A comprehensive survey of Rhinonyssid mites (Mesostigmata: Rhinonyssidae) in Northwest Russia: New mite-host associations and prevalence data Manuel De Rojas , Jorge Doña , Ivan Dimov ‡ University of Seville, Seville, Spain § Illinois Natural History Survey, Champaign, United States of America | State Pediatric Medical University, St. Petersburg, Russia Corresponding author: Manuel De Rojas ([email protected]), Jorge Doña ([email protected]) Academic editor: Jenő Kontschán Received: 19 Dec 2019 | Accepted: 26 Feb 2020 | Published: 28 Feb 2020 Citation: De Rojas M, Doña J, Dimov I (2020) A comprehensive survey of Rhinonyssid mites (Mesostigmata: Rhinonyssidae) in Northwest Russia: New mite-host associations and prevalence data. Biodiversity Data Journal 8: e49535. https://doi.org/10.3897/BDJ.8.e49535 Abstract Background Rhinonyssid mites are permanent parasites of birds that inhabit their respiratory tract. There are around 600 species described worldwide and almost all species of birds are found to have embedded rhinonyssid mites. Despite their presumed relevance, these mites are largely unstudied due to the difficulty in sampling them and, therefore, the majority of mite-host associations and species-prevalence data are unknown. New information In this study, 179 mite specimens belonging to 27 species and eight genera were identified. Notably, 18 new mite-bird associations were documented for the first time, thus increasing the known host range for these mite species. In addition, mite-host associations found in ‡ § | © De Rojas M et al. This is an open access article distributed under the terms of the Creative Commons Attribution License (CC BY 4.0), which permits unrestricted use, distribution, and reproduction in any medium, provided the original author and source are credited. this study were compared with known associations from these species of birds in the European part of Russia and in Europe. Overall, this study represents the largest survey to date carried out on rhinonyssid mites in Russia and one of the most comprehensive datasets on rhinonyssid host-range. Keywords birds, checklist, ectoparasites, parasites, symbionts Introduction Nasal mites of the family Rhinonyssidae are permanent haematophagous endoparasites of birds that inhabit their respiratory tract (Vitzthum 1935; George 1961; Fain 1994; Dimov and de Rojas 2012). Most species live in the nasal cavity on the vascularised epithelial tissue; nevertheless, some species occupy the lungs, tracheal tissues and body cavity of their hosts (Lindquist et al. 2009; Krantz and Walter 2009). Rhinonyssids can not only cause damage to their hosts in a direct way (Rhinonyssidosis avium disease) (Dimov 2011), but could also be reservoirs or vectors of other infections like West Nile fever, Q fever, avian influenza and Lyme disease, as have been shown in mites from the family Dermanyssidae (Reeves et al. 2006). Despite their ecological relevance, most aspects of the basic biology, ecology and evolution of these mites are still poorly understood. This lack of knowledge, amongst other reasons, is owing to their being very challenging to study; for example, due to most species having typically low prevalence on their hosts and being difficult to sample (being only possible to collect from dead birds). The family Rhinonyssidae currently includes about 600 described species arranged in eleven genera (Domrow 1969; Fain 1994; Dimov et al. 2015; Dimov 2018). In Parasitology, parasite host-specificity and prevalence are widely-studied parameters as they are informative of relevant processes, such as parasite degree of specialisation, population dynamics or transmission efficiency (Poulin 2011). Host specificity of rhinonyssid mites has been found to vary from one genus to another by surveys across different geographic areas (e.g. USA, Spicer 1987; Canada, Knee et al. 2008). In particular, some genera have been found to be constrained to a single host family, while others can inhabit hosts from different orders (Pence 1975; Butenko 1984). In addition, these studies have found that the prevalence of these mites varies across geographic areas (although the estimates may be biased by a low sample size; Spicer 1987). However, the host-specificity and prevalence of most rhinonyssid mites are still poorly understood, thus hampering further studies on the ecology and evolution of this host-parasite system. In this study, 2,107 bird specimens from northwest of Russia, belonging to 75 species from 55 genera, 30 families and 10 orders were examined for rhinonyssid mites. The mites were identified and the prevalence (including confidence intervals to show how accurate the estimates are) of these species was calculated. Additionally, the mite-host associations, found in our study, were compared with the known rhinonyssids from these species of birds 2De Rojas M et al in the European part of Russia and Europe. Overall, this study represents the largest survey to date carried out on rhinonyssid mites in Russia and one of the most comprehensive datasets on rhinonyssid host-range and prevalence. General description Purpose: In this survey, 2,107 individual birds were collected in Russia representing 75 species belonging to 55 genera, 30 families and 10 orders and analysed for rhinonyssid mites. The mites were identified and the prevalence (including 95% confidence intervals to show how accurate the estimates are) of these species was calculated. Additionally, the mite-host associations found in our study were compared with the known rhinonyssids from these species of birds in the European part of Russia and Europe. Sampling methods Study extent: Individual birds were collected during four years (2010-2013). Sampling description: Mites were collected from birds that died under various circumstances. Specifically, most of the surveyed hosts were birds found dead on the roads or that died because of high-voltage transmission lines. Host birds were morphologically classified according to Malchevsky and Pukinsky 1983. All the birds were examined for rhinonyssid mites and when they were found, a complete morphometrical study was conducted to identify each specimen (Butenko 1984; Pence 1975; Fain 1956; Dimov and de Rojas 2012; Dimov and Knee 2012; Dimov and Mironov 2012). Quality control: The nasal cavity of birds was opened following Butenko's method (Butenko 1984), with some changes introduced by us. In particular, 1) the nasal cavity was opened with a scalpel and scissors under a binocular stereomicroscope. 2) The ossa mandibulae were removed along with the hyoid brush apparatus (apparatus hyobranchialis); the eyeballs were removed with tweezers. 3) Then, two incisions were made: a transverse incision in the region of the papillae pharyngeales and a medial incision, from the rima infundibuli (through the choana) to the ruga palatina mediana area. 4) Nostrils in the area of operculum were examined and then the os maxillare were opened. 5) Lastly, the maxillary bone was removed and the nasal cavity with three conchs was opened, including the largest of all rostral - concha nasalis rostralis, the middle - concha nasalis media and the caudal - concha nasalis caudalis. The examination of all nasal cavities was performed with tweezers and a dissecting needle under a binocular stereomicroscope. Rhinonyssid mites were placed in tubes with 70% ethanol for storage. Each tube was labelled with data on the type of host and a detailed description of the collection site. Mites were then cleared in lactic acid and mounted on slides with Fora- Berlese liquid, according to the generally-accepted technique for small mites (Walter and Krantz 2009; Krantz and Walter 2009). Finally, individual mites were identified, based on morphometrics analysis. A comprehensive survey of Rhinonyssid mites (Mesostigmata: Rhinonyssidae) ... 3 Geographic coverage Description: Individual birds were collected across the northwest of the European part of Russia, mainly in the territory of the Leningrad Region, at 41 points and, to a lesser extent, in territories of Arkhangelsk, Kaliningrad and Pskov regions (Suppl. material 1; Table 2). Bird order Bird families Bird genera Bird species Analysed individuals Infected individuals Prevalence Anseriformes 1 3 4 105 13 12.38% (7.38-20.04) Caprimulgiformes 1 1 1 2 1 50% (2.56-97.44) Charadriiformes 4 5 6 117 8 6.8% (3.51-12.91) Columbiformes 1 1 1 262 10 3.8% (2.09-6.88) Cuculiformes 1 1 1 13 1 7.69% (0.39-33.31) Accipitriformes 1 2 2 3 0 0% (0-56.15) Galliformes 1 2 2 32 0 0% (0-10.71) Gruiformes 1 2 2 8 2 25% (4.44-59.07) Passeriformes 18 37 55 1549 58 3.7% (2.91-4.81) Piciformes 1 1 1 16 0 0% (0-19.36) Total 30 55 75 2107 93 4.41% (3.62-5.38) Bird taxa Mite species N Ni I Region (previous records) Locality (this study) Previous records Accipiter nisus (Linnaeus, 1758) NA 2 0 0 NA NA NA Table 1. Data of prevalence of rhinonyssid mites in different orders of hosts and the number of families, genera and species of birds studied. Confidence intervals of prevalence (95%) are provided between parentheses. Table 2. Data on the mite-host associations detected. Each row depicts an individual bird from mite-host associations, in which more than one mite was found. N = number of birds examined per mite-host association, Ni = Number of infected birds, I = number of mites isolated. Region (previous records) = NR: Northwest Russia, ER: European part of Russia, WE: Western Europe and * indicates a previously unrecorded mite host association. Note that values from N and Ni columns are duplicated between individual birds that belong to the same mite-host association. 4De Rojas M et al Bird taxa Mite species N Ni I Region (previous records) Locality (this study) Previous records Buteo buteo (Linnaeus, 1758) NA 1 0 0 NA NA NA Anas platyrhynchos Linnaeus, 1758 Rhinonyssus kadrae Dimov, 2013 38 1 2 NR* Dubrovka, Leningrad Oblast, Russia NA Anas platyrhynchos Linnaeus, 1758 Rhinonyssus levinseni (Tragardh, 1904) 38 1 1 NR* Volosovo, Leningrad Region, Russia NA Aythya nyroca (Güldenstädt, 1770) Rhinonyssus nyrocae Butenko, 1971 1 1 1 NR, WE Pskov, Pskov region, Russia Butenko 1971, Butenko 1984 Somateria mollissima Linnaeus, 1758 Rhinonyssus polystictae Butenko, 1984 11 9 1 NR* Ermilovo, Leningrad Region, Russia NA Somateria mollissima Linnaeus, 1758 Rhinonyssus polystictae Butenko, 1984 11 9 1 NR* Ermilovo, Leningrad Region, Russia NA Somateria mollissima Linnaeus, 1758 Rhinonyssus polystictae Butenko, 1984 11 9 2 NR* Ermilovo, Leningrad Region, Russia NA Somateria mollissima Linnaeus, 1758 Rhinonyssus polystictae Butenko, 1984 11 9 1 NR* Ermilovo, Leningrad Region, Russia NA Somateria mollissima Linnaeus, 1758 Rhinonyssus polystictae Butenko, 1984 11 9 2 NR* Ermilovo, Leningrad Region, Russia NA Somateria mollissima Linnaeus, 1758 Rhinonyssus polystictae Butenko, 1984 11 9 1 NR* Ermilovo, Leningrad Region, Russia NA Somateria mollissima Linnaeus, 1758 Rhinonyssus polystictae Butenko, 1984 11 9 2 NR* Ermilovo, Leningrad Region, Russia NA Somateria mollissima Linnaeus, 1758 Rhinonyssus polystictae Butenko, 1984 11 9 1 NR* Ermilovo, Leningrad Region, Russia NA A comprehensive survey of Rhinonyssid mites (Mesostigmata: Rhinonyssidae) ... 5 Bird taxa Mite species N Ni I Region (previous records) Locality (this study) Previous records Somateria mollissima Linnaeus, 1758 Rhinonyssus polystictae Butenko, 1984 11 9 1 NR* Ermilovo, Leningrad Region, Russia NA Anas crecca Linnaeus, 1758 Rhinonyssus subrhinolethrum Butenko, 1971 17 1 1 NR Pikalevo, Leningrad Region, Russia Butenko 1971, Butenko 1984 Caprimulgus europeus Linnaeus, 1758 Vitznyssus tsachevi Dimov et Rojas 2012 2 1 2 NR* Pikalevo, Leningrad region, Russia NA Charadrius dubius Scopoli, 1786 Rhinonyssus bregetovae Butenko, 1974 27 3 4 NR Voypala, Leningrad Region, Russia Butenko 1984 Charadrius dubius Scopoli, 1786 Rhinonyssus bregetovae Butenko, 1974 27 3 2 NR Voypala, Leningrad Region, Russia Butenko 1984 Charadrius dubius Scopoli, 1786 Rhinonyssus bregetovae Butenko, 1974 27 3 1 NR Voypala, Leningrad Region, Russia Butenko 1984 Charadrius dubius Scopoli, 1786 Rhinonyssus neglectus Hirst 1921 27 1 1 NR* Lavrovo, Leningrad Region, Russia NA Vanellus vanellus Linnaeus, 1758 Rhinonyssus dobromiri Dimov et Spicer, 2013 4 1 2 NR* Leningrad Region, Russia NA Larus argentatus Pontoppidan, 1763 Larinyssus orbicularis Strandtmann, 1948 22 1 2 NR* Voybokalo, Leningrad Oblast, Russia NA Larus ridibundus Linnaeus, 1766 NA 19 0 0 NA NA NA Scolopax rusticola Linnaeus, 1758 NA 1 0 0 NA NA NA 6De Rojas M et al Bird taxa Mite species N Ni I Region (previous records) Locality (this study) Previous records Sterna hirundo Linnaeus, 1758 Larinyssus iohanssenae Dimov, 2013 17 2 1 NR* Kronstad, Leningrad Region, Russia; Apraksin, Leningrad Region, Russia NA Sterna hirundo Linnaeus, 1758 Larinyssus iohanssenae Dimov, 2013 17 2 2 NR* Kronstad, Leningrad Region, Russia; Apraksin, Leningrad Region, Russia NA Columba livia Gmelin, 1789 Mesonyssus columbae Crossley, 1950 262 5 3 NR, WE St. Petersburg, Russia; Voybokalo, Leningrad Oblast, Russia; Slantsy, Leningrad Region, Russia; Luga, Leningradskaya, Russia; Radogosh, Leningrad Region, Russia Butenko 1984; Cerny 1970; Crossley 1951; Domrow 1965; Domrow 1966a; Domrow 1966b; Fain 1956; Fain 1957; Fain 1958; Fain 1962b; Fain et al. 1974; Pence 1975; Sixl 1971; Wilson 1964; Wilson 1966; Zumpt and Till 1955 Columba livia Gmelin, 1789 Mesonyssus columbae Crossley, 1950 262 5 1 NR, WE St. Petersburg, Russia; Voybokalo, Leningrad Oblast, Russia; Slantsy, Leningrad Region, Russia; Luga, Leningradskaya, Russia; Radogosh, Leningrad Region, Russia Butenko 1984; Cerny 1970; Crossley 1951; Domrow 1965; Domrow 1966a; Domrow 1966b; Fain 1956; Fain 1957; Fain 1958; Fain 1962b; Fain et al. 1974; Pence 1975; Sixl 1971; Wilson 1964; Wilson 1966; Zumpt and Till 1955 Columba livia Gmelin, 1789 Mesonyssus columbae Crossley, 1950 262 5 2 NR, WE St. Petersburg, Russia; Voybokalo, Leningrad Oblast, Russia; Slantsy, Leningrad Region, Russia; Luga, Leningradskaya, Russia; Radogosh, Leningrad Region, Russia Butenko 1984; Cerny 1970; Crossley 1951; Domrow 1965; Domrow 1966a; Domrow 1966b; Fain 1956; Fain 1957; Fain 1958; Fain 1962b; Fain et al. 1974; Pence 1975; Sixl 1971; Wilson 1964; Wilson 1966; Zumpt and Till 1955 A comprehensive survey of Rhinonyssid mites (Mesostigmata: Rhinonyssidae) ... 7 Bird taxa Mite species N Ni I Region (previous records) Locality (this study) Previous records Columba livia Gmelin, 1789 Mesonyssus columbae Crossley, 1950 262 5 2 NR, WE St. Petersburg, Russia; Voybokalo, Leningrad Oblast, Russia; Slantsy, Leningrad Region, Russia; Luga, Leningradskaya, Russia; Radogosh, Leningrad Region, Russia Butenko 1984; Cerny 1970; Crossley 1951; Domrow 1965; Domrow 1966a; Domrow 1966b; Fain 1956; Fain 1957; Fain 1958; Fain 1962b; Fain et al. 1974; Pence 1975; Sixl 1971; Wilson 1964; Wilson 1966; Zumpt and Till 1955 Columba livia Gmelin, 1789 Mesonyssus columbae Crossley, 1950 262 5 3 NR, WE St. Petersburg, Russia; Voybokalo, Leningrad Oblast, Russia; Slantsy, Leningrad Region, Russia; Luga, Leningradskaya, Russia; Radogosh, Leningrad Region, Russia Butenko 1984; Cerny 1970; Crossley 1951; Domrow 1965; Domrow 1966a; Domrow 1966b; Fain 1956; Fain 1957; Fain 1958; Fain 1962b; Fain et al. 1974; Pence 1975; Sixl 1971; Wilson 1964; Wilson 1966; Zumpt and Till 1955 Columba livia Gmelin, 1789 Mesonyssus melloi Castro, 1948 262 3 4 NR, WE St. Petersburg, Russia; Voybokalo, Leningrad Oblast, Russia; Slantsy, Leningrad Region, Russia. Butenko 1984; Castro 1948; Domrow 1966a; Domrow 1969; Domrow 1972a; Domrow 1972b; Fain 1959; Fain 1962b; Pence 1979; Sixl 1969; Wilson 1964; Wilson 1966; Zumpt and Till 1955 Columba livia Gmelin, 1789 Mesonyssus melloi Castro, 1948 262 3 4 NR, WE St. Petersburg, Russia; Voybokalo, Leningrad Oblast, Russia; Slantsy, Leningrad Region, Russia. Butenko 1984; Castro 1948; Domrow 1966a; Domrow 1969; Domrow 1972a; Domrow 1972b; Fain 1959; Fain 1962b; Pence 1979; Sixl 1969; Wilson 1964; Wilson 1966; Zumpt and Till 1955 8De Rojas M et al Bird taxa Mite species N Ni I Region (previous records) Locality (this study) Previous records Columba livia Gmelin, 1789 Mesonyssus melloi Castro, 1948 262 3 2 NR, WE St. Petersburg, Russia; Voybokalo, Leningrad Oblast, Russia; Slantsy, Leningrad Region, Russia. Butenko 1984; Castro 1948; Domrow 1966a; Domrow 1969; Domrow 1972a; Domrow 1972b; Fain 1959; Fain 1962b; Pence 1979; Sixl 1969; Wilson 1964; Wilson 1966; Zumpt and Till 1955 Cuculus canorus Linnaeus, 1758 Sternostoma zini Dimov et Knee, 2012 13 1 2 NR* Vyritsa, Leningrad Region, Russia NA Coturnix coturnix Linnaeus, 1758 NA 31 0 0 NA NA NA Tetrao urogallus Linnaeus, 1758 NA 1 0 0 NA NA NA Gallinula chloropus Linnaeus, 1758 Rallinyssus caudistigmus Strandtmann, 1948 6 1 3 NR, WE Gatchina, Leningrad Region, Russia; Sosnovy Bor, Leningrad Region, Russia Domrow 1966a; Domrow 1969; Fain 1957; Fain et al. 1974; Pence 1975; Strandtmann 1948 Fulica atra Linnaeus, 1758 Rallinyssus caudistigmus Strandtmann, 1948 2 1 2 NR, ER, WE Gatchina, Leningrad Region, Russia; Sosnovy Bor, Leningrad Region, Russia Bregetova 1951; Butenko 1984; Domrow 1966a; Domrow 1969; Fain 1957; Fain 1959; Fain et al. 1974; Pence 1972d; Pence 1975; Strandtmann 1948 Alauda arvensis Linnaeus, 1758 Ptilonyssus schumili Butenko et Lavrovskaya, 1980 35 6 2 NR, ER Rjbachii, Leningrad Region, Russia Butenko and Lavroskaya 1980a; Butenko and Lavroskaya 1980b Alauda arvensis Linnaeus, 1758 Ptilonyssus schumili Butenko et Lavrovskaya, 1980 35 6 1 NR, ER Rjbachii, Leningrad Region, Russia Butenko and Lavroskaya 1980a; Butenko and Lavroskaya 1980b A comprehensive survey of Rhinonyssid mites (Mesostigmata: Rhinonyssidae) ... 9 Bird taxa Mite species N Ni I Region (previous records) Locality (this study) Previous records Parus major Linnaeus, 1758 Ptilonyssus pari Fain et Hyland 1963 118 9 1 NR, WE St. Petersburg, Russia Fain and Bafort 1963; Fain and Hyland 1963; Fain et al. 1974; Kadosaka et al. 1983; Pence 1972a; Pence 1972b; Pence 1975; Pence and Casto 1976; Shumilo and Lunkashu 1970; Sixl 1969; Sixl 1970; Spicer 1977; Spicer 1978 Parus major Linnaeus, 1758 Ptilonyssus pari Fain et Hyland 1963 118 9 1 NR, WE St. Petersburg, Russia Fain and Bafort 1963; Fain and Hyland 1963; Fain et al. 1974; Kadosaka et al. 1983; Pence 1972a; Pence 1972b; Pence 1975; Pence and Casto 1976; Shumilo and Lunkashu 1970; Sixl 1969; Sixl 1970; Spicer 1977; Spicer 1978 Parus major Linnaeus, 1758 Ptilonyssus pari Fain et Hyland 1963 118 9 2 NR, WE St. Petersburg, Russia Fain and Bafort 1963; Fain and Hyland 1963; Fain et al. 1974; Kadosaka et al. 1983; Pence 1972a; Pence 1972b; Pence 1975; Pence and Casto 1976; Shumilo and Lunkashu 1970; Sixl 1969; Sixl 1970; Spicer 1977; Spicer 1978 16 De Rojas M et al Bird taxa Mite species N Ni I Region (previous records) Locality (this study) Previous records Parus major Linnaeus, 1758 Ptilonyssus pari Fain et Hyland 1963 118 9 1 NR, WE St. Petersburg, Russia Fain and Bafort 1963; Fain and Hyland 1963; Fain et al. 1974; Kadosaka et al. 1983; Pence 1972a; Pence 1972b; Pence 1975; Pence and Casto 1976; Shumilo and Lunkashu 1970; Sixl 1969; Sixl 1970; Spicer 1977; Spicer 1978 Parus ater Linnaeus, 1758 NA 2 0 0 NA NA NA Parus cristatus Linnaeus, 1758 NA 9 0 0 NA NA NA Parus montanus (Conrad von Baldenstein, 1827) NA 29 0 0 NA NA NA Parus palustris Linnaeus, 1758 NA 6 0 0 NA NA NA Passer domesticus Linnaeus, 1758 Ptilonyssus degtiarevae Dimov et Mironov, 2012 74 1 5 NR* Gavrilovo, Leningradskaya Region, Russia NA Passer montanus Linnaeus, 1758 Ptilonyssus lovottiae Dimov et Mironov, 2012 91 3 18 NR* Leningrad Region Russia; Boronichevo, Leningrad Region, Russia; Novaya LadogaLeningrad Oblast, Russia NA Passer montanus Linnaeus, 1758 Ptilonyssus lovottiae Dimov et Mironov, 2012 91 3 3 NR* Leningrad Region Russia; Boronichevo, Leningrad Region, Russia; Novaya LadogaLeningrad Oblast, Russia NA A comprehensive survey of Rhinonyssid mites (Mesostigmata: Rhinonyssidae) ... 17 Bird taxa Mite species N Ni I Region (previous records) Locality (this study) Previous records Passer montanus Linnaeus, 1758 Ptilonyssus lovottiae Dimov et Mironov, 2012 91 3 5 NR* Leningrad Region Russia; Boronichevo, Leningrad Region, Russia; Novaya LadogaLeningrad Oblast, Russia NA Regulus regulus Linnaeus, 1758 NA 27 0 0 NA NA NA Sitta europea Linnaeus, 1758 NA 3 0 0 NA NA NA Phylloscopus sibilatrix (Bechstein, 1793) NA 1 0 0 NA NA NA Phylloscopus trochilus Linnaeus, 1758 NA 16 0 0 NA NA NA Dendrocopos major Linnaeus, 1758 NA 16 0 0 NA NA NA Sturnus vulgaris Linnaeus, 1758 NA 39 0 0 NA NA NA Sylvia atricapilla Linnaeus, 1758 NA 7 0 0 NA NA NA Sylvia borin (Boddaert, 1783) NA 3 0 0 NA NA NA Sylvia curruca Linnaeus, 1758 NA 41 0 0 NA NA NA Turdus iliacus Linnaeus, 1766 Ptilonyssus euroturdi Fain et Hyland, 1963 27 7 1 NR* Arkhangelsk Region, Russia NA 18 De Rojas M et al Bird taxa Mite species N Ni I Region (previous records) Locality (this study) Previous records Turdus iliacus Linnaeus, 1766 Ptilonyssus euroturdi Fain et Hyland, 1963 27 7 2 NR* Arkhangelsk Region, Russia NA Turdus iliacus Linnaeus, 1766 Ptilonyssus euroturdi Fain et Hyland, 1963 27 7 1 NR* Arkhangelsk Region, Russia NA Turdus iliacus Linnaeus, 1766 Ptilonyssus euroturdi Fain et Hyland, 1963 27 7 3 NR* Arkhangelsk Region, Russia NA Turdus iliacus Linnaeus, 1766 Ptilonyssus euroturdi Fain et Hyland, 1963 27 7 2 NR* Arkhangelsk Region, Russia NA Turdus iliacus Linnaeus, 1766 Ptilonyssus euroturdi Fain et Hyland, 1963 27 7 2 NR* Arkhangelsk Region, Russia NA Turdus iliacus Linnaeus, 1766 Ptilonyssus euroturdi Fain et Hyland, 1963 27 7 1 NR* Arkhangelsk Region, Russia NA Turdus merula Linnaeus, 1758 Sternostoma dureni Fain, 1956 78 1 1 NR* Hervir, Leningrad Region, Russia NA Turdus philomelos Brehm, 1831 Sternostoma turdi Zumpt et Till, 1955 19 7 2 NR, ER WE Severodvinsk, Arkhangelsk, Russia Butenko 1965; Fain 1956; Fain 1957; Fain 1959; Fain 1962a; Fain 1963; Fain and Aikten 1967; Fain et al. 1974; Furman 1957; Pence 1972c; Shumilo and Lunkashu 1970; Sixl 1971; Spicer 1984; Spicer 1987; Zumpt and Till 1955 A comprehensive survey of Rhinonyssid mites (Mesostigmata: Rhinonyssidae) ... 19 Bird taxa Mite species N Ni I Region (previous records) Locality (this study) Previous records Turdus philomelos Brehm, 1831 Sternostoma turdi Zumpt et Till, 1955 19 7 2 NR, ER WE Severodvinsk, Arkhangelsk, Russia Butenko 1965; Fain 1956; Fain 1957; Fain 1959; Fain 1962a; Fain 1963; Fain and Aikten 1967; Fain et al. 1974; Furman 1957; Pence 1972c; Shumilo and Lunkashu 1970; Sixl 1971; Spicer 1984; Spicer 1987; Zumpt and Till 1955 Turdus philomelos Brehm, 1831 Sternostoma turdi Zumpt et Till, 1955 19 7 1 NR, ER WE Severodvinsk, Arkhangelsk, Russia Butenko 1965; Fain 1956; Fain 1957; Fain 1959; Fain 1962a; Fain 1963; Fain and Aikten 1967; Fain et al. 1974; Furman 1957; Pence 1972c; Shumilo and Lunkashu 1970; Sixl 1971; Spicer 1984; Spicer 1987; Zumpt and Till 1955 Turdus philomelos Brehm, 1831 Sternostoma turdi Zumpt et Till, 1955 19 7 1 NR, ER WE Severodvinsk, Arkhangelsk, Russia Butenko 1965; Fain 1956; Fain 1957; Fain 1959; Fain 1962a; Fain 1963; Fain and Aikten 1967; Fain et al. 1974; Furman 1957; Pence 1972c; Shumilo and Lunkashu 1970; Sixl 1971; Spicer 1984; Spicer 1987; Zumpt and Till 1955 20 De Rojas M et al Bird taxa Mite species N Ni I Region (previous records) Locality (this study) Previous records Turdus philomelos Brehm, 1831 Sternostoma turdi Zumpt et Till, 1955 19 7 1 NR, ER WE Severodvinsk, Arkhangelsk, Russia Butenko 1965; Fain 1956; Fain 1957; Fain 1959; Fain 1962a; Fain 1963; Fain and Aikten 1967; Fain et al. 1974; Furman 1957; Pence 1972c; Shumilo and Lunkashu 1970; Sixl 1971; Spicer 1984; Spicer 1987; Zumpt and Till 1955 Turdus philomelos Brehm, 1831 Sternostoma turdi Zumpt et Till, 1955 19 7 1 NR, ER WE Severodvinsk, Arkhangelsk, Russia Butenko 1965; Fain 1956; Fain 1957; Fain 1959; Fain 1962a; Fain 1963; Fain and Aikten 1967; Fain et al. 1974; Furman 1957; Pence 1972c; Shumilo and Lunkashu 1970; Sixl 1971; Spicer 1984; Spicer 1987; Zumpt and Till 1955 Turdus philomelos Brehm, 1831 Sternostoma turdi Zumpt et Till, 1955 19 7 1 NR, ER WE Severodvinsk, Arkhangelsk, Russia Butenko 1965; Fain 1956; Fain 1957; Fain 1959; Fain 1962a; Fain 1963; Fain and Aikten 1967; Fain et al. 1974; Furman 1957; Pence 1972c; Shumilo and Lunkashu 1970; Sixl 1971; Spicer 1984; Spicer 1987; Zumpt and Till 1955 Turdus pilaris Linnaeus, 1758 NA 121 0 0 NA NA NA Dendrocopos major Linnaeus, 1758 NA 16 0 0 NA NA NA Coordinates: 60.05; 31.75. A comprehensive survey of Rhinonyssid mites (Mesostigmata: Rhinonyssidae) ... 21 Taxonomic coverage Description: A total of 2,107 individual birds were collected, representing 75 species belonging to 55 genera, 30 families and 10 orders. The majority of the studied hosts (59 species from 36 genera) belonged to the order Passeriformes (the most numerous and widely-distributed order of birds of the northwest of Russia) (Table 1). Specifically, sampled passerine species comprise up to 58% out of the total number of species of passerine species inhabiting the territory of the northwest of Russia (Malchevsky and Pukinsky 1983). In general, 179 mite specimens belonging to 27 species and 8 genera were identified. From a total of 27 host-mite associations, we report 18 novel host-mite associations (Table 2). As expected, due to the higher sampling effort in Passerifoms, the number of new hostmite associations was the highest in this order (9 out of 18), followed by waterbirds belonging to Charadriiformes and Anseriformes (with 4 and 3 new host-mite associations, respectively). Only a single new host-mite association was detected in Caprimulgiformes and Cuculiformes. Lastly, no new association was found in Columbiformes, Galliformes or Piciformes. Taxa included: Rank Scientific Name Common Name kingdom Animalia Animals subkingdom Eumetazoa phylum Chordata subphylum Vertebrata class Aves Birds subclass Galloanserae subclass Passerae superorder Anserimorphae superorder Columbimorphae superorder Cuculimorphae superorder Gallomorphae superorder Passerimorphae order Accipitriformes order Anseriformes order Caprimulgiformes order Charadriiformes order Columbiformes 22 De Rojas M et al order Cuculiformes order Galliformes order Gruiformes order Passeriformes suborder Accipitri suborder Anseri suborder Caprimulgi suborder Charadrii suborder Columbi suborder Cuculi suborder Passeri suborder Phasiani suborder Ralli superfamily Accipitroidea superfamily Aegithaloidea superfamily Alaudoidea superfamily Anatoidea superfamily Bombycilloidea superfamily Caprimulgoidea superfamily Charadrioidea superfamily Columboidea superfamily Corvoidea superfamily Cuculoidea superfamily Fringilloidea superfamily Gruoidea superfamily Hirundinoidea superfamily Laroidea superfamily Muscicapoidea superfamily Passeroidea superfamily Phasianoidea superfamily Ralloidea A comprehensive survey of Rhinonyssid mites (Mesostigmata: Rhinonyssidae) ... 23 superfamily Reguloidea superfamily Scolopacoidea superfamily Sittoidea superfamily Sturnoidea superfamily Sylvioidea family Accipitridae family Anatidae family Caprimulgidae family Charadriidae family Columbidae family Corvidae family Cuculidae family Emberizidae family Fringillidae family Laridae family Motacillidae family Muscicapidae family Paridae family Ploceidae family Rallidae family Regulidae family Scolopacidae family Sturnidae family Sylviidae family Turdidae species Acanthis canabina species Acanthis flammea species Aegithalos caudatus species Accipiter nisus species Alauda arvensis species Anas crecca 24 De Rojas M et al species Anas platyrhynchos species Anthus pratensis species Anthus trivialis species Aythya nyroca species Bombycilla garrulus species Buteo buteo species Caprimulgus europeus species Carduelis carduelis species Carpodacus erythrinus species Charadrius dubius species Chloris chloris species Columba livia species Corvus cornix species Coturnix coturnix species Cuculus canorus species Delichon urbica species Dendrocopos major species Emberiza citrinella species Emberiza hortulana species Emberiza rustica species Emberiza schoeniclus species Ficedula hypoleuca species Fringilla coelebs species Fringilla montifringilla species Fulica atra species Gallinula chloropus species Garrulus glandarius species Hirundo rustica species Larus ridibundus species Loxia curvirostra species Luscinia luscinia A comprehensive survey of Rhinonyssid mites (Mesostigmata: Rhinonyssidae) ... 25 On the other hand, some species that have been found in the European part of Russia and Europe were not found in this study (e.g. Mesonyssus hirsutus from Columba livia; Ptilonyssus nudus from Fringilla coelebs; Ptilonyssus pari from Parus ater; Parus caeruleus or Parus europea; Rhinonyssus vanellus from Vanellus vanellus). Overall, these differences in diversity could be the outcome of the lack of knowledge about these mites in these regions. Indeed, differences in sampling effort (i.e. some groups have been more extensively sampled in Northwest Russia than in other geographic areas) may be biasing these interpretations. Overall, further studies aimed at ascertaining whether specific rhinonyssid mite species are found throughout all their host distribution are encouraged. In this vein, global syntheses are needed to draw more general conclusions on the distribution of rhinonyssid mites. Additionally, future studies describing new species are required to catalogue the unknown diversity of this group of mites. In addition, new molecular approaches (e.g. DNA metabarcoding of complex samples; Doña et al. 2019) would help to accelerate the discovery of new species along with validating the species status of previously-described rhinonyssid species, as cryptic species are known in this group (de Rojas et al. 2018). Acknowledgements The authors are very grateful to the head of the laboratory of parasitology ZIN RAS Medvedev DSc. and employees of the laboratory of parasitology: Mironov DSc., Shatrov DSc., Bochkov DSc. and Leonovich DSc. for the comprehensive assistance and support provided in the process of carrying out the research. We thank Centro Municipal Zoosanitario de Sevilla and especially Mr. Francisco Peña Fernández and Mr. Rafael Cuadrado Nieto for their technical support. Our thanks to Geoffrey Giddings for proofreading the manuscript. Author contributions MDR, JD, and ID conceived the study. ID collected the samples. MR and ID made the morphobiometric assessments. 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Journal of the Entomological Society of South Africa 18: 60‑92. Supplementary materials Suppl. material 1: Figure S1 Authors: de Rojas M, Doña J, Dimov I Data type: Figure Brief description: Map of sampling localities. Note that specific locations, as well as their coordinates, can be found in Table 2. Download file (390.42 kb) 36 De Rojas M et al Suppl. material 2: Table S1 Authors: de Rojas M, Doña J, Dimov I Data type: Table Brief description: Comparison of species of the family Rhinonyssidae located in Northwest Russia, the European part of Russia and Western Europe; plus and minus signs indicate presence and absence, respectively. Mite species found in this study are marked with * Download file (6.31 kb) Suppl. material 3: Table S2 Authors: de Rojas, M., Doña, J., Dimov, I Data type: Table Brief description: Rhinonyssid mites from Russian birds Download file (15.10 kb) A comprehensive survey of Rhinonyssid mites (Mesostigmata: Rhinonyssidae) ... 37