Orlova, Maria V. ¹ ; Halliday, Bruce ² ; Reeves, Will K. ³ ; Doronin, Igor V. ⁴ ; Mishchenko, Vladimir A. ⁵ ; Vyalykh, Ivan V. ⁶ and Kidov, Artem A. ⁷

¹✉ Tyumen State Medical University, Tyumen, Russia & National Research Tomsk State University, Tomsk, Russia & Federal Scientific Research Institute of Viral Infections «Virome» of Federal Service for Surveillance on Consumer Rights Protection and Human Wellbeing, Ekaterinburg, Russia.
²Australian National Insect Collection, CSIRO, Canberra, Australia.
³C. P. Gillette Museum of Arthropod Diversity, Fort Collins, USA.
⁴Zoological Institute of Russian Academy of Science, Saint Petersburg, Russia.
⁵Federal Scientific Research Institute of Viral Infections Virome of Federal Service for Surveillance on Consumer Rights Protection and Human Wellbeing, Ekaterinburg, Russia.
⁶Federal Scientific Research Institute of Viral Infections Virome of Federal Service for Surveillance on Consumer Rights Protection and Human Wellbeing, Ekaterinburg, Russia.
⁷Russian State Agrarian University – Moscow Agricultural Academy named after. K. A. Timiryazev, Moscow, Russia.

2024 - Volume: 64 Issue: 2 pages: 637-653

Original research

Keywords

Abstract

Introduction

Mites in the family Macronyssidae (Acari: Mesostigmata: Gamasina) are mostly obligate blood-sucking ectoparasites of mammals, birds and reptiles. The family currently includes 34 genera with approximately 240 species (Radovsky, 2010). The genus Ophionyssus Mégnin, 1884a currently includes 14 species worldwide (Beron, 2014). The most extensively studied species in the genus is the snake mite Ophionyssus natricis (Gervais, 1844), which naturally infests snakes and lizards in Africa (Till, 1957; Evans & Till, 1966), but in other parts of the world it is associated primarily with animals in captivity (Miranda et al., 2017; Norval et al., 2020). Our purpose in this paper is to review the published data on the geographic distribution and host range of the snake mite, and to summarise the available information about its clinical significance.

Material and methods

Most of the information presented here was derived from a search of the literature using Web of Science ® and Google Scholar ®, and from the personal libraries of the authors. We collected new information by examining specimens in the Reptile Collection of the Zoological Museum of the Moscow State University (ZMMU) and in the terrarium of the Russian State Agrarian University – Moscow Agricultural Academy. Mites were collected from the body surface of lizards by forceps, transferred to 70% alcohol, and mounted on microscope slides in Faure-Berlese medium. Mites were identified based on keys and descriptions given by Bregetova (1956) and Moraza et al. (2009). Measurements of body parts of imaginal and pre-imaginal stages are presented in Table 1. External measurements were taken in micrometres (μm). Slide-mounted voucher specimens were deposited in the collection of the Parasitological Collection of the Tyumen State Medical University (Tyumen, Russia). The classification of reptiles is based on Uetz et al. (2023).

New material examined

Five females, two protonymphs from Ophisops elegans Ménétries, 1832 from Azerbaijan, Lerik District, Zuvand settlement vicinity 38°47′ N, 48°25′ E, April 2009, leg. A.A. Kidov, identified by M.V. Orlova (Figure 1, Table 2). Additional specimens from Darevskia brauneri (Méhely, 1909) from Russia, Krasnodar region, Ubinskaya settlement, 44°42′N, 38°31′ E, leg. and det. A.A. Kidov; and from Python regius (Shaw, 1802) and Telescopus fallax Fleischmann, 1831, April 2009, from terrarium of Russian State Agrarian University (Moscow), identified by A.A. Kidov. It is most likely that the snakes and lizards acquired O. natricis during their stay in the terrarium in Moscow, so we attributed this record to Russia, not to Azerbaijan (Table 3). Our records of O. natricis for Ophisops elegans, Darevskia brauneri and Telescopus fallax are new host records.

Results

Taxonomic background

The first published reference to the snake mite appears to be that of Metaxa (1823), who observed undentified mites on captive snakes in Italy. Dugès (1834) found it in France and noted its similarity to the bird mite Dermanyssus avium Dugès, 1834. It was first named as Dermanyssus natricis by Gervais (1844), based on specimens collected on captive reptiles in Paris. The same species was also described by several other authors under different names. Camin (1949), Till (1957), Fain (1962) and Micherdzinski (1980) reviewed the earlier literature on the species and listed these junior synonyms, and that information need not be repeated here.

Ophionyssus natricis has been throughly described and illustrated several times, notably by Camin (1953) and Micherdzinski (1980). Moraza et al. (2009) provided detailed information on how O. natricis can be distinguished from other species of Ophionyssus. The morphological recognition of the species is supported by molecular sequence data (Alfonso-Toledo & Paredes-León, 2021). We now provide a list of some of the major taxonomic references to the species, including the important recent works by Moraza et al. (2009), Radovsky (2010) and Beron (2014).

Ophionyssus natricis (Gervais, 1844)

Dermanyssus natricis Gervais, 1844: 223.

Ophionyssus natricis Mégnin, 1884a: 617; 1884b: 110; André, 1937: 62; Vitzthum, 1943; 771; Fonseca, 1948: 313; Camin, 1949: 583; 1953; 3; Zemskaya, 1951: 49; Baker et al., 1956: 33; Bregetova, 1956: 223; Keegan, 1956: 219; Womersley, 1956: 599; Till, 1957: 126; Schweizer, 1961: 155; Fain, 1962: 107; Domrow, 1963: 214; 1974: 17; 1985: 152; 1988: 857; Evans & Till, 1966: 337; Beron, 1966: 52; Costa, 1966: 75; Hallas, 1978: 28; Mehl, 1979: 33; Arutunjan & Ohandjanian, 1983: 312; Zhang & Uchikawa, 1993: 76; Moraza et al., 2009: 65; Radovsky, 2010: 108; Beron, 2014: 130.

At least five other species names are now considered to be junior synonyms of O. natricis (see Beron, 2014 for details).

Misidentifications of Ophionyssus natricis

Some published records of O. natricis actually refer to other species. Biological Services (2015) includes an illustration of a mite on the head of a snake, but the mite appears to be Ophiomegistus Banks, 1914 and not Ophionyssus. The illustration of a snake mite in Maxwell (2022) shows an oribatid mite. Sabu et al. (2002) reported a mite they identified as O. natricis on snakes in India. Their illustration shows an unidentified species of Astigmata, not O. natricis, and the occurrence of O. natricis inside nodules of necrotic tissue as they reported would be extremely unusual.

Geographic distribution

The data listed in Table 2 show that Ophionyssus natricis is near cosmopolitan, and has been found in every continent except Antarctica. Records for Russia are listed in more detail in Table 3. The apparent absence of O. natricis in China is surprising. Su et al. (2010) and Ma & Bai (2012) reported other species of Ophionyssus in China, and O. natricis will almost certainly be found there when further studies are carried out.

Life cycle and behaviour

The life cycle and behaviour of Ophionyssus natricis were examined in detail by Camin (1953). Individuals pass through the egg, larva, protonymph, and deutonymph stages before developing into adult males and females. Protonymphs and adults are hematophagous, feeding on the host and then molting in the environment, but larvae and deutonymphs do not feed. The life cycle can be completed within 7 to 14 days when environmental and host conditions are optimal, in temperatures ranging from 20º C to 30º C and humidity higher than 75%. Oliver (1971) summarised previous results showing that O. natricis reproduces by arrhentokous parthenogenesis, with nine chromosomes in males and 18 in females. Aspects of the biology and behaviour of O. natricis are described in a large number of publications, including the useful summaries by Reinert & Brandstätter (1993), Wozniak & DeNardo (2000), Fitzgerald & Vera (2006) and Schilliger et al. (2013).

Bannert et al. (2000) provided a detailed description of the life cycle of Ophionyssus galloticolus Fain & Bannert, 2000, which appears to be very similar to that of O. natricis.

Occurrence on aquatic hosts

Captive snakes infested by O. natricis seek relief from irritation by immersing themselves in water (for example Page, 1966; Šlapeta et al., 2018), and dead mites can be found floating in water bowls in terraria, suggesting that mites are easily killed by immersion in water. It therefore seems unlikely that reptiles that spend significant time in water would be acceptable hosts for this parasite. We conducted a Web of Science® search for records of O. natricis on marine and aquatic snakes in the following genera: Acrochordus Hornstedt, 1787; Afronatrix Rossman & Eberle, 1977; Agkistrodon Palisot de Beauvois, 1799; Aipysurus Lacépède, 1804; Cerberus Cuvier, 1829; Emydocephalus Krefft, 1869; Enhydris Sonnini & Latreille, 1802; Ephalophis Smith, 1931; Farancia Gray, 1842; Fowlea Theobald, 1868; Grayia Günther, 1858; Helicops Wagler, 1828; Homalopsis Kuhl & Hasselt, 1822; Hydrelaps Boulenger, 1896; Hydrophis Latreille, 1801; Hydrops Wagler, 1830; Laticauda Laurenti, 1768; Leptodeira Fitzinger, 1843; Myron Gray, 1849; Myrrophis Kumar et al., 2012; Natrix Laurenti, 1768; Nerodia Baird & Girard, 1853; Opisthotropis Günther, 1872; Parahydrophis Burger & Natsuno, 1974; Pseudoeryx Fitzinger, 1826; Ptychophis Gomes, 1915; Tretanorhinus Duméril et al., 1854; Trimerodytes Cope, 1895. Positive results were returned for Helicops, Leptideira, and Natrix. Mauri (1967) reported O. natricis on Helicops leopardinus (Schlegel, 1837) and Leptodeira annulata (Linnaeus, 1758) in Argentina. Both these species have been referred to as water snakes, but they are not completely aquatic (Ávila et al., 2006; Thaler et al., 2022). Feider & Solomon (1963), Markov et al. (1964), Bogdanov (1965), and Beron (1966) reported the snake mite on Natrix tessellata Laurenti, 1768 in Uzbekistan, Bulgaria, Romania and Russia (Astrakhan') but did not provide detailed collecting data. Chiodini et al. (1983) and Dik (2012) reported O. natricis on Natrix spp. in captivity but these hosts also cannot be considered as fully aquatic.

Some documents report the occurrence of O. natricis on crocodiles, but these all appear to be the result of mistakes or misunderstandings. Dhooria (2016) reported this host association without any reference to its source. It is also found on many internet sites including Biological Services (2015), Maxwell (2022), and Reptiles Cove (2022), but these records are not supported by evidence.

Wozniak & DeNardo (2000) referred to the host range of O. natricis as follows ''The mite has been shown to thrive on most snakes and some lizards including southern alligator lizards, Elgaira mulicarnata (Wozniak, personal observations), blue-tongue skinks Tiliqua scincoides (Wozniak, personal observations) and side-blotched lizards Uta stansburiana (Goldberg and Bursey, 1991)''. Mendoza-Roldan (2019) misquoted that information when describing O. natricis as ''a cosmopolitan inhabitant of captive snakes, but also infest captive lizards, turtles, crocodiles and other reptiles (Wozniak & DeNardo, 2000)''. This incorrect information was not repeated in Mendoza-Roldan et al. (2020a, 2020b), who reported O. natricis only on snakes and lizards.

Numerous publications refer to ectoparasites of crocodilians, including ticks and leeches, but none of these mention Ophionyssus sp. on this host (e.g. Jacobson, 1984; Magnusson, 1985; Huchzermeyer, 2003; Leslie et al., 2011; Tellez, 2013; Divers & Stahl, 2019; Partyka, 2019; Pereira & Colli, 2023). We have been unable to find any confirmed records of Ophionyssus sp. on any species of crocodilian.

Pest control

Snake mites are highly mobile, and can quickly infest and re-infest terraria or enclosures. They can survive for extended periods without feeding (Wozniak & DeNardo, 2000), so routine environmental hygiene alone is not an effective method of control. Fitzgerald & Vera (2006) reviewed the chemical pesticides used for control of snake mites, as well as a variety of cultural methods for limiting their populations. Commonly used insecticides and acaricides, including those previously reviewed by Camin et al. (1964), are potentially harmful to reptiles, and are not recommended for control of snake mites. Alternative methods of mite control have included the use of sorptive dust, which operates by causing dehydration (Tarshis, 1960). Fitzgerald (2019) listed the currently available methods for controlling snake mites, and the methods and materials that are no longer recommended. A new generation of isoxazoline compounds appears to have considerable potential to provide safe and effective mite control when administered to snakes orally without side effects and without the need to treat the environment of the snake (Fuantos-Gámez et al., 2020; Gobble, 2022; Mendoza-Roldan et al., 2023).

Some authors have proposed the use of predatory mites to control O. natricis on captive reptiles. Rotter (1963) used Cheyletus eruditus Schrank, 1781 (Acari: Cheyletidae) to control the snake mite on captive lizards, and C. eruditus is now available commercially for that purpose (Schilliger et al., 2013; APPI Biological Control, 2021). Stratiolaelaps scimitus (Womersley, 1956) (Acari: Laelapidae) may also have some effect as a predator in terraria, but its value is limited by its low tempertature optimum (Mendyk, 2015). Maslova & Dochevoy (2016) reported a carabid beetle (Carabus granulatus telluris Bates, 1883) moving around on the scales of a pitviper (Gloydius ussuriensis (Emelianov, 1929)) and feeding on ectoparasites, apparently including O. natricis.

Clinical significance

Infestation of reptiles with Ophionyssus natricis can cause dehydration, lethargy, growth impairment (Wozniak & DeNardo, 2000) and in severe infestations, anaemia and dysecdysis (DeNardo & Wozniak, 1997; Mendoza-Roldan et al., 2023). Affected individuals suffer hyperaemic and oedematous skin, and seek relief from the resulting pruritus by soaking in water (Wozniak & DeNardo, 2000). There have been reports of loreal pit inflammations and impactions associated with heavy infestations (Garrett & Harwell, 1991). Histopathologic evaluation of feeding sites shows infiltration with neutrophils, lymphocytes and plasma cells, with multifocal perivascular aggregates of lymphocytes and plasma cells in the adjacent dermis (DeNardo & Wozniak, 1997).

Ophionyssus natricis is a mechanical vector of Aeromonas hydrophila Chester, 1901, the causative agent of hemorrhagic disease in reptiles (Mendoza-Roldan et al., 2021). It can be a vector of blood-borne protozoa and viral pathogens of snakes (Camin, 1948; Chiodini et al, 1983; Schumacher et al, 1994; Mendoza-Roldan et al., 2023). In South America it was identified as a vector of Hepatozoon sp. and Rickettsia sp. (Mendoza-Roldan et al., 2020a, b). It has also been implicated as vector of Arenavirus sp., the etiological agent of the Inclusion Body Disease in boid snakes (Chang & Jacobson, 2010; Divers & Stahl, 2019). Ophionyssus natricis collected from Boa constrictor in Italy also contained Wolbachia sp. (Manoj et al., 2021). Further vector competency studies are needed to characterise the overall role of this parasite in the transmission of other infectious agents, such as filariids.

This parasite can become a pest to humans due to the aggressive feeding of the protonymphs, which swarm and bite humans, causing papular vesiculo-bullous eruptions in the skin (Schultz, 1975; McClain et al., 2009), other bite-associated dermatitis (Beck, 1996; Amanatfard et al., 2014), and the risk of zoonotic transmission of pathogens.

Ecological significance

Schroeder (1934) presented a very clear description of the life cycle of O. natricis, and its importance as a pest of snakes in captivity. He also pointed out the agricultural value of snakes as a natural means of reducing rodent populations. Fonseca (1948), citing Schroeder (1934), speculated that O. natricis was harmful to wild snake populations, and therefore indirectly influenced wild rodent populations. That conclusion is not supported by the observation that O. natricis rarely reaches heavy levels of infestation of snakes in natural habitats.

Discussion

The catalogue of the family Macronyssidae by Beron (2014) provides a good taxonomic background to the systematics of Ophionyssus natricis. The genus Ophionyssus currently includes 14 valid species. Several other species have been placed in Ophionyssus at some time, but have since been transferred to other genera, including Thigmonyssus myrmecophagus (Fonseca, 1954), Trichonyssus ehmanni (Domrow, 1985), T. galeotes (Domrow et al., 1980), and T. scincorum (Domrow et al., 1980).

Most species of Ophionyssus have a limited geographic distribution but O. saurarum (Oudemans, 1901) is widespread in Europe, Asia, and Africa, and O. natricis is cosmopolitan. If we exclude O. natricis, the highest level of species diversity in the genus is found in Africa, with five species in southern Africa and a further three in the Canary Islands. Two species have been described from the Indo-Australian Region, and four occur in Europe and the Western Palaearctic.

Ophionyssus natricis occurs on a very wide range of snake and reptile hosts, while other species of Ophionyssus parasitise lizards in the families Lacertidae, Scincidae, Cordylidae, Agamidae, and Pygopodidae. Reptilian hosts of O. natricis include three families of turtles, 10 families of lizards, and seven families of snakes. The snake family with the largest number of infested species (46) is Colubridae, reflecting the fact that it is the most numerous group of modern snakes, with more than half of all known species. Approximately 20% of the hosts are terrarium species that are not typical of the countries where the infested specimens were found.

The evolutionary history of O. natricis has been obscured by human-assisted dispersal. Till (1957) recorded its presence on both snakes and lizards in wild conditions in South Africa. Field records of O. natricis in other countries are rare, suggesting that the species may have arisen in southern Africa, possibly following a host shift from a lizard to a snake. Nieri-Bastos et al. (2011) and Gomes-Almeida & Pepato (2021) demonstrated the potential value of molecular systematics in Macronyssidae. The required sequence data is not yet available for Ophionyssus, so a molecular analysis of the phylogenetic background of the snake mite is not yet possible.

Ophionyssus natricis has recently been found on wild populations of lizards in Australia (Norval et al., 2020, 2021). Research is needed to determine whether this parasite represents a serious threat to wild populations of reptiles, both in Australia and elsewhere.

Acknowledgments

The study was supported by the Ministry of Science and Higher Education of the Russian Federation (topic no. 122031100282-2). Especial thank to Nikolay V. Anisimov (Tyumen State University, Russia) for his help with pictures. Authors contributed equally to this work.

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