# Review of the distribution and biology of the snake mite
## Ophionyssus natricis (Acari: Macronyssidae)

Maria V Orlova, Bruce Halliday, Will K Reeves, Igor V Doronin, Vladimir A Mishchenko, Ivan V Vyalykh, Artem A Kidov

► To cite this version:

Maria V Orlova, Bruce Halliday, Will K Reeves, Igor V Doronin, Vladimir A Mishchenko, et al. Review of the distribution and biology of the snake mite Ophionyssus natricis (Acari: Macronyssidae). Acarologia, 2024, 64 (2), pp.637-653. 10.24349/gmr0-8m9o. hal-04582892

HAL Id: hal-04582892

https://hal.science/hal-04582892

Submitted on 22 May 2024

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# Review of the distribution and biology of the snake mite
## Ophionyssus natricis (Acari: Macronyssidae)

Maria V Orlova, Bruce Halliday, Will K Reeves, Igor V Doronin, Vladimir A Mishchenko, Ivan V Vyalykh, Artem A Kidov

<figure><img src="image_1.png" /></figure>

► To cite this version:

Maria V Orlova, Bruce Halliday, Will K Reeves, Igor V Doronin, Vladimir A Mishchenko, et al. Review of the distribution and biology of the snake mite Ophionyssus natricis (Acari: Macronyssidae). Acarologia, 2024, 64 (2), pp.637-653. 10.24349/gmr0-8m9o. hal-04582892

HAL Id: hal-04582892

https://hal.science/hal-04582892v1

Submitted on 22 May 2024

HAL is a multi-disciplinary open access
archive for the deposit and dissemination of sci-
entific research documents, whether they are pub-
lished or not. The documents may come from
teaching and research institutions in France or
abroad, or from public or private research centers.

L'archive ouverte pluridisciplinaire HAL, est
destinée au dépôt et à la diffusion de documents
scientifiques de niveau recherche, publiés ou non,
émanant des établissements d'enseignement et de
recherche français ou étrangers, des laboratoires
publics ou privés.

<figure><img src="image_2.png" /></figure>
Acarologia

Open Science
in Acarology

# Review of the distribution and biology of the snake mite *Ophionyssus natricis* (Acari: Macronyssidae)

Maria V. Orlova IDa,b,c, Bruce Halliday Id, Will K. Reeves Id,e, Igor V. Doronin Id,f, Vladimir A. Mishchenko Id,g, Ivan V. Vyalykh Id,g, Artem A. Kidov Id,h

## Original research

### ABSTRACT

The mite *Ophionyssus natricis* (Gervais, 1844) (Macronyssidae) is a widespread ectoparasite of reptiles and a vector of some important veterinary diseases. This paper reviews the geographic distribution and host range of *O. natricis* and the available information about its clinical significance. Some incorrect identifications and host records are also identified. The snake mite has been collected from 144 species of reptiles, mostly snakes, as well as one amphibian and three species of mammals. It has been recorded in 36 countries in Europe, Asia, Africa, North and South America, and Australia. Its geographic range has undoubtedly been extended by human activities. Our records of *O. natricis* on the lizards *Darevskia brauneri* (Méhely, 1909) and *Ophisops elegans* Ménétries 1832, and on the snake *Telescopus fallax* Fleischmann, 1831, are new host records.

**Keywords** reptile; lizard; snake; ectoparasite

### 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.

Received 27 January 2024
Accepted 16 April 2024
Published 06 May 2024

Corresponding author
Maria V. Orlova ID:
masha_orlova@mail.ru

Academic editor
Roy, Lise

https://doi.org/10.24349/gmr0-8m90

ISSN 0044-586X (print)
ISSN 2107-7207 (electronic)

<figure><img src="image_2.png" /></figure>

Orlova M. V. et al.

Licensed under
Creative Commons CC-BY 4.0

<footnote>a Tyumen State Medical University, Tyumen, Russia.</footnote>

<footnote>b National Research Tomsk State University, Tomsk, Russia.</footnote>

<footnote>c Federal Scientific Research Institute of Viral Infections «Virome» of Federal Service for Surveillance on Consumer Rights Protection and Human Wellbeing, Ekaterinburg, Russia.</footnote>

<footnote>d Australian National Insect Collection, CSIRO, Canberra, Australia.</footnote>

<footnote>e C. P. Gillette Museum of Arthropod Diversity, Fort Collins, USA.</footnote>

<footnote>f Zoological Institute of Russian Academy of Science, Saint Petersburg, Russia.</footnote>

<footnote>g Federal Scientific Research Institute of Viral Infections Virome of Federal Service for Surveillance on Consumer Rights Protection and Human Wellbeing, Ekaterinburg, Russia.</footnote>

<footnote>h Russian State Agrarian University - Moscow Agricultural Academy named after. K. A. Timiryazev, Moscow, Russia.</footnote>
Acarologia

Open Science
in Acarology

# Review of the distribution and biology of the snake mite *Ophionyssus natricis* (Acari: Macronyssidae)

Maria V. Orlova IDa,b,c, Bruce Halliday Id, Will K. Reeves Id,e, Igor V. Doronin Id,f, Vladimir A. Mishchenko Id,g, Ivan V. Vyalykh Id,g, Artem A. Kidov Id,h

## Original research

### ABSTRACT

The mite *Ophionyssus natricis* (Gervais, 1844) (Macronyssidae) is a widespread ectoparasite of reptiles and a vector of some important veterinary diseases. This paper reviews the geographic distribution and host range of *O. natricis* and the available information about its clinical significance. Some incorrect identifications and host records are also identified. The snake mite has been collected from 144 species of reptiles, mostly snakes, as well as one amphibian and three species of mammals. It has been recorded in 36 countries in Europe, Asia, Africa, North and South America, and Australia. Its geographic range has undoubtedly been extended by human activities. Our records of *O. natricis* on the lizards *Darevskia brauneri* (Méhely, 1909) and *Ophisops elegans* Ménétries 1832, and on the snake *Telescopus fallax* Fleischmann, 1831, are new host records.

**Keywords** reptile; lizard; snake; ectoparasite

### 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.

Received 27 January 2024
Accepted 16 April 2024
Published 06 May 2024

Corresponding author
Maria V. Orlova ID:
masha_orlova@mail.ru

Academic editor
Roy, Lise

https://doi.org/10.24349/gmr0-8m90

ISSN 0044-586X (print)
ISSN 2107-7207 (electronic)

<figure><img src="image_3.png" /></figure>

Orlova M. V. et al.

Licensed under
Creative Commons CC-BY 4.0

<footnote>ªTyumen State Medical University, Tyumen, Russia.</footnote>

<footnote>ᵇNational Research Tomsk State University, Tomsk, Russia.</footnote>

<footnote>ᶜFederal Scientific Research Institute of Viral Infections «Virome» of Federal Service for Surveillance on Consumer Rights Protection and Human Wellbeing, Ekaterinburg, Russia.</footnote>

<footnote>ᵈAustralian National Insect Collection, CSIRO, Canberra, Australia.</footnote>

<footnote>ᵉC. P. Gillette Museum of Arthropod Diversity, Fort Collins, USA.</footnote>

<footnote>fZoological Institute of Russian Academy of Science, Saint Petersburg, Russia.</footnote>

<footnote>gFederal Scientific Research Institute of Viral Infections Virome of Federal Service for Surveillance on Consumer Rights Protection and Human Wellbeing, Ekaterinburg, Russia.</footnote>

<footnote>hRussian State Agrarian University - Moscow Agricultural Academy named after. K. A. Timiryazev, Moscow, Russia.</footnote>
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).

<table><caption>**Table 1** Measurements of *Ophionyssus natricis* (Gervais, 1844) from different parts of the distribution area.</caption><thead><tr><th></th><th>Brazil<br>(Micherdzinski, 1980)</th><th>Japan<br>(Zhang & Uchikawa, 1993)</th><th>Romania<br>(Miron & Ivan, 2003)</th><th>Azerbaijan<br>(this study, n=5)</th></tr></thead><tbody><tr><td colspan="5" style="text-align:center;">Female</td></tr><tr><td colspan="5" style="text-align:center;">Idiosoma</td></tr><tr><td>Length</td><td>990</td><td>—</td><td>985 (944–1056)</td><td>654 (622–689)</td></tr><tr><td>Width</td><td>667</td><td>—</td><td>—</td><td>314 (300–322)</td></tr><tr><td colspan="5" style="text-align:center;">Propodosomal shield</td></tr><tr><td>Length</td><td>310</td><td>270–310</td><td>274 (252–289)</td><td>289 (271–294)</td></tr><tr><td>Width</td><td>263</td><td>265–325</td><td>—</td><td>261 (255–267)</td></tr><tr><td colspan="5" style="text-align:center;">Pygidial shield</td></tr><tr><td>Length</td><td>—</td><td>50–60</td><td>70 (60–78)</td><td>55–61</td></tr><tr><td>Width</td><td>—</td><td>50–60</td><td>—</td><td>52–59</td></tr><tr><td colspan="5" style="text-align:center;">Sternal shield</td></tr><tr><td>Length</td><td>45</td><td>40–45</td><td>46 (36–54)</td><td>49 (43–53)</td></tr><tr><td>Width</td><td>112</td><td>103–110</td><td>—</td><td>91 (86–94)</td></tr><tr><td colspan="5" style="text-align:center;">Genital shield</td></tr><tr><td>Length</td><td>275</td><td>290–320</td><td>—</td><td>273 (267–279)</td></tr><tr><td>Width</td><td>288</td><td>—</td><td>—</td><td>—</td></tr><tr><td colspan="5" style="text-align:center;">Stigmata and peritreme</td></tr><tr><td>Length</td><td>144</td><td>128–145</td><td>—</td><td>—</td></tr><tr><td colspan="5" style="text-align:center;">Anal shield</td></tr><tr><td>Length</td><td>—</td><td>110–145</td><td>125 (114–144)</td><td>121 (120–123)</td></tr><tr><td>Width</td><td>—</td><td>68–85</td><td>—</td><td>67 (65–68)</td></tr><tr><td colspan="5" style="text-align:center;">Gnathosoma</td></tr><tr><td>Length</td><td>180–183</td><td>—</td><td>—</td><td>176 (171–179)</td></tr><tr><td colspan="5" style="text-align:center;">Chelicera, fixed digit</td></tr><tr><td>Length</td><td>—</td><td>45–50</td><td>—</td><td>44–50</td></tr><tr><td colspan="5" style="text-align:center;">Protonymph (n=2)</td></tr><tr><td colspan="5" style="text-align:center;">Idiosoma</td></tr><tr><td>Length</td><td>366–381</td><td>—</td><td>—</td><td>361, 369</td></tr><tr><td>Width</td><td>240–251</td><td>—</td><td>—</td><td>221, 227</td></tr><tr><td colspan="5" style="text-align:center;">Propodosomal shield</td></tr><tr><td>Length</td><td>—</td><td>180–235</td><td>—</td><td>177, 181</td></tr><tr><td>Width</td><td>—</td><td>190–240</td><td>—</td><td>182, 189</td></tr><tr><td colspan="5" style="text-align:center;">Pygidial shield</td></tr><tr><td>Length</td><td>—</td><td>50–75</td><td>—</td><td>48, 52</td></tr><tr><td>Width</td><td>—</td><td>75–90</td><td>—</td><td>71, 79</td></tr><tr><td colspan="5" style="text-align:center;">Sternal shield</td></tr><tr><td>Length</td><td>—</td><td>110</td><td>—</td><td>101</td></tr><tr><td>Width</td><td>—</td><td>75–80</td><td>—</td><td>72</td></tr><tr><td colspan="5" style="text-align:center;">Anal shield</td></tr><tr><td>Length</td><td>—</td><td>55–65</td><td>—</td><td>41, 48</td></tr><tr><td>Width</td><td>—</td><td>42–55</td><td>—</td><td>36, 38</td></tr></tbody></table>
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).

<table><caption>**Table 1** Measurements of *Ophionyssus natricis* (Gervais, 1844) from different parts of the distribution area.</caption><thead><tr><th></th><th>Brazil<br>(Micherdzinski, 1980)</th><th>Japan<br>(Zhang & Uchikawa, 1993)</th><th>Romania<br>(Miron & Ivan, 2003)</th><th>Azerbaijan<br>(this study, n=5)</th></tr></thead><tbody><tr><td colspan="5" style="text-align: center;">Female</td></tr><tr><td colspan="5" style="text-align: center;">Idiosoma</td></tr><tr><td>Length</td><td>990</td><td>—</td><td>985 (944–1056)</td><td>654 (622–689)</td></tr><tr><td>Width</td><td>667</td><td>—</td><td>—</td><td>314 (300–322)</td></tr><tr><td colspan="5" style="text-align: center;">Propodosomal shield</td></tr><tr><td>Length</td><td>310</td><td>270–310</td><td>274 (252–289)</td><td>289 (271–294)</td></tr><tr><td>Width</td><td>263</td><td>265–325</td><td>—</td><td>261 (255–267)</td></tr><tr><td colspan="5" style="text-align: center;">Pygidial shield</td></tr><tr><td>Length</td><td>—</td><td>50–60</td><td>70 (60–78)</td><td>55–61</td></tr><tr><td>Width</td><td>—</td><td>50–60</td><td>—</td><td>52–59</td></tr><tr><td colspan="5" style="text-align: center;">Sternal shield</td></tr><tr><td>Length</td><td>45</td><td>40–45</td><td>46 (36–54)</td><td>49 (43–53)</td></tr><tr><td>Width</td><td>112</td><td>103–110</td><td>—</td><td>91 (86–94)</td></tr><tr><td colspan="5" style="text-align: center;">Genital shield</td></tr><tr><td>Length</td><td>275</td><td>290–320</td><td>—</td><td>273 (267–279)</td></tr><tr><td>Width</td><td>288</td><td>—</td><td>—</td><td>—</td></tr><tr><td colspan="5" style="text-align: center;">Stigmata and peritreme</td></tr><tr><td>Length</td><td>144</td><td>128–145</td><td>—</td><td>—</td></tr><tr><td colspan="5" style="text-align: center;">Anal shield</td></tr><tr><td>Length</td><td>—</td><td>110–145</td><td>125 (114–144)</td><td>121 (120–123)</td></tr><tr><td>Width</td><td>—</td><td>68–85</td><td>—</td><td>67 (65–68)</td></tr><tr><td colspan="5" style="text-align: center;">Gnathosoma</td></tr><tr><td>Length</td><td>180–183</td><td>—</td><td>—</td><td>176 (171–179)</td></tr><tr><td colspan="5" style="text-align: center;">Chelicera, fixed digit</td></tr><tr><td>Length</td><td>—</td><td>45–50</td><td>—</td><td>44–50</td></tr><tr><td colspan="5" style="text-align: center;">Protonymph (n=2)</td></tr><tr><td colspan="5" style="text-align: center;">Idiosoma</td></tr><tr><td>Length</td><td>366–381</td><td>—</td><td>—</td><td>361, 369</td></tr><tr><td>Width</td><td>240–251</td><td>—</td><td>—</td><td>221, 227</td></tr><tr><td colspan="5" style="text-align: center;">Propodosomal shield</td></tr><tr><td>Length</td><td>—</td><td>180–235</td><td>—</td><td>177, 181</td></tr><tr><td>Width</td><td>—</td><td>190–240</td><td>—</td><td>182, 189</td></tr><tr><td colspan="5" style="text-align: center;">Pygidial shield</td></tr><tr><td>Length</td><td>—</td><td>50–75</td><td>—</td><td>48, 52</td></tr><tr><td>Width</td><td>—</td><td>75–90</td><td>—</td><td>71, 79</td></tr><tr><td colspan="5" style="text-align: center;">Sternal shield</td></tr><tr><td>Length</td><td>—</td><td>110</td><td>—</td><td>101</td></tr><tr><td>Width</td><td>—</td><td>75–80</td><td>—</td><td>72</td></tr><tr><td colspan="5" style="text-align: center;">Anal shield</td></tr><tr><td>Length</td><td>—</td><td>55–65</td><td>—</td><td>41, 48</td></tr><tr><td>Width</td><td>—</td><td>42–55</td><td>—</td><td>36, 38</td></tr></tbody></table>
<figure><img src="image_3.png" /><figcaption>**Figure 1** *Ophionyssus natricis* (ex *Ophisops elegans* from Azerbaijan): A – Female, propodosomal shield, scale bar = 50 µm; B – Female, pygidial shield, scale bar = 100 µm; C – Female, sternal and genital shields, scale bar = 50 µm; D – Protonymph, ventral idiosoma, scale bar = 100 µm.</figcaption></figure>

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.
<figure><img src="image_4.png" /><figcaption>**Figure 1** *Ophionyssus natricis* (ex *Ophisops elegans* from Azerbaijan): A – Female, propodosomal shield, scale bar = 50 µm; B – Female, pygidial shield, scale bar = 100 µm; C – Female, sternal and genital shields, scale bar = 50 µm; D – Protonymph, ventral idiosoma, scale bar = 100 µm.</figcaption></figure>

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 unidentified 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.

<table><caption>**Table 2** Published records of *Ophionyssus natricis*. Non-reptile hosts are shown in square brackets.</caption><thead><tr><th>Country</th><th>Host</th><th>References</th><th>Notes</th><th>Species range</th></tr></thead><tbody><tr><td colspan="5" style="text-align:center; font-weight:bold;">Europe</td></tr><tr><td>Norway</td><td>Not stated</td><td>Mehl, 1979</td><td></td><td></td></tr><tr><td>Sweden</td><td>Not stated</td><td>Ahl, 1994</td><td></td><td></td></tr><tr><td>Denmark</td><td>[Homo sapiens]</td><td>Schultz, 1975</td><td></td><td></td></tr><tr><td>Bulgaria</td><td>Natrix tessellata</td><td>Beron, 1966</td><td></td><td>Native species</td></tr><tr><td rowspan="5">Romania</td><td><strong>Snakes</strong></td><td></td><td></td><td></td></tr><tr><td>Natrix natrix</td><td>Feider & Solomon, 1963</td><td></td><td>Native species</td></tr><tr><td>Natrix tessellata</td><td>Miron & Ivan, 2003</td><td>Mite as Ophionyssus viperae</td><td>Native species</td></tr><tr><td>Zamenis longissimus</td><td>Miron & Ivan, 2003</td><td>As Elaphe longissima</td><td>Native species</td></tr><tr><td>Dolichophis caspius</td><td>Miron & Ivan, 2003</td><td>As Coluber jugularis caspius</td><td>Native species</td></tr><tr><td rowspan="5">Great Britain</td><td>Vipera ursinii</td><td>Miron & Ivan, 2003</td><td></td><td>Native species</td></tr><tr><td><strong>Lizards</strong></td><td></td><td></td><td></td></tr><tr><td>Uta stansburiana hesperis</td><td>Burrage, 1966</td><td></td><td>Non-native species</td></tr><tr><td>Sceloporus occidentalis occidentalis</td><td>Burrage, 1966</td><td></td><td>Non-native species</td></tr><tr><td>Pogona vitticeps</td><td>Castro, 2019</td><td></td><td>Non-native species</td></tr><tr><td rowspan="5">Poland</td><td>Not identified</td><td>Evans & Till, 1966</td><td></td><td></td></tr><tr><td><strong>Snakes</strong></td><td></td><td></td><td></td></tr><tr><td>Thamnophis ordinoides</td><td>André, 1937</td><td>As Uta stansburiana elegans</td><td>Non-native species</td></tr><tr><td>Cerastes vipera</td><td>André, 1937</td><td>As Cerastes aegyptiacus</td><td>Non-native species</td></tr><tr><td>Malayopython reticulatus</td><td>André, 1937</td><td>As Python reticulatus</td><td>Non-native species</td></tr><tr><td rowspan="2">Germany</td><td>Epicrates cenchria</td><td>Mendoza-Roldan et al., 2019</td><td></td><td>Non-native species</td></tr><tr><td><strong>Turtles</strong></td><td></td><td></td><td></td></tr><tr><td></td><td>Testudo hermanni boettgeri</td><td>Castro, 2019</td><td></td><td>Non-native species</td></tr><tr><td rowspan="2">Poland</td><td><strong>Snakes</strong></td><td></td><td></td><td></td></tr><tr><td>Boa constrictor</td><td>Pawelczyk et al., 2016</td><td></td><td>Non-native species</td></tr><tr><td rowspan="3">Germany</td><td><strong>Lizards</strong></td><td></td><td></td><td></td></tr><tr><td>Iguana iguana</td><td>Beck & Pantchev, 2006; Beck & Pfister, 2006</td><td></td><td>Non-native species</td></tr><tr><td><strong>Turtles</strong></td><td></td><td></td><td></td></tr><tr><td></td><td>Testudo hermanni boettgeri</td><td>Wiechert, 2007</td><td></td><td>Non-native species</td></tr><tr><td rowspan="3">France</td><td><strong>Snakes</strong></td><td></td><td></td><td></td></tr><tr><td>Natrix natrix</td><td>André, 1937</td><td>As Tropidonotus natrix</td><td>Native species</td></tr><tr><td>Hemorrhös hippocrepis</td><td>André, 1937</td><td>As Periops hippocrepis</td><td>Non-native species</td></tr><tr><td>Netherland</td><td>Snakes (not identified)</td><td>Buitendijk, 1945</td><td></td><td></td></tr><tr><td>Austria</td><td>Captive snakes, occasionally captive</td><td>Hassl, 2016</td><td></td><td></td></tr><tr><td rowspan="2">Switzerland</td><td>Agamidae (not identified)</td><td></td><td></td><td></td></tr><tr><td><strong>Snakes</strong></td><td></td><td></td><td></td></tr><tr><td></td><td>Vipera aspis</td><td>Schweizer, 1961</td><td></td><td>Native species</td></tr><tr><td rowspan="10">Italy</td><td><strong>Snakes</strong></td><td></td><td></td><td></td></tr><tr><td>Pantherophis guttatus</td><td>Cervone et al., 2016</td><td></td><td>Non-native species</td></tr><tr><td>Python regius</td><td>Cervone et al., 2016</td><td></td><td>Non-native species</td></tr><tr><td>P. bivittatus</td><td>Cervone et al., 2016</td><td></td><td>Non-native species</td></tr><tr><td>Epicrates cenchria</td><td>Cervone et al., 2016</td><td></td><td>Non-native species</td></tr><tr><td>Morelia spilota</td><td>Mendoza-Roldán et al., 2019</td><td></td><td>Non-native species</td></tr><tr><td>Eunectes murinus</td><td>Mendoza-Roldán et al., 2019</td><td></td><td>Non-native species</td></tr><tr><td>Boa constrictor</td><td>Mendoza-Roldán et al., 2019</td><td></td><td>Non-native species</td></tr><tr><td>Podarcis siculus</td><td>Mendoza-Roldán et al., 2019</td><td></td><td>Native species</td></tr><tr><td>Podarcis siculus klemmeri</td><td>Mendoza-Roldán et al., 2019</td><td></td><td>Native species</td></tr><tr><td rowspan="2">Spain (Canary islands)</td><td><strong>Snakes</strong></td><td></td><td></td><td></td></tr><tr><td>Lampropeltis californiae</td><td>Santana-Hernández et al., 2021</td><td></td><td>Non-native species</td></tr><tr><td rowspan="2">Slovenia</td><td><strong>Snakes</strong></td><td></td><td></td><td></td></tr><tr><td>Boa constrictor</td><td>Rataj et al., 2011</td><td></td><td>Non-native species</td></tr></tbody></table>
# Results

## Taxonomic background

The first published reference to the snake mite appears to be that of Metaxa (1823), who observed unidentified 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.

<table><caption>**Table 2** Published records of *Ophionyssus natricis*. Non-reptile hosts are shown in square brackets.</caption><thead><tr><th>Country</th><th>Host</th><th>References</th><th>Notes</th><th>Species range</th></tr></thead><tbody><tr><td colspan="5" style="text-align:center; font-weight:bold;">Europe</td></tr><tr><td>Norway</td><td>Not stated</td><td>Mehl, 1979</td><td></td><td></td></tr><tr><td>Sweden</td><td>Not stated</td><td>Ahl, 1994</td><td></td><td></td></tr><tr><td>Denmark</td><td>[Homo sapiens]</td><td>Schultz, 1975</td><td></td><td></td></tr><tr><td>Bulgaria</td><td>Natrix tessellata</td><td>Beron, 1966</td><td></td><td>Native species</td></tr><tr><td rowspan="5">Romania</td><td><strong>Snakes</strong></td><td></td><td></td><td></td></tr><tr><td>Natrix natrix</td><td>Feider & Solomon, 1963</td><td></td><td>Native species</td></tr><tr><td>Natrix tessellata</td><td>Miron & Ivan, 2003</td><td>Mite as Ophionyssus viperae</td><td>Native species</td></tr><tr><td>Zamenis longissimus</td><td>Miron & Ivan, 2003</td><td>As Elaphe longissima</td><td>Native species</td></tr><tr><td>Dolichophis caspius</td><td>Miron & Ivan, 2003</td><td>As Coluber jugularis caspius</td><td>Native species</td></tr><tr><td rowspan="5">Great Britain</td><td>Vipera ursinii</td><td>Miron & Ivan, 2003</td><td></td><td>Native species</td></tr><tr><td><strong>Lizards</strong></td><td></td><td></td><td></td></tr><tr><td>Uta stansburiana hesperis</td><td>Burrage, 1966</td><td></td><td>Non-native species</td></tr><tr><td>Sceloporus occidentalis occidentalis</td><td>Burrage, 1966</td><td></td><td>Non-native species</td></tr><tr><td>Pogona vitticeps</td><td>Castro, 2019</td><td></td><td>Non-native species</td></tr><tr><td></td><td>Not identified</td><td>Evans & Till, 1966</td><td></td><td></td></tr><tr><td rowspan="5">Poland</td><td><strong>Snakes</strong></td><td></td><td></td><td></td></tr><tr><td>Thamnophis ordinoides</td><td>André, 1937</td><td>As Uta stansburiana elegans</td><td>Non-native species</td></tr><tr><td>Cerastes vipera</td><td>André, 1937</td><td>As Cerastes aegyptiacus</td><td>Non-native species</td></tr><tr><td>Malayopython reticulatus</td><td>André, 1937</td><td>As Python reticulatus</td><td>Non-native species</td></tr><tr><td>Episcrates cenchria</td><td>Mendoza-Roldan et al., 2019</td><td></td><td>Non-native species</td></tr><tr><td rowspan="2">Germany</td><td><strong>Turtles</strong></td><td></td><td></td><td></td></tr><tr><td>Testudo hermanni boettgeri</td><td>Castro, 2019</td><td></td><td>Non-native species</td></tr><tr><td rowspan="2">Poland</td><td><strong>Snakes</strong></td><td></td><td></td><td></td></tr><tr><td>Boa constrictor</td><td>Pawelczyk et al., 2016</td><td></td><td>Non-native species</td></tr><tr><td rowspan="3">Germany</td><td><strong>Lizards</strong></td><td></td><td></td><td></td></tr><tr><td>Iguana iguana</td><td>Beck & Pantchev, 2006; Beck & Pfister, 2006</td><td></td><td>Non-native species</td></tr><tr><td><strong>Turtles</strong></td><td></td><td></td><td></td></tr><tr><td rowspan="2">France</td><td>Testudo hermanni boettgeri</td><td>Wiechert, 2007</td><td></td><td>Non-native species</td></tr><tr><td><strong>Snakes</strong></td><td></td><td></td><td></td></tr><tr><td rowspan="2">Netherland</td><td>Natrix natrix</td><td>André, 1937</td><td>As Tropidonotus natrix</td><td>Native species</td></tr><tr><td>Hemorrhös hippocrepis</td><td>André, 1937</td><td>As Periops hippocrepis</td><td>Non-native species</td></tr><tr><td>Austria</td><td>Snakes (not identified)</td><td>Buitendijk, 1945</td><td></td><td></td></tr><tr><td></td><td>Captive snakes, occasionally captive</td><td>Hassl, 2016</td><td></td><td></td></tr><tr><td>Switzerland</td><td>Agamidae (not identified)</td><td></td><td></td><td></td></tr><tr><td rowspan="2">Italy</td><td><strong>Snakes</strong></td><td></td><td></td><td></td></tr><tr><td>Vipera aspis</td><td>Schweizer, 1961</td><td></td><td>Native species</td></tr><tr><td rowspan="10">Italy</td><td><strong>Snakes</strong></td><td></td><td></td><td></td></tr><tr><td>Pantherophis guttatus</td><td>Cervone et al., 2016</td><td></td><td>Non-native species</td></tr><tr><td>Python regius</td><td>Cervone et al., 2016</td><td></td><td>Non-native species</td></tr><tr><td>P. bivittatus</td><td>Cervone et al., 2016</td><td></td><td>Non-native species</td></tr><tr><td>Episcrates cenchria</td><td>Cervone et al., 2016</td><td></td><td>Non-native species</td></tr><tr><td>Morelia spilota</td><td>Mendoza-Roldán et al., 2019</td><td></td><td>Non-native species</td></tr><tr><td>Eunectes murinus</td><td>Mendoza-Roldán et al., 2019</td><td></td><td>Non-native species</td></tr><tr><td>Boa constrictor</td><td>Mendoza-Roldán et al., 2019</td><td></td><td>Non-native species</td></tr><tr><td>Podarcis siculus</td><td>Mendoza-Roldán et al., 2019</td><td></td><td>Native species</td></tr><tr><td>Podarcis siculus klemmeri</td><td>Mendoza-Roldán et al., 2019</td><td></td><td>Native species</td></tr><tr><td>Spain (Canary islands)</td><td><strong>Snakes</strong></td><td></td><td></td><td></td></tr><tr><td rowspan="2">Slovenia</td><td>Lampropeltis californiae</td><td>Santana-Hernández et al., 2021</td><td></td><td>Non-native species</td></tr><tr><td><strong>Snakes</strong></td><td></td><td></td><td></td></tr><tr><td></td><td>Boa constrictor</td><td>Rataj et al., 2011</td><td></td><td>Non-native species</td></tr></tbody></table>
*Ophionyssus natricis* has been thoroughly 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-

<table><caption>**Table 2** Continued.</caption><thead><tr><th>Country</th><th>Host</th><th>References</th><th>Notes</th><th>Species range</th></tr></thead><tbody><tr><td colspan="5" style="text-align: center; font-weight: bold;">Asia</td></tr><tr><td>Armenia</td><td><strong>Lizards</strong><br>Darevskia sp.</td><td>Arutunian & Ohandjanian, 1983</td><td>As Lacerta saxicola</td><td></td></tr><tr><td></td><td><strong>Snakes</strong><br>Natrix natrix</td><td>Arutunian & Ohandjanian, 1983</td><td></td><td>Native species</td></tr><tr><td>Uzbekistan</td><td><strong>Snakes</strong><br>Natrix tessellata</td><td>Abdushukurova et al., 1966</td><td></td><td>Native species</td></tr><tr><td>Turkmenistan</td><td><strong>Snakes</strong><br>Psammophis lineolatus</td><td>Markov & Bogdanov, 1960</td><td></td><td>Native species</td></tr><tr><td>(Soviet Central Asia)</td><td><strong>Snakes</strong><br>Echis carinatus<br>Spalerosophis diadema</td><td>Bogdanov, 1965<br>Bogdanov, 1965</td><td></td><td>Native species<br>Native species</td></tr><tr><td>Turkey</td><td><strong>Snakes</strong><br>Natrix tessellata<br>Pantherophis guttatus</td><td>Dik, 2012<br>Keskin, 2021</td><td></td><td>Native species<br>Non-native species</td></tr><tr><td>Iran</td><td>[Homo sapiens]<br><strong>Snakes</strong><br>Python sp.</td><td>Amanatfard et al., 2014<br>Amanatfard et al., 2014</td><td></td><td></td></tr><tr><td>Israel</td><td><strong>Snakes</strong><br>Hemorrhois nummifer<br>Snakes (not identified)</td><td>Costa, 1966<br>Shulov, 1957</td><td>As Coluber ravergieri</td><td>Native species</td></tr><tr><td>Korea</td><td><strong>Snakes</strong><br>Snakes (not identified)</td><td>Whan, 1966</td><td></td><td></td></tr><tr><td>Japan</td><td><strong>Snakes</strong><br>Chondropython viridis</td><td>Zhang & Uchikawa, 1993</td><td></td><td>Non-native species</td></tr><tr><td>Malaysia</td><td><strong>Snakes</strong><br>Python bivittatus</td><td>Mariana et al., 2011</td><td>As Python molurus bivittatus</td><td>Native species</td></tr><tr><td>India</td><td><strong>Snakes</strong><br>Naja naja naja<br>Daboa russelii<br>Chrysopelea ornata</td><td>Khaire et al., 1987<br>Khaire et al., 1987<br>Khaire et al., 1987</td><td>As Vipera russelii</td><td>Native species<br>Native species<br>Native species</td></tr><tr><td>Pakistan</td><td><strong>Snakes</strong><br>Python molurus</td><td>Sumaira et al., 2020</td><td></td><td>Native species</td></tr><tr><td colspan="5" style="text-align: center; font-weight: bold;">Africa</td></tr><tr><td>Egypt</td><td><strong>Snakes</strong><br>Platyceps florulentus<br>Psammophis leightoni<br>Psammophis schokari<br>Spalerosophis diadema cliffordii<br>Naja haje<br>Telescopus obtusus<br>[Sclerophrys mauritanica]</td><td>Keegan, 1956; Yunker, 1956<br>Keegan, 1956; Yunker, 1956<br>Keegan, 1956; Yunker, 1956<br>Keegan, 1956; Yunker, 1956<br>Keegan, 1956; Yunker, 1956<br>Keegan, 1956; Yunker, 1956<br>Blanc & Ascione, 1959</td><td>As Coluber florulentus<br>As Psammophis sibilans<br>As Spalerosophis cliffordii<br>As Telescopus dhara obtusus<br>As [Bufo mauritanicus]</td><td>Native species<br>Native species<br>Native species<br>Native species<br>Native species<br>Native species<br>Native species</td></tr><tr><td>Morocco</td><td><strong>Lizards</strong><br>Mesalina simoni<br><strong>Snakes</strong><br>Malpolon monspessulanus<br>Natrix maura<br>Coluber hippocrepis</td><td>Er-Rguibi et al., 2023<br>Blanc & Ascione, 1959<br>Blanc & Ascione, 1959<br>Blanc & Ascione, 1959</td><td></td><td>Native species</td></tr><tr><td>Congo</td><td><strong>Snakes</strong><br>Bothrophthalmus lineatus</td><td>Fain, 1962</td><td></td><td>Native species</td></tr><tr><td>Republic of South Africa</td><td><strong>Snakes</strong><br>Bitis arietans<br>Boaedon lineatus<br>Causus rhombeatus<br>Dispholidus typus<br>Naja nivea<br>Psammophis leightoni<br>Hemachatus haemachates</td><td>Till, 1957<br>Till, 1957<br>Till, 1957<br>Till, 1957<br>Till, 1957<br>Till, 1957<br>Till, 1957</td><td>As Psammophis sibilans trinasalis</td><td>Native species<br>Native species<br>Native species<br>Native species<br>Native species<br>Native species<br>Native species</td></tr><tr><td>West Africa (country not stated)</td><td><strong>Snakes</strong><br>Boodon fuliginosus</td><td>Phisalix, 1914</td><td></td><td>Native species</td></tr></tbody></table>
*Ophionyssus natricis* has been thoroughly 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-

<table><caption>**Table 2** Continued.</caption><thead><tr><th>Country</th><th>Host</th><th>References</th><th>Notes</th><th>Species range</th></tr></thead><tbody><tr><td colspan="5" style="text-align:center; font-weight:bold;">Asia</td></tr><tr><td>Armenia</td><td><strong>Lizards</strong><br>Darevskia sp.</td><td>Arutunian & Ohandjanian, 1983</td><td>As Lacerta saxicola</td><td></td></tr><tr><td></td><td><strong>Snakes</strong><br>Natrix natrix</td><td>Arutunian & Ohandjanian, 1983</td><td></td><td>Native species</td></tr><tr><td>Uzbekistan</td><td><strong>Snakes</strong><br>Natrix tessellata</td><td>Abdushukurova et al., 1966</td><td></td><td>Native species</td></tr><tr><td>Turkmenistan</td><td><strong>Snakes</strong><br>Psammophis lineolatus</td><td>Markov & Bogdanov, 1960</td><td></td><td>Native species</td></tr><tr><td>(Soviet Central Asia)</td><td><strong>Snakes</strong><br>Echis carinatus<br>Spalerosophis diadema</td><td>Bogdanov, 1965<br>Bogdanov, 1965</td><td></td><td>Native species<br>Native species</td></tr><tr><td>Turkey</td><td><strong>Snakes</strong><br>Natrix tessellata<br>Pantherophis guttatus</td><td>Dik, 2012<br>Keskin, 2021</td><td></td><td>Native species<br>Non-native species</td></tr><tr><td>Iran</td><td>[Homo sapiens]<br><strong>Snakes</strong><br>Python sp.</td><td>Amanatfard et al., 2014<br>Amanatfard et al., 2014</td><td></td><td></td></tr><tr><td>Israel</td><td><strong>Snakes</strong><br>Hemorrhois nummifer<br>Snakes (not identified)</td><td>Costa, 1966<br>Shulov, 1957</td><td>As Coluber ravergieri</td><td>Native species</td></tr><tr><td>Korea</td><td><strong>Snakes</strong><br>Snakes (not identified)</td><td>Whan, 1966</td><td></td><td></td></tr><tr><td>Japan</td><td><strong>Snakes</strong><br>Chondropython viridis</td><td>Zhang & Uchikawa, 1993</td><td></td><td>Non-native species</td></tr><tr><td>Malaysia</td><td><strong>Snakes</strong><br>Python bivittatus</td><td>Mariana et al., 2011</td><td>As Python molurus bivittatus</td><td>Native species</td></tr><tr><td>India</td><td><strong>Snakes</strong><br>Naja naja naja<br>Daboia russelii<br>Chrysopelea ornata</td><td>Khaire et al., 1987<br>Khaire et al., 1987<br>Khaire et al., 1987</td><td>As Vipera russelii</td><td>Native species<br>Native species<br>Native species</td></tr><tr><td>Pakistan</td><td><strong>Snakes</strong><br>Python molurus</td><td>Sumaira et al., 2020</td><td></td><td>Native species</td></tr><tr><td colspan="5" style="text-align:center; font-weight:bold;">Africa</td></tr><tr><td>Egypt</td><td><strong>Snakes</strong><br>Platyceps florulentus<br>Psammophis leightoni<br>Psammophis schokari<br>Spalerosophis diadema cliffordii<br>Naja haje<br>Telescopus obtusus<br>[Sclerophrys mauritanica]</td><td>Keegan, 1956; Yunker, 1956<br>Keegan, 1956; Yunker, 1956<br>Keegan, 1956; Yunker, 1956<br>Keegan, 1956; Yunker, 1956<br>Keegan, 1956; Yunker, 1956<br>Keegan, 1956; Yunker, 1956<br>Blanc & Ascione, 1959</td><td>As Coluber florulentus<br>As Psammophis sibilans<br>As Spalerosophis cliffordii<br>As Telescopus dhara obtusus<br>As [Bufo mauritanicus]</td><td>Native species<br>Native species<br>Native species<br>Native species<br>Native species<br>Native species<br>Native species</td></tr><tr><td>Morocco</td><td><strong>Lizards</strong><br>Mesalina simoni<br><strong>Snakes</strong><br>Malpolon monspessulanus<br>Natrix maura<br>Coluber hippocrepis</td><td>Er-Rguibi et al., 2023<br>Blanc & Ascione, 1959<br>Blanc & Ascione, 1959<br>Blanc & Ascione, 1959</td><td></td><td>Native species<br>Native species<br>Native species<br>Native species</td></tr><tr><td>Congo</td><td><strong>Snakes</strong><br>Bothrophthalmus lineatus</td><td>Fain, 1962</td><td></td><td>Native species</td></tr><tr><td>Republic of South Africa</td><td><strong>Snakes</strong><br>Bitis arietans<br>Boaedon lineatus<br>Causus rhombeatus<br>Dispholidus typus<br>Naja nivea<br>Psammophis leightoni<br>Hemachatus haemachates</td><td>Till, 1957<br>Till, 1957<br>Till, 1957<br>Till, 1957<br>Till, 1957<br>Till, 1957<br>Till, 1957</td><td>As Psammophis sibilans trinasalis</td><td>Native species<br>Native species<br>Native species<br>Native species<br>Native species<br>Native species<br>Native species</td></tr><tr><td>West Africa (country not stated)</td><td><strong>Snakes</strong><br>Boodon fuliginosus</td><td>Phisalix, 1914</td><td></td><td>Native species</td></tr></tbody></table>
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).

<table><caption>**Table 2** Continued.</caption><thead><tr><th>Country</th><th>Host</th><th>References</th><th>Notes</th><th>Species range</th></tr></thead><tbody><tr><td colspan="5" style="text-align: center; font-weight: bold;">America</td></tr><tr><td>Canada</td><td>[Condylura cristata]</td><td>Micherdzinski, 1980</td><td></td><td></td></tr><tr><td rowspan="10">USA (including Hawaii)</td><td><strong>Snakes</strong></td><td></td><td></td><td></td></tr><tr><td>Heterodon platirhinos</td><td>Thomasson & Blouin-Demers, 2015</td><td></td><td>Native species</td></tr><tr><td><strong>Lizards</strong></td><td></td><td></td><td></td></tr><tr><td>Uta stansburiana</td><td>Goldberg & Bursey, 1991</td><td></td><td>Native species</td></tr><tr><td>Elgaria multicarinatus</td><td>DeNardo & Wozniak, 1997; Wozniak & DeNardo, 2000</td><td></td><td>Native species</td></tr><tr><td>Tiliqua scincoides</td><td>DeNardo & Wozniak, 1997; Wozniak & DeNardo, 2000</td><td></td><td>Non-native species</td></tr><tr><td><strong>Snakes</strong></td><td></td><td></td><td></td></tr><tr><td>Malayopython reticulatus</td><td>André 1937</td><td>As Python reticulatus</td><td>Non-native species</td></tr><tr><td>Boa constrictor</td><td>Radovsky 1971; DeNardo & Wozniak, 1997</td><td></td><td>Non-native species</td></tr><tr><td>Crotalus ruber</td><td>Schroeder, 1934</td><td></td><td>Native species</td></tr><tr><td rowspan="10">Mexico</td><td><strong>Lizards</strong></td><td></td><td></td><td></td></tr><tr><td>Thamnophis sirtalis infernalis</td><td>Schroeder, 1934</td><td>As C. confiuentus oreganus</td><td>Native species</td></tr><tr><td>Pituophis catenifer annectens</td><td>Schroeder, 1934</td><td></td><td>Native species</td></tr><tr><td>Drymarchon couperi</td><td>Schroeder, 1934</td><td></td><td>Native species</td></tr><tr><td>Antaresia maculosa</td><td>Reeves et al., 2006</td><td></td><td>Native species</td></tr><tr><td>Antaresia maculosa</td><td>Reeves et al., 2006</td><td></td><td>Native species</td></tr><tr><td><strong>Lizards</strong></td><td></td><td></td><td></td></tr><tr><td>Iguana iguana</td><td>Salazar, 2014</td><td></td><td>Native species</td></tr><tr><td><strong>Snakes</strong></td><td></td><td></td><td></td></tr><tr><td>Crotalus triseriatus</td><td>Paredes-Leon et al., 2008</td><td></td><td>Native species</td></tr><tr><td>Crotalus durissus</td><td>Rodriguez & Lazcano, 1992</td><td></td><td>Native species</td></tr><tr><td>Crotalus cerastes cercobombus</td><td>Rodriguez & Lazcano, 1992</td><td></td><td>Native species</td></tr><tr><td>Crotalus lepidus lepidus</td><td>Rodriguez & Lazcano, 1992</td><td></td><td>Native species</td></tr><tr><td>Crotalus atrox</td><td>Rodriguez & Lazcano, 1992</td><td></td><td>Native species</td></tr><tr><td>Crotalus ravus</td><td>Rodriguez & Lazcano, 1992</td><td></td><td>Native species</td></tr><tr><td>Pituophis deppei deppei</td><td>Rodriguez & Lazcano, 1992</td><td></td><td>Native species</td></tr><tr><td>Boa constrictor</td><td>Rodriguez & Lazcano, 1992</td><td></td><td>Native species</td></tr><tr><td>Gerrhonotus liocephalus</td><td>Rodriguez & Lazcano, 1992</td><td></td><td>Native species</td></tr><tr><td>Lampropeltis triangulum</td><td>Rodriguez & Lazcano, 1992</td><td></td><td>Native species</td></tr><tr><td>Barisia imbricate</td><td>Rodriguez & Lazcano, 1992</td><td></td><td>Native species</td></tr><tr><td>Barisia ciliaris</td><td>Rodriguez & Lazcano, 1992</td><td></td><td>Native species</td></tr><tr><td>Corytophanes cristatus</td><td>Rodriguez & Lazcano, 1992</td><td></td><td>Native species</td></tr><tr><td>Laemanchus serratus</td><td>Fuantos-Gámez et al., 2020</td><td></td><td>Native species</td></tr><tr><td>Python bivittatus</td><td>Fuantos-Gámez et al., 2020</td><td>As Python molurus bivittatus</td><td>Non-native species</td></tr><tr><td rowspan="10">Panama</td><td><strong>Turtles</strong></td><td></td><td></td><td></td></tr><tr><td>Staurotypus triporcatus</td><td>Paredes-León et al., 2008</td><td></td><td>Native species</td></tr><tr><td>Trachemys scripta elegans</td><td>Paredes-León et al., 2008</td><td></td><td>Native species</td></tr><tr><td>Trachemys venusta</td><td>Paredes-León et al., 2008</td><td></td><td>Native species</td></tr><tr><td><strong>Snakes</strong></td><td></td><td></td><td></td></tr><tr><td>Boa constrictor</td><td>Miranda et al., 2017</td><td></td><td>Native species</td></tr><tr><td>Epicerates maurus</td><td>Miranda et al., 2017</td><td></td><td>Native species</td></tr><tr><td>Corallus ruschenbergerii</td><td>Miranda et al., 2017</td><td></td><td>Native species</td></tr><tr><td>Corallus caninus</td><td>Miranda et al., 2017</td><td></td><td>Native species</td></tr><tr><td>Python regius</td><td>Miranda et al., 2017</td><td></td><td>Non-native species</td></tr><tr><td rowspan="2">Nicaragua</td><td><strong>Snakes</strong></td><td></td><td></td><td></td></tr><tr><td>Boa constrictor</td><td>Rimbaud et al., 2006</td><td></td><td>Non-native species</td></tr><tr><td rowspan="2">Antigua</td><td><strong>Snakes</strong></td><td></td><td></td><td></td></tr><tr><td>Alsophis antiguae</td><td>Daltry et al., 2001</td><td></td><td>Native species</td></tr><tr><td rowspan="6">Brazil</td><td>[Mesomys ecaudatus]</td><td>Ferreira da Silva et al., 2018</td><td></td><td></td></tr><tr><td><strong>Lizards</strong></td><td></td><td></td><td></td></tr><tr><td>Enyalius iheringii</td><td>Mendoza-Roldan et al., 2020b</td><td></td><td>Native species</td></tr><tr><td>Pogona vitticeps</td><td>Mendoza-Roldan et al., 2020b</td><td></td><td>Non-native species</td></tr><tr><td><strong>Snakes</strong></td><td></td><td></td><td></td></tr><tr><td>Boa constrictor constrictor</td><td>Barbosa et al., 2006</td><td>As Python reticulatus</td><td>Native species</td></tr><tr><td>Malayopython reticulatus</td><td>Fonseca, 1933; Micherdzinski, 1980</td><td></td><td>Non-native species</td></tr><tr><td>Corallus hortullana</td><td>Mendoza-Roldan et al., 2020b</td><td>As Corallus hortullanus</td><td>Native species</td></tr><tr><td>Crotalus durissus</td><td>Mendoza-Roldan et al., 2020b</td><td>As Crotalus durissus terrificus</td><td>Native species</td></tr><tr><td>Pantherophis guttatus</td><td>Almeida Pereira et al., 2019</td><td>As Pantherophis guttata</td><td>Native species</td></tr></tbody></table>
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).

<table><caption>**Table 2** Continued.</caption><thead><tr><th>Country</th><th>Host</th><th>References</th><th>Notes</th><th>Species range</th></tr></thead><tbody><tr><td colspan="5" style="text-align: center; font-weight: bold;">America</td></tr><tr><td>Canada</td><td>[Condylura cristata]</td><td>Micherdzinski, 1980</td><td></td><td></td></tr><tr><td rowspan="10">USA (including Hawaii)</td><td><strong>Snakes</strong></td><td></td><td></td><td></td></tr><tr><td>Heterodon platirhinos</td><td>Thomasson & Blouin-Demers, 2015</td><td></td><td>Native species</td></tr><tr><td><strong>Lizards</strong></td><td></td><td></td><td></td></tr><tr><td>Uta stansburiana</td><td>Goldberg & Bursey, 1991</td><td></td><td>Native species</td></tr><tr><td>Elgaria multicarinatus</td><td>DeNardo & Wozniak, 1997; Wozniak & DeNardo, 2000</td><td></td><td>Native species</td></tr><tr><td>Tiliqua scincoides</td><td>DeNardo & Wozniak, 1997; Wozniak & DeNardo, 2000</td><td></td><td>Non-native species</td></tr><tr><td><strong>Snakes</strong></td><td></td><td></td><td></td></tr><tr><td>Malayopython reticulatus</td><td>André 1937</td><td>As Python reticulatus</td><td>Non-native species</td></tr><tr><td>Boa constrictor</td><td>Radovsky 1971; DeNardo & Wozniak, 1997</td><td></td><td>Non-native species</td></tr><tr><td>Crotalus ruber</td><td>Schroeder, 1934</td><td></td><td>Native species</td></tr><tr><td>Crotalus oreganus</td><td>Schroeder, 1934</td><td>As C. confuientus oreganus</td><td>Native species</td></tr><tr><td>Masticophis flagellum</td><td>Schroeder, 1934</td><td></td><td>Native species</td></tr><tr><td>Thamnophis sirtalis infernalis</td><td>Schroeder, 1934</td><td></td><td>Native species</td></tr><tr><td>Thamnophis hammondii</td><td>Schroeder, 1934</td><td>As Thamnophis ordinoides hammondii</td><td>Native species</td></tr><tr><td>Pituophis catenifer annectens</td><td>Schroeder, 1934</td><td></td><td>Native species</td></tr><tr><td>Drymarchon couperi</td><td>Reeves et al., 2006</td><td></td><td>Native species</td></tr><tr><td>Antaresia maculosa</td><td>Reeves et al., 2006</td><td></td><td>Native species</td></tr><tr><td rowspan="20">Mexico</td><td><strong>Lizards</strong></td><td></td><td></td><td></td></tr><tr><td>Iguana iguana</td><td>Salazar, 2014</td><td></td><td>Native species</td></tr><tr><td><strong>Snakes</strong></td><td></td><td></td><td></td></tr><tr><td>Crotalus triseriatus</td><td>Paredes-Leon et al., 2008</td><td></td><td>Native species</td></tr><tr><td>Crotalus durissus</td><td>Rodriguez & Lazcano, 1992</td><td></td><td>Native species</td></tr><tr><td>Crotalus cerastes cercobombus</td><td>Rodriguez & Lazcano, 1992</td><td></td><td>Native species</td></tr><tr><td>Crotalus lepidus lepidus</td><td>Rodriguez & Lazcano, 1992</td><td></td><td>Native species</td></tr><tr><td>Crotalus atrox</td><td>Rodriguez & Lazcano, 1992</td><td></td><td>Native species</td></tr><tr><td>Crotalus ravus</td><td>Rodriguez & Lazcano, 1992</td><td></td><td>Native species</td></tr><tr><td>Pituophis deppei deppei</td><td>Rodriguez & Lazcano, 1992</td><td></td><td>Native species</td></tr><tr><td>Boa constrictor</td><td>Rodriguez & Lazcano, 1992</td><td></td><td>Native species</td></tr><tr><td>Gerrhonotus liocephalus</td><td>Rodriguez & Lazcano, 1992</td><td></td><td>Native species</td></tr><tr><td>Lampropeltis triangulum</td><td>Rodriguez & Lazcano, 1992</td><td></td><td>Native species</td></tr><tr><td>Barisia imbricate</td><td>Rodriguez & Lazcano, 1992</td><td></td><td>Native species</td></tr><tr><td>Barisia ciliaris</td><td>Rodriguez & Lazcano, 1992</td><td></td><td>Native species</td></tr><tr><td>Corytophanes cristatus</td><td>Rodriguez & Lazcano, 1992</td><td></td><td>Native species</td></tr><tr><td>Laemanchus serratus</td><td>Fuantos-Gámez et al., 2020</td><td></td><td>Native species</td></tr><tr><td>Python bivittatus</td><td>Fuantos-Gámez et al., 2020</td><td>As Python molurus bivittatus</td><td>Non-native species</td></tr><tr><td><strong>Turtles</strong></td><td></td><td></td><td></td></tr><tr><td>Staurotypus triporcatus</td><td>Paredes-León et al., 2008</td><td></td><td>Native species</td></tr><tr><td>Trachemys scripta elegans</td><td>Paredes-León et al., 2008</td><td></td><td>Native species</td></tr><tr><td>Trachemys venusta</td><td>Paredes-León et al., 2008</td><td></td><td>Native species</td></tr><tr><td rowspan="5">Panama</td><td><strong>Snakes</strong></td><td></td><td></td><td></td></tr><tr><td>Boa constrictor</td><td>Miranda et al., 2017</td><td></td><td>Native species</td></tr><tr><td>Epicerates maurus</td><td>Miranda et al., 2017</td><td></td><td>Native species</td></tr><tr><td>Corallus ruschenbergerii</td><td>Miranda et al., 2017</td><td></td><td>Native species</td></tr><tr><td>Corallus caninus</td><td>Miranda et al., 2017</td><td></td><td>Native species</td></tr><tr><td>Nicaragua</td><td><strong>Snakes</strong></td><td></td><td></td><td></td></tr><tr><td rowspan="2">Antigua</td><td>Boa constrictor</td><td>Rimbaud et al., 2006</td><td></td><td>Non-native species</td></tr><tr><td><strong>Snakes</strong></td><td></td><td></td><td></td></tr><tr><td rowspan="2">Brazil</td><td>Alsophis antiguae</td><td>Daltry et al., 2001</td><td></td><td>Native species</td></tr><tr><td>[Mesomys ecaudatus]</td><td>Ferreira da Silva et al., 2018</td><td></td><td></td></tr><tr><td colspan="5" style="text-align: center; font-weight: bold;">Lizards</td></tr><tr><td></td><td>Enyalius iheringii</td><td>Mendoza-Roldan et al., 2020b</td><td></td><td>Native species</td></tr><tr><td></td><td>Pogona vitticeps</td><td>Mendoza-Roldan et al., 2020b</td><td></td><td>Non-native species</td></tr><tr><td colspan="5" style="text-align: center; font-weight: bold;">Snakes</td></tr><tr><td></td><td>Boa constrictor constrictor</td><td>Barbosa et al., 2006</td><td>As Python reticulatus</td><td>Native species</td></tr><tr><td></td><td>Malayopython reticulatus</td><td>Fonseca, 1933; Micherdzinski, 1980</td><td></td><td>Non-native species</td></tr><tr><td></td><td>Corallus hortullana</td><td>Mendoza-Roldan et al., 2020b</td><td>As Corallus hortullanus</td><td>Native species</td></tr><tr><td></td><td>Crotalus durissus</td><td>Mendoza-Roldan et al., 2020b</td><td>As Crotalus durissus terrificus</td><td>Native species</td></tr><tr><td></td><td>Pantherophis guttatus</td><td>Almeida Pereira et al., 2019</td><td>As Pantherophis guttata</td><td>Native species</td></tr></tbody></table>
## 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,

<table><caption>**Table 2** Continued.</caption><thead><tr><th>Country</th><th>Host</th><th>References</th><th>Notes</th><th>Species range</th></tr></thead><tbody><tr><td colspan="5" style="text-align:center; font-weight:bold;">America</td></tr><tr><td rowspan="20">Argentina</td><td colspan="4" style="font-weight:bold;">Lizards</td></tr><tr><td>Leiosaurus bellii</td><td>Mauri, 1967</td><td rowspan="3">As Ophiodes striatus</td><td>Native species</td></tr><tr><td>Teious teyou</td><td>Mauri, 1967</td><td>Native species</td></tr><tr><td>Ophiodes fragilis</td><td>Mauri, 1967</td><td>Native species</td></tr><tr><td colspan="4" style="font-weight:bold;">Snakes</td></tr><tr><td>Epicrates cenchria crassus</td><td>Mauri, 1967</td><td rowspan="5">As Leimadophis poecilogyrus</td><td>Native species</td></tr><tr><td>Helicops leopardine</td><td>Mauri, 1967</td><td>Native species</td></tr><tr><td>Chironius carinatus</td><td>Mauri, 1967</td><td>Native species</td></tr><tr><td>Erythrolamrus poecilogyrus</td><td>Mauri, 1967</td><td>Native species</td></tr><tr><td>Xenodon merremii</td><td>Mauri, 1967</td><td>Native species</td></tr><tr><td>Lystrophis dorbignyi</td><td>Mauri, 1967</td><td rowspan="2">As Sibynomorphus turgidus</td><td>Native species</td></tr><tr><td>Lystrophis semcinctus</td><td>Mauri, 1967</td><td>Native species</td></tr><tr><td>Dipsas turgida</td><td>Mauri, 1967</td><td rowspan="2">As Pseudoboa cloelia</td><td>Native species</td></tr><tr><td>Imantodes cenchoa</td><td>Mauri, 1967</td><td>Native species</td></tr><tr><td>Leptodeira annulata annulata</td><td>Mauri, 1967</td><td rowspan="2">As Pseudoboa rhombifera</td><td>Native species</td></tr><tr><td>Clelia clelia</td><td>Mauri, 1967</td><td>Native species</td></tr><tr><td>Oxyrhopus rhombifer</td><td>Mauri, 1967</td><td rowspan="2">As Tomodon ocellatus</td><td>Native species</td></tr><tr><td>Tomodon dorsatus</td><td>Mauri, 1967</td><td>Native species</td></tr><tr><td>Tachymenis ocellata</td><td>Mauri, 1967</td><td rowspan="2">As Philodryas schotti</td><td>Native species</td></tr><tr><td>Philodryas baroni</td><td>Mauri, 1967</td><td>Native species</td></tr><tr><td>Philodryas patagoniensis</td><td>Mauri, 1967</td><td rowspan="10">As Philodryas schotti</td><td>Native species</td></tr><tr><td>Erhythrolamptus aesculapii</td><td>Mauri, 1967</td><td>Native species</td></tr><tr><td>Micrurus frontalis</td><td>Mauri, 1967</td><td>Native species</td></tr><tr><td>Micrurus lemniscatus</td><td>Mauri, 1967</td><td>Native species</td></tr><tr><td>Bothrops alternata</td><td>Mauri, 1967</td><td>Native species</td></tr><tr><td>Bothrops ammodythoides</td><td>Mauri, 1967</td><td>Native species</td></tr><tr><td>Bothrops jararacussu</td><td>Mauri, 1967</td><td>Native species</td></tr><tr><td>Bothrops neuwiedii meridionalis</td><td>Mauri, 1967</td><td>Native species</td></tr><tr><td>Crotalus terrificus terrificus</td><td>Mauri, 1967</td><td>Native species</td></tr><tr><td colspan="5" style="text-align:center; font-weight:bold;">Australia and Oceania</td></tr><tr><td rowspan="15">Australia</td><td colspan="4" style="font-weight:bold;">Lizards</td></tr><tr><td>Amphibolurus muricatus</td><td>Domrow, 1988</td><td rowspan="5">As Python molurus bivittatus</td><td>Native species</td></tr><tr><td>Carlia longipes</td><td>Domrow, 1988</td><td>Native species</td></tr><tr><td>Chlamydosaurus kingie</td><td>Domrow, 1988</td><td>Native species</td></tr><tr><td>Lacerta viridis</td><td>Domrow, 1988</td><td>Non-native species</td></tr><tr><td>Tiliqua rugosa</td><td>Norval et al., 2020, 2021</td><td>Native species</td></tr><tr><td>Tiliqua scincoides</td><td>Norval et al., 2020, 2021</td><td>Native species</td></tr><tr><td colspan="4" style="font-weight:bold;">Snakes</td></tr><tr><td>Acanthophis antarcticus</td><td>Norval et al., 2020, 2021</td><td rowspan="10">As Python molurus bivittatus</td><td>Native species</td></tr><tr><td>Antaresia children</td><td>Norval et al., 2020, 2021</td><td>Native species</td></tr><tr><td>Aspidites ramsayi</td><td>Norval et al., 2020, 2021</td><td>Native species</td></tr><tr><td>Hoplocephalus bitorquatus</td><td>Norval et al., 2020, 2021</td><td>Native species</td></tr><tr><td>Hoplocephalus stephensii</td><td>Norval et al., 2020, 2021</td><td>Native species</td></tr><tr><td>Liasis fuscus</td><td>Norval et al., 2020, 2021</td><td>Native species</td></tr><tr><td>Morelia viridis</td><td>Norval et al., 2020, 2021</td><td>Native species</td></tr><tr><td>Morelia spilota metcalfei</td><td>Norval et al., 2020, 2021</td><td>Native species</td></tr><tr><td>Notechis scutatus</td><td>Norval et al., 2020, 2021</td><td>Native species</td></tr><tr><td>Pseudechis porphyriacus</td><td>Norval et al., 2020, 2021</td><td>Native species</td></tr><tr><td>Pseudonaja textilis</td><td>Norval et al., 2020, 2021</td><td>Non-native species</td></tr><tr><td>Python bivittatus</td><td>Norval et al., 2020, 2021</td><td>Native species</td></tr><tr><td>Tropidechis carinatus</td><td>Norval et al., 2020, 2021</td><td>Native species</td></tr><tr><td rowspan="4">New Zealand</td><td colspan="4" style="font-weight:bold;">Lizards</td></tr><tr><td>Tiliqua scincoides</td><td>Heath, 1986</td><td rowspan="4">As Coluber viridiflavus viridiflavus</td><td>Non-native species</td></tr><tr><td colspan="4" style="font-weight:bold;">Snakes</td></tr><tr><td>Sceloporus graciosus graciosus</td><td>Radford, 1950</td><td>Native species</td></tr><tr><td>Hierophis viridiflavus</td><td>Radford, 1950</td><td>Native species</td></tr></tbody></table>
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

<table><caption>**Table 3** Records of *Ophionyssus natricis* in Russia.</caption><thead><tr><th>Date</th><th>Locality</th><th>Host</th><th>References</th><th>Notes</th><th>Species range</th></tr></thead><tbody><tr><td>1950</td><td>Moscow Zoo, host was captured in Middle Asia</td><td><strong>Snakes</strong></td><td></td><td></td><td></td></tr><tr><td rowspan="3">Undated</td><td rowspan="3">Moscow Zoo</td><td><em>Echis carinatus</em></td><td>Zemskaya, 1951</td><td></td><td>Non-native species</td></tr><tr><td><strong>Snakes</strong></td><td></td><td></td><td></td></tr><tr><td><em>Macrovipera lebetinus</em><br><em>Platiceps karelini</em></td><td>Bregetova, 1956</td><td>As <em>Vipera lebetina</em><br><em>As Coluber karelini</em></td><td>Native species<br>Non-native species</td></tr><tr><td>1954–1955</td><td>Damchik section, Astrakhan’ Nature Reserve, Astrakhan’ oblast’ (Lower Volga region)</td><td><strong>Snakes</strong></td><td></td><td></td><td></td></tr><tr><td rowspan="3">1959–1960</td><td rowspan="3">Manure pile, Cherniy Mys settlement, Kolyvan’ district, Novosibirsk oblast’ (Western Siberia)</td><td><em>Natrix natrix</em></td><td>Markov et al., 1964</td><td></td><td>Native species</td></tr><tr><td><em>Natrix tessellata</em></td><td></td><td></td><td>Native species</td></tr><tr><td><em>Elaphe dione</em></td><td></td><td></td><td>Native species</td></tr><tr><td>1973–1976</td><td>Bogdinsk-Baskunchak Nature Reserve, Astrakhan’ oblast’ (Lower Volga region)</td><td><strong>Snakes</strong></td><td></td><td></td><td></td></tr><tr><td>2005</td><td>Bogdinsk-Baskunchak Nature Reserve, Astrakhan’ oblast’ (Lower Volga region)</td><td><em>Natrix natrix</em></td><td>Belova & Grigoriev, 1981</td><td></td><td>Native species</td></tr><tr><td></td><td></td><td><strong>Snakes</strong></td><td></td><td></td><td></td></tr><tr><td>23–30 August 2008</td><td>South-Eastern part of Novosibirsk oblast’ (Western Siberia)</td><td><em>Dolichophis caspius</em></td><td>Bakiev, 2007</td><td></td><td>Native species</td></tr><tr><td rowspan="3">Undated</td><td rowspan="3">Moscow Zoo</td><td><em>Gloydius halys</em></td><td>Simonov & Zinchenko, 2010</td><td></td><td>Native species</td></tr><tr><td><strong>Snakes</strong></td><td></td><td></td><td></td></tr><tr><td><em>Simalia boeleni</em></td><td>Vasiliev & Balakina, undated</td><td>As <em>Morelia boeleni</em></td><td>Non-native species</td></tr><tr><td rowspan="5">April 2009</td><td rowspan="5">Moscow</td><td><strong>Lizards</strong></td><td></td><td></td><td></td></tr><tr><td><em>Darevskia brauneri</em></td><td>this paper, new record</td><td></td><td>Native species</td></tr><tr><td><em>Ophisops elegans</em></td><td>this paper, new record</td><td></td><td>Native species</td></tr><tr><td><strong>Snakes</strong></td><td></td><td></td><td></td></tr><tr><td><em>Python regius</em></td><td>this paper, new record</td><td></td><td>Non-native species</td></tr><tr><td></td><td><em>Telescopus fallax</em></td><td></td><td></td><td>Native species</td></tr></tbody></table>
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

<table><caption>**Table 3** Records of *Ophionyssus natricis* in Russia.</caption><thead><tr><th>Date</th><th>Locality</th><th>Host</th><th>References</th><th>Notes</th><th>Species range</th></tr></thead><tbody><tr><td>1950</td><td>Moscow Zoo, host was captured in Middle Asia</td><td><strong>Snakes</strong></td><td></td><td></td><td></td></tr><tr><td rowspan="3">Undated</td><td rowspan="3">Moscow Zoo</td><td><em>Echis carinatus</em></td><td>Zemskaya, 1951</td><td></td><td>Non-native species</td></tr><tr><td><strong>Snakes</strong></td><td></td><td></td><td></td></tr><tr><td><em>Macrovipera lebetinus</em><br><em>Platiceps karelini</em></td><td>Bregetova, 1956</td><td>As <em>Vipera lebetina</em><br><em>As Coluber karelini</em></td><td>Native species<br>Non-native species</td></tr><tr><td>1954–1955</td><td>Damchik section, Astrakhan’ Nature Reserve, Astrakhan’ oblast’ (Lower Volga region)</td><td><strong>Snakes</strong></td><td></td><td></td><td></td></tr><tr><td rowspan="3">1959–1960</td><td rowspan="3">Manure pile, Cherniy Mys settlement, Kolyvan’ district, Novosibirsk oblast’ (Western Siberia)</td><td><em>Natrix natrix</em></td><td>Markov et al., 1964</td><td></td><td>Native species</td></tr><tr><td><em>Natrix tessellata</em></td><td></td><td></td><td>Native species</td></tr><tr><td><em>Elaphe dione</em><br><strong>Snakes</strong></td><td></td><td></td><td>Native species</td></tr><tr><td>1973–1976</td><td>Bogdinsk-Baskunchak Nature Reserve, Astrakhan’ oblast’ (Lower Volga region)</td><td><em>Natrix natrix</em><br><strong>Snakes</strong></td><td>Belova & Grigoriev, 1981</td><td></td><td>Native species</td></tr><tr><td>2005</td><td>Bogdinsk-Baskunchak Nature Reserve, Astrakhan’ oblast’ (Lower Volga region)</td><td><em>Dolichophis caspius</em><br><strong>Snakes</strong></td><td>Bakiev, 2007</td><td></td><td>Native species</td></tr><tr><td rowspan="3">23–30 August 2008</td><td rowspan="3">South-Eastern part of Novosibirsk oblast’ (Western Siberia)</td><td><em>Gloydius halys</em></td><td>Simonov & Zinchenko, 2010</td><td></td><td>Native species</td></tr><tr><td><strong>Snakes</strong></td><td></td><td></td><td></td></tr><tr><td><em>Simalia boeleni</em><br><strong>Lizards</strong></td><td>Vasiliev & Balakina, undated</td><td>As <em>Morelia boeleni</em></td><td>Non-native species</td></tr><tr><td rowspan="3">April 2009</td><td rowspan="3">Moscow Zoo</td><td><em>Darevskia brauneri</em></td><td>this paper, new record</td><td></td><td>Native species</td></tr><tr><td><em>Ophisops elegans</em></td><td>this paper, new record</td><td></td><td>Native species</td></tr><tr><td><strong>Snakes</strong></td><td></td><td></td><td></td></tr><tr><td></td><td></td><td><em>Python regius</em></td><td>this paper, new record</td><td></td><td>Non-native species</td></tr><tr><td></td><td></td><td><em>Telescopus fallax</em></td><td>this paper, new record</td><td></td><td>Native species</td></tr></tbody></table>
(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*.
(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* 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
*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.

ORCID

Maria V. Orlova https://orcid.org/0000-0001-7811-4364

Bruce Halliday https://orcid.org/0000-0003-1909-998X

Will K. Reeves https://orcid.org/0000-0001-9025-5071

Igor V. Doronin https://orcid.org/0000-0003-1000-3144

Vladimir A. Mishchenko https://orcid.org/0000-0003-4280-283X

Ivan V. Vyalykh https://orcid.org/0000-0002-3123-8359

Artem A. Kidov https://orcid.org/0000-0001-9328-2470

References

Abdushukurova R.U., Markov G.S., Bogdanov O.P. 1966. On the parasitic fauna of the dice snake. Vertebrates of Central Asia (Collection of articles dedicated to the 60th anniversary of the academician of the Academy of Sciences of the Uzbek SSR T.Z. Zahidov). Tashkent, Fan Publishing House. p. 215-220. [In Russian]

Ahl I. 1994. Snake mites *Ophionyssus natricis* (Gervais 1844) more common than snakes in the terrarium? Litteratura Serpentium, 14: 9-24.

Alfonso-Toledo J.A., Paredes-León R. 2021. Molecular and morphological identification of dermanyssoid mites (Parasitiformes: Mesostigmata: Dermanyssoidea) causatives of a parasitic outbreak on captive snakes. Journal of Medical Entomology, 58: 246-251. https://doi.org/10.1093/jme/tjaa164

Almeida Pereira J., de Araújo Barreto L., Vianna C.A.B., de Oliveira Henriques M., de Sousa Oliveira Batista Cirne L.C. 2019. Diagnóstico e tratamento de serpentes *Pantherophis guttata* (corn snake) infestadas com *Ophionyssus natricis*: relato de caso. Revista Agrária Acadêmica, 3: 202-206. https://doi.org/10.32406/v2n32019/202-206/agrariacad

Amanatfard E., Youssefi M.R., Barimani A. 2014. Human dermatitis caused by *Ophionyssus natricis*, a snake mite. Iranian Journal of Parasitology, 9: 594-596.

André M. 1937. Sur l'Ophionyssus natricis Gervais, acarien parasite des reptiles. Bulletin du Muséum National d'Histoire Naturelle, 9: 62-65.

APBI Biological Control. 2021. TAURRUS Natural Enemies of reptile mites. https://www.taurrus.co.uk/. Version dated 2021, date of access 19 October 2023.

Arutunjan E.S., Ohandjanian A.M. 1983. Parasitic gamasid mites from the Sevan Basin. Academy of Science of Armenian SSR, Institute of Zoology, Zoological Papers, 19: 300-318. [In Russian]

Ávila R.W., Ferreira V.L., Arruda, J.A.O. 2006. Natural history of the South American water snake *Helicops leopardinus* (Colubridae: Hydropsini) in the Pantanal, Central Brazil. Journal of Herpetology, 40: 274-279. https://doi.org/10.1670/113-05N.1

Baker E.W., Evans T.M., Gould D.J., Hull W.B., Keegan H.L. 1956. A Manual of Parasitic Mites of Medical or Economic Importance. National Pest Control Association, New York. pp. 170.

Bakiev A.G. 2007. Overall results of snakes' parasites investigation in the Volga River Basin. Communicate 2: Parasitiformes. Bulletin of Mordovia University, Biological Sciences, 4: 68-70. [In Russian]

Banks, N. 1914. New Acarina. Journal of Entomology and Zoology, 6: 55-66.

Bannert, B., Karaca H.Y., Wohltmann A. 2000. Life cycle and parasitic interaction of the lizard-parasitizing mite *Ophionyssus galloticolus* (Acari: Gamasida: Macronyssidae), with remarks about the evolutionary consequences of parasitism in mites. Experimental and Applied Acarology, 24: 597-613. https://doi.org/10.1023/A:1026504627926

Barbosa A.R., Silva H., Albuquerque H.N., Ribeiro I.A.M. 2006. Contribuição ao estudo parasitológico de jibóias, *Boa constrictor constrictor* Linnaeus, 1758, em cativeiro. Revista de Biologia e Ciências da Terra, 6: 1-19.

Beck W. 1996. Tierische Milben als Epizoonoseerreger und ihre Bedeutung in der Dermatologie. Der Hautarzt, 47: 744-748. https://doi.org/10.1007/s001050050501

Beck W., Pantchev N. 2006. Schlangenmilbenbefall (*Ophionyssus natricis*) beim Grünen Leguan (*Iguana iguana*) - Ein Fallbericht. Kleintierpraxis, 51: 648-652.

Beck W., Pfister N. 2006. Humanpathogene Milben als Zoonoseerreger. Wiener Klinische Wochenschrift, 118, Supplement 3, 27-32. https://doi.org/10.1007/s00508-006-0678-y

Belova O.S., Grigoriev O.V. 1981. Occurrence of gamasid and ixodid ticks of reptiles of Western Siberia. In: Borkin L.J. (ed.) Herpetological Investigations in Siberia and the Far East. Leningrad: Zoological Institute of the USSR Academy of Sciences. p. 16-18.

Beron P. 1966. Contribution à l'étude des acariens parasites des reptiles en Bulgarie. Bulletin de l'Institut de Zoologie et Musée, 22: 51-53. [in Bulgarian]

Beron P. 2014. Acarorum Catalogus. III. Parasitiformes: Opilioacarida, Holothyrida, Mesostigmata. Sofia: Pensoft. pp. 286.

Biological Services. 2015. Snake/reptile mite *Ophionyssus natricis*. Available from https://biologicalservices.com.au/pests/snake-reptile-mite-92.html. [13 Oct 2023].
University, Russia) for his help with pictures. Authors contributed equally to this work.

ORCID

Maria V. Orlova https://orcid.org/0000-0001-7811-4364

Bruce Halliday https://orcid.org/0000-0003-1909-998X

Will K. Reeves https://orcid.org/0000-0001-9025-5071

Igor V. Doronin https://orcid.org/0000-0003-1000-3144

Vladimir A. Mishchenko https://orcid.org/0000-0003-4280-283X

Ivan V. Vyalykh https://orcid.org/0000-0002-3123-8359

Artem A. Kidov https://orcid.org/0000-0001-9328-2470

References

Abdushukurova R.U., Markov G.S., Bogdanov O.P. 1966. On the parasitic fauna of the dice snake. Vertebrates of Central Asia (Collection of articles dedicated to the 60th anniversary of the academician of the Academy of Sciences of the Uzbek SSR T.Z. Zahidov). Tashkent, Fan Publishing House. p. 215-220. [In Russian]

Ahl I. 1994. Snake mites *Ophionyssus natricis* (Gervais 1844) more common than snakes in the terrarium? Litteratura Serpentium, 14: 9-24.

Alfonso-Toledo J.A., Paredes-León R. 2021. Molecular and morphological identification of dermanyssoid mites (Parasitiformes: Mesostigmata: Dermanyssoidea) causatives of a parasitic outbreak on captive snakes. Journal of Medical Entomology, 58: 246-251. https://doi.org/10.1093/jme/tjaa164

Almeida Pereira J., de Araújo Barreto L., Vianna C.A.B., de Oliveira Henriques M., de Sousa Oliveira Batista Cirne L.C. 2019. Diagnóstico e tratamento de serpentes *Pantherophis guttata* (corn snake) infestadas com *Ophionyssus natricis*: relato de caso. Revista Agrária Acadêmica, 3: 202-206. https://doi.org/10.32406/v2n32019/202-206/agrariacad

Amanatfard E., Youssefi M.R., Barimani A. 2014. Human dermatitis caused by *Ophionyssus natricis*, a snake mite. Iranian Journal of Parasitology, 9: 594-596.

André M. 1937. Sur l'Ophionyssus natricis Gervais, acarien parasite des reptiles. Bulletin du Muséum National d'Histoire Naturelle, 9: 62-65.

APBI Biological Control. 2021. TAURRUS Natural Enemies of reptile mites. https://www.taurrus.co.uk/. Version dated 2021, date of access 19 October 2023.

Arutunjan E.S., Ohandjanian A.M. 1983. Parasitic gamasid mites from the Sevan Basin. Academy of Science of Armenian SSR, Institute of Zoology, Zoological Papers, 19: 300-318. [In Russian]

Ávila R.W., Ferreira V.L., Arruda, J.A.O. 2006. Natural history of the South American water snake *Helicops leopardinus* (Colubridae: Hydropsini) in the Pantanal, Central Brazil. Journal of Herpetology, 40: 274-279. https://doi.org/10.1670/113-05N.1

Baker E.W., Evans T.M., Gould D.J., Hull W.B., Keegan H.L. 1956. A Manual of Parasitic Mites of Medical or Economic Importance. National Pest Control Association, New York. pp. 170.

Bakiev A.G. 2007. Overall results of snakes' parasites investigation in the Volga River Basin. Communicate 2: Parasitiformes. Bulletin of Mordovia University, Biological Sciences, 4: 68-70. [In Russian]

Banks, N. 1914. New Acarina. Journal of Entomology and Zoology, 6: 55-66.

Bannert, B., Karaca H.Y., Wohltmann A. 2000. Life cycle and parasitic interaction of the lizard-parasitizing mite *Ophionyssus galloticolus* (Acari: Gamasida: Macronyssidae), with remarks about the evolutionary consequences of parasitism in mites. Experimental and Applied Acarology, 24: 597-613. https://doi.org/10.1023/A:1026504627926

Barbosa A.R., Silva H., Albuquerque H.N., Ribeiro I.A.M. 2006. Contribuição ao estudo parasitológico de jibóias, *Boa constrictor constrictor* Linnaeus, 1758, em cativeiro. Revista de Biologia e Ciências da Terra, 6: 1-19.

Beck W. 1996. Tierische Milben als Epizoonoseerreger und ihre Bedeutung in der Dermatologie. Der Hautarzt, 47: 744-748. https://doi.org/10.1007/s001050050501

Beck W., Pantchev N. 2006. Schlangenmilbenbefall (*Ophionyssus natricis*) beim Grünen Leguan (*Iguana iguana*) - Ein Fallbericht. Kleintierpraxis, 51: 648-652.

Beck W., Pfister N. 2006. Humanpathogene Milben als Zoonoseerreger. Wiener Klinische Wochenschrift, 118, Supplement 3, 27-32. https://doi.org/10.1007/s00508-006-0678-y

Belova O.S., Grigoriev O.V. 1981. Occurrence of gamasid and ixodid ticks of reptiles of Western Siberia. In: Borkin L.J. (ed.) Herpetological Investigations in Siberia and the Far East. Leningrad: Zoological Institute of the USSR Academy of Sciences. p. 16-18.

Beron P. 1966. Contribution à l'étude des acariens parasites des reptiles en Bulgarie. Bulletin de l'Institut de Zoologie et Musée, 22: 51-53. [in Bulgarian]

Beron P. 2014. Acarorum Catalogus. III. Parasitiformes: Opilioacarida, Holothyrida, Mesostigmata. Sofia: Pensoft. pp. 286.

Biological Services. 2015. Snake/reptile mite *Ophionyssus natricis*. Available from: https://biologicalservices.com.au/pests/snake-reptile-mite-92.html. [13 Oct 2023].
Blanc G., Ascione L. 1959. Sur la présence d' *Ophionyssus natricis* (Gervais 1844) sur trois serpents du Maroc des Forets de Nefifik et du Cherrat. Archives de l'Institut Pasteur du Maroc, 5: 666-668.

Bogdanov O.P. 1965. Ecology of reptiles in Middle Asia. Tashkent, Science. p. 213. [in Russian]

Bregetova N.G. 1956. Gamasid mites (Gamasoidea), a brief guide. Moscow: Zoological Institute, USSR Academy of Sciences. p. 247. [in Russian]

Buitendijk A.M. 1945. Voorloopige catalogus van de Acari in die collectie-Oudemans. Zoologische Mededelingen, 24: 281-391.

Burrage B.R. 1966. Observations on the macronyssid mite (order Acarina), *Ophionyssus natricis* (Gervais), on the two iguanid lizards, *Uta stansburiana hesperis* and *Sceloporus occidentalis occidentalis*. British Journal of Herpetology, 3: 275-278.

Camin J.H. 1948. Mite trasmission of a hemorrhagic septicemia in snakes. Journal of Parasitology, 34: 345-354. https://doi.org/10.2307/3273698

Camin J.H. 1949. An attempt to clarify the status of the species in the genus *Ophionyssus* Mégnin (Acarina: Macronyssidae). Journal of Parasitology, 35: 583-589. https://doi.org/10.2307/3273637

Camin J.H. 1953. Observations on the life history and sensory behaviour of the snake mite, *Ophionyssus natricis* (Gervais) (Acarina: Macronyssidae). Special Publications, The Chicago Academy of Sciences, 10: 1-75 + Plates I-III.

Camin J.H., Clarke G.K., Goodson, L.H., Shuyler, H.R. 1964. Control of the snake mite, *Ophionyssus natricis* (Gervais), in captive reptile collections. Zoologica: Scientific Contributions of the New York Zoological Society, 49: 65-79. https://doi.org/10.5962/p.203293

Castro P.D.J. 2019. Ectoparasites in captive reptiles. Avaiable from https://www.theveterinarynurse.com/review/article/ectoparasites-in-captive-reptiles [26 June 2023] https://doi.org/10.12968/vetn.2019.10.1.33

Cervone M., Fichi G., Lami A., Lanza A., Damiani G.M., Perucci S. 2016. Internal and external parasitic infections of pet reptiles in Italy. Journal of Herpetological Medicine and Surgery, 26: 122-130. https://doi.org/10.5818/1529-9651.26.3.4.122

Chang L., Jacobson E.R. 2010. Inclusion body disease, a worldwide infectious disease of booid snakes: a review. Journal of Exotic Pet Medicine, 19: 216-225. https://doi.org/10.1053/j.jepm.2010.07.014

Chiodini R.J., Holdeman L.V., Gandelman A.L. 1983. Infectious necrotic hepatitis caused by an unclassified anaerobic bacillus in the water snake. Journal of Comparative Pathology, 93: 235-242. https://doi.org/10.1016/0021-9975(83)90010-5

Costa M. 1966. The present stage of knowledge of mesostigmatic mites in Israel (Acari, Mesostigmata). Israel Journal of Zoology, 15: 69-82.

Daltry J.C., Bloxam Q., Cooper G., Day M.L., Hartley J., Henry M., Lindsay K., Smith B.E. 2001. Five years of conserving the 'world's rarest snake', the Antiguan racer *Alsophis antiguae*. Oryx, 35: 119-127. https://doi.org/10.1046/j.1365-3008.2001.00169.x

DeNardo D.F., Wozniak E.J. 1997. Understanding the snake mite and current therapies for its control. Proceedings of the Fourth Annual Conference of the Association of Reptilian and Amphibian Veterinarians, 1997, pp. 137-147.

Dhooria M.S. 2016. Fundamentals of Applied Acarology. Singapore: Springer. pp. 470. https://doi.org/10.1007/978-981-10-1594-6

Dik B. 2012. Türkiye'de Bir Su Yıldanında (*Natrix tessellata*, Laurente 1768) (Reptilia: Squamata: Colubridae) ilk *Ophionyssus natricis* (Gervais, 1844) Olgusu. Turkiye Parazitolji Dergisi, 36: 112-115. https://doi.org/10.5152/tpd.2012.27

Dipineto L., Raia P., Varriale L., Borrelli L., Botta V., Serio C., Capasso M., Rinaldi L. 2018. Bacteria and parasites in *Podarcis sicula* and *P. sicula klemmerii*. BMC Veterinary Research, 14: 392. https://doi.org/10.1186/s12917-018-1708-5

Divers S.J., Stahl S.J. 2019. Mader's Reptile And Amphibian Medicine And Surgery, Third Edition, Saint Louis: Elsevier. pp. 1537.

Domrow R. 1963. New records and species of Austromalayan laelapid mites. Proceedings of the Linnean Society of New South Wales, 88: 199-220

Domrow R. 1974. Miscellaneous mites from Australian vertebrates. Proceedings of the Linnean Society of New South Wales, 99: 15-35.

Domrow R. 1985. Species of *Ophionyssus* Mégnin from Australian lizards and snakes (Acari: Dermanyssidae). Journal of the Australian Entomological Society, 24: 149-153. https://doi.org/10.1111/j.1440-6055.1985.tb00213.x

Domrow R. 1988. Acari Mesostigmata parasitic on Australian vertebrates: an annotated checklist, keys and bibliography. Invertebrate Taxonomy, 1: 817-948. https://doi.org/10.1071/IT9870817

Domrow R., Heath, A.C.G., Kennedy, C. 1980. Two new species of *Ophionyssus* (Acari : Dermanyssidae) from New Zealand lizards. New Zealand Journal of Zoology, 7: 291-297. https://doi.org/10.1080/03014223.1980.10423790

Dugès A. 1834. Recherches sur l'Ordre des Acariens. Annales des Sciences Naturelles, 2: 18-63 + Plates 7-8.

Er-Rguibi O., Laghzaoui E.-L., Aglagane A., Kimdil L., Stekolnikov A.A., Abbad A., El Mouden E.H. 2023. New locality and host records of mites and ticks (Chelicerata: Acari) parasitizing lizards of Morocco. Acarologia, 63: 464-479. https://doi.org/10.24349/j4lz-jxdk

Evans G.O., Till W.M. 1966. Studies on the British Dermanyssidae (Acari: Mesostigmata). Part II Classification. Bulletin of the British Museum (Natural History) Zoology, 14: 109-370.

Fain A. 1962. Les acariens mesostigmatiques ectoparasites des serpents. Bulletin de l'Institut Royal des Sciences Naturelles de Belgique, 38: 1-149.

Fain, A., Bannert B. 2000. Two new species of *Ophionyssus* Mégnin (Acari: Macronyssidae) parasitic on lizards of the genus *Gallotia* Boulenger (Reptilia: Lacertidae) from the Canary Islands. International Journal of Acarology, 26: 41-50. https://doi.org/10.1080/01647950008683634
Blanc G., Ascione L. 1959. Sur la présence d' *Ophionyssus natricis* (Gervais 1844) sur trois serpents du Maroc des Forets de Nefifik et du Cherrat. Archives de l'Institut Pasteur du Maroc, 5: 666-668.

Bogdanov O.P. 1965. Ecology of reptiles in Middle Asia. Tashkent, Science. p. 213. [in Russian]

Bregetova N.G. 1956. Gamasid mites (Gamasoidea), a brief guide. Moscow: Zoological Institute, USSR Academy of Sciences. p. 247. [in Russian]

Buitendijk A.M. 1945. Voorloopige catalogus van de Acari in die collectie-Oudemans. Zoologische Mededelingen, 24: 281-391.

Burrage B.R. 1966. Observations on the macronyssid mite (order Acarina), *Ophionyssus natricis* (Gervais), on the two iguanid lizards, *Uta stansburiana hesperis* and *Sceloporus occidentalis occidentalis*. British Journal of Herpetology, 3: 275-278.

Camin J.H. 1948. Mite trasmission of a hemorrhagic septicemia in snakes. Journal of Parasitology, 34: 345-354. https://doi.org/10.2307/3273698

Camin J.H. 1949. An attempt to clarify the status of the species in the genus *Ophionyssus* Mégnin (Acarina: Macronyssidae). Journal of Parasitology, 35: 583-589. https://doi.org/10.2307/3273637

Camin J.H. 1953. Observations on the life history and sensory behaviour of the snake mite, *Ophionyssus natricis* (Gervais) (Acarina: Macronyssidae). Special Publications, The Chicago Academy of Sciences, 10: 1-75 + Plates I-III.

Camin J.H., Clarke G.K., Goodson, L.H., Shuyler, H.R. 1964. Control of the snake mite, *Ophionyssus natricis* (Gervais), in captive reptile collections. Zoologica: Scientific Contributions of the New York Zoological Society, 49: 65-79. https://doi.org/10.5962/p.203293

Castro P.D.J. 2019. Ectoparasites in captive reptiles. Avaiable from https://www.theveterinarynurse.com/review/article/ectoparasites-in-captive-reptiles [26 June 2023] https://doi.org/10.12968/vetn.2019.10.1.33

Cervone M., Fichi G., Lami A., Lanza A., Damiani G.M., Perucci S. 2016. Internal and external parasitic infections of pet reptiles in Italy. Journal of Herpetological Medicine and Surgery, 26: 122-130. https://doi.org/10.5818/1529-9651.26.3.4.122

Chang L., Jacobson E.R. 2010. Inclusion body disease, a worldwide infectious disease of booid snakes: a review. Journal of Exotic Pet Medicine, 19: 216-225. https://doi.org/10.1053/j.jepm.2010.07.014

Chiodini R.J., Holdeman L.V., Gandelman A.L. 1983. Infectious necrotic hepatitis caused by an unclassified anaerobic bacillus in the water snake. Journal of Comparative Pathology, 93: 235-242. https://doi.org/10.1016/0021-9975(83)90010-5

Costa M. 1966. The present stage of knowledge of mesostigmatic mites in Israel (Acari, Mesostigmata). Israel Journal of Zoology, 15: 69-82.

Daltry J.C., Bloxam Q., Cooper G., Day M.L., Hartley J., Henry M., Lindsay K., Smith B.E. 2001. Five years of conserving the 'world's rarest snake', the Antiguan racer *Alsophis antiguae*. Oryx, 35: 119-127. https://doi.org/10.1046/j.1365-3008.2001.00169.x

DeNardo D.F., Wozniak E.J. 1997. Understanding the snake mite and current therapies for its control. Proceedings of the Fourth Annual Conference of the Association of Reptilian and Amphibian Veterinarians, 1997, pp. 137-147.

Dhooria M.S. 2016. Fundamentals of Applied Acarology. Singapore: Springer. pp. 470. https://doi.org/10.1007/978-981-10-1594-6

Dik B. 2012. Türkiye'de Bir Su Yıldanında (*Natrix tessellata*, Laurente 1768) (Reptilia: Squamata: Colubridae) ilk *Ophionyssus natricis* (Gervais, 1844) Olgusu. Turkiye Parazitolji Dergisi, 36: 112-115. https://doi.org/10.5152/tpd.2012.27

Dipineto L., Raia P., Varriale L., Borrelli L., Botta V., Serio C., Capasso M., Rinaldi L. 2018. Bacteria and parasites in *Podarcis sicula* and *P. sicula klemmerii*. BMC Veterinary Research, 14: 392. https://doi.org/10.1186/s12917-018-1708-5

Divers S.J., Stahl S.J. 2019. Mader's Reptile And Amphibian Medicine And Surgery, Third Edition, Saint Louis: Elsevier. pp. 1537.

Domrow R. 1963. New records and species of Austromalayan laelapid mites. Proceedings of the Linnean Society of New South Wales, 88: 199-220

Domrow R. 1974. Miscellaneous mites from Australian vertebrates. Proceedings of the Linnean Society of New South Wales, 99: 15-35.

Domrow R. 1985. Species of *Ophionyssus* Mégnin from Australian lizards and snakes (Acari: Dermanyssidae). Journal of the Australian Entomological Society, 24: 149-153. https://doi.org/10.1111/j.1440-6055.1985.tb00213.x

Domrow R. 1988. Acari Mesostigmata parasitic on Australian vertebrates: an annotated checklist, keys and bibliography. Invertebrate Taxonomy, 1: 817-948. https://doi.org/10.1071/IT9870817

Domrow R., Heath, A.C.G., Kennedy, C. 1980. Two new species of *Ophionyssus* (Acari : Dermanyssidae) from New Zealand lizards. New Zealand Journal of Zoology, 7: 291-297. https://doi.org/10.1080/03014223.1980.10423790

Dugès A. 1834. Recherches sur l'Ordre des Acariens. Annales des Sciences Naturelles, 2: 18-63 + Plates 7-8.

Er-Rguibi O., Laghzaoui E.-L., Aglagane A., Kimdil L., Stekolnikov A.A., Abbad A., El Mouden E.H. 2023. New locality and host records of mites and ticks (Chelicerata: Acari) parasitizing lizards of Morocco. Acarologia, 63: 464-479. https://doi.org/10.24349/j4lz-jxdk

Evans G.O., Till W.M. 1966. Studies on the British Dermanyssidae (Acari: Mesostigmata). Part II Classification. Bulletin of the British Museum (Natural History) Zoology, 14: 109-370.

Fain A. 1962. Les acariens mesostigmatiques ectoparasites des serpents. Bulletin de l'Institut Royal des Sciences Naturelles de Belgique, 38: 1-149.

Fain, A., Bannert B. 2000. Two new species of *Ophionyssus* Mégnin (Acari: Macronyssidae) parasitic on lizards of the genus *Gallotia* Boulenger (Reptilia: Lacertidae) from the Canary Islands. International Journal of Acarology, 26: 41-50. https://doi.org/10.1080/01647950008683634
Feider Z., Solomon L. 1963. *Ophionyssus natricis* (Gervais) 1844 (Acari) o nouă specie pentru R.P.R. parazită pe şerpi. Studii şi Cercetări Științifice Academia R.P. Române Filiala Iași. Biologie și Științe Agricole, 14: 281-286.

Ferreira da Silva A., Teixeira Pinto Z., Hidalgo Teixeira R., Armando Cunha R., Carriço C., Leal Caetano R., Salles Gazeta G., Amorim M. 2018. First record of *Ophionyssus natricis* (Gervais) (Acari: Macronyssidae) on *Python reticulatus* (Schneider) (Pythonidae) in Brazil. EntomoBrasilis, 11: 41-44.
https://doi.org/10.12741/ebrasilis.v11i1.768

Fitzgerald K.T. 2019. Acariasis. In: Divers S.J., Stahl S.J. (Eds.). Mader's Reptile and Amphibian Medicine and Surgery, third Edition. Saint Louis: Elsevier. p. 1290-1291. https://doi.org/10.1016/B978-0-323-48253-0-00139-2

Fitzgerald K.T., Vera R. 2006. Acariasis. In: Mader D.R. (Ed.). Reptile Medicine and Surgery. Saint Louis: Elsevier. p. 720-738. https://doi.org/10.1016/B0-72-169327-X/50047-X

Fonseca F. 1933. Presença do *Ophionyssus serpentium* (Hirst, 1915) (Acarina, Dermanyssidae) no serpentaria do Instituto Butantan. Memórias do Instituto Butantan, 7: 145-146.

Fonseca F. 1948. A monograph of the genera and species of Macronyssidae Oudemans, 1936 (synom.: Liponissidae Vitzthum, 1931) (Acari). Proceedings of the Zoological Society of London, 118: 249-334. https://doi.org/10.1111/j.1096-3642.1948.tb00378.x

Fonseca F. 1954. Notas de acarologia. XXXVI. Aquisições novas para a fauna Brasileira de ácaros haematófagos (Acari, Macronyssidae). Revista Brasileira de Entomologia, 1: 79-92.

Fuantos-Gámez B.A, Romero-Núñez C., Sheinberg-Waisburd G., Bautista-Gómez L.G, Yarto-Jaramillo E., Heredia-Cardenas R., Miranda-Contreras L. 2020. Successful treatment of *Ophionyssus natricis* with afoxolaner in two Burmese pythons (*Python molurus bivittatus*). Veterinary Dermatology, 31: 496-498. https://doi.org/10.1111/vde.12898

Garrett C.M., Harwell G. 1991. Loreal pit impaction in a black speckled palm-pitviper (*Bothriechis nigroviridis*). Journal of Zoo and Wildlife Medicine, 22: 249-251.

Gervais F.L.P. 1844. Acarides. In: Walckenaer C.A. (Ed.). Histoire Naturelle des Insectes. Aptères. III. Paris: Roret. p. 132-288.

Gobble J. 2022. Oral Fluralaner (Bravecto®) use in the control of mites in 20 Ball Pythons (*Python regius*). Journal of Herpetological Medicine and Surgery, 32: 111-112. https://doi.org/10.5818/JHMS-D-21-00003

Goldberg S.R., Bursey C.R. 1991. Integumental lesions caused by ectoparasites in a wild population of the Side-blotched lizard *Uta stansburiana*. Journal of Wildlife Diseases, 27: 68-73. https://doi.org/10.7589/0090-3558-27.1.68

Gomes-Almeida, B.K., Pepato, A.R. 2021. A new genus and new species of macronyssid mite (Mesostigmata: Gamasina: Macronyssidae) from Brazilian caves including molecular data and key for genera occurring in Brazil. Acarologia, 61: 501-526. https://doi.org/10.24349/acarologia/2021447

Hallas T.E. 1978. Fortegnelse over Danske mider (Acari). Entomologiske Meddelelser, 46: 27-45.

Hassl A.R. 2016. Ticks and mites parasitizing free-ranging reptiles in Austria - with an identification key to Central European herpetophagous Acarina. Herpetozoa, 29: 77-83.

Heath A.C.G. 1986. First occurrence of the reptile mite, *Ophionyssus natricis* (Acari: Dermanyssidae) in New Zealand. New Zealand Veterinary Journal, 34: 78. https://doi.org/10.1080/00480169.1986.35304

Huchzermeyer E.W. 2003. Crocodiles. Biology, Husbandry and Diseases. Wallingford: CAB Publishing. pp. 353. https://doi.org/10.1079/9780851996561.0000

Jacobson E.R. 1984. Immobilization, blood sampling, necropsy techniques and diseases of crocodilians: a review. Journal of Zoo Animal Medicine, 15: 38-45. https://doi.org/10.2307/20094678

Keegan, H.L. 1956. Ectoparasitic lealaptid and dermanyssid mites of Egypt, Kenya and: the Sudan, primarily based on Namru 3 Collections, 1948 - 1953. Journal of the Egyptian Public Health Association, 31: 199-272 + Plates I-XIII.

Keskin A. 2021. Occurrence of *Ophionyssus natricis* (Acari: Macronyssidae) on the captive corn snake, *Pantherophis guttatus*, (Squamata: Colubridae) in Turkey. Acarological Studies, 3: 89-95.
https://doi.org/10.47121/acarolstud.907114

Khaire A., Khaire N., Joshi P.V., Bhat, H.R. 1987. An outbreak of infestation of *Ophionyssus natricis* (Gervais, 1844) in the snake park at Poona, India. The Snake, 19: 70-71.

Leslie A.J., Lovely C.J., Pittman J.M. 2011. A preliminary disease survey in the wild Nile crocodile (*Crocodylus niloticus*) population in the Okavango Delta, Botswana. Journal of the South African Veterinary Association, 82: 155-159. https://doi.org/10.4102/jsava.v82i3.54

Ma L.-M., Bai X.-L. 2012. Discovery of the genus *Ophionyssus* Mégnin in China and description of a new species (Acari: Mesostigmata: Macronyssidae). Acta Archnologica Sinica, 21: 83-86.

Magnusson W.E. 1985. Habitat selection, parasites and injuries in Amazonian crocodilians. Amazoniana, 9: 193-204.

Manoj R.R.S., Latrofa M.S., Mendoza-Roldan J.A., Otranto D. 2021. Molecular detection of *Wolbachia* endosymbiont in reptiles and their ectoparasites. Parasitology Research, 120: 3255-3261. https://doi.org/10.1007/s00436-021-07237-1

Mariana A., Vellayan S., Halimaton I., Ho T.M. 2011. Acariasis on pet Burmese python, *Python molurus bivittatus* in Malaysia. Asian Pacific Journal of Tropical Medicine, 4: 227-228. https://doi.org/10.1016/S1995-7645(11)60075-8

Markov G.S., Bogdanov O.P. 1960. Helminths and mites – parasites of snakes in Central Asia. Uzbek biological journal, 1: 35-41.

Markov G.S., Ivanov V.P., Kryuchkov G.P., Lukianova Zh.F., Nikulin V.P., Chernobay V.F. 1964. Protozoan and parasitic mites of reptiles from the Caspian region. Scientific notes of the Volgograd State Pedagogical Institute Named after A.S. Serafimovich, 16: 106-110. [In Russian]

Maslova I.V., Dochevoy Y.E. 2016. A case of protocooperation between the pitviper *Gloydius ussuriensis* and the ground beetle *Carabus granulatus telluris*. Russian Journal of Herpetology, 23: 231-234.
https://doi.org/10.30906/1026-2296-2016-23-3-231-234
Feider Z., Solomon L. 1963. *Ophionyssus natricis* (Gervais) 1844 (Acari) o nouă specie pentru R.P.R. parazită pe şerpi. Studii şi Cercetări Științifice Academia R.P. Române Filiala Iași. Biologie și Științe Agricole, 14: 281-286.

Ferreira da Silva A., Teixeira Pinto Z., Hidalgo Teixeira R., Armando Cunha R., Carriço C., Leal Caetano R., Salles Gazeta G., Amorim M. 2018. First record of *Ophionyssus natricis* (Gervais) (Acari: Macronyssidae) on *Python reticulatus* (Schneider) (Pythonidae) in Brazil. EntomoBrasilis, 11: 41-44.
https://doi.org/10.12741/ebrasilis.v11i1.768

Fitzgerald K.T. 2019. Acariasis. In: Divers S.J., Stahl S.J. (Eds.). Mader's Reptile and Amphibian Medicine and Surgery, third Edition. Saint Louis: Elsevier. p. 1290-1291. https://doi.org/10.1016/B978-0-323-48253-0-00139-2

Fitzgerald K.T., Vera R. 2006. Acariasis. In: Mader D.R. (Ed.). Reptile Medicine and Surgery. Saint Louis: Elsevier. p. 720-738. https://doi.org/10.1016/B0-72-169327-X/50047-X

Fonseca F. 1933. Presença do *Ophionyssus serpentium* (Hirst, 1915) (Acarina, Dermanyssidae) no serpentaria do Instituto Butantan. Memórias do Instituto Butantan, 7: 145-146.

Fonseca F. 1948. A monograph of the genera and species of Macronyssidae Oudemans, 1936 (synom.: Liponissidae Vitzthum, 1931) (Acari). Proceedings of the Zoological Society of London, 118: 249-334. https://doi.org/10.1111/j.1096-3642.1948.tb00378.x

Fonseca F. 1954. Notas de acarologia. XXXVI. Aquisições novas para a fauna Brasileira de ácaros haematófagos (Acari, Macronyssidae). Revista Brasileira de Entomologia, 1: 79-92.

Fuantos-Gámez B.A, Romero-Núñez C., Sheinberg-Waisburd G., Bautista-Gómez L.G, Yarto-Jaramillo E., Heredia-Cardenas R., Miranda-Contreras L. 2020. Successful treatment of *Ophionyssus natricis* with afoxolaner in two Burmese pythons (*Python molurus bivittatus*). Veterinary Dermatology, 31: 496-498. https://doi.org/10.1111/vde.12898

Garrett C.M., Harwell G. 1991. Loreal pit impaction in a black speckled palm-pitviper (*Bothriechis nigroviridis*). Journal of Zoo and Wildlife Medicine, 22: 249-251.

Gervais F.L.P. 1844. Acarides. In: Walckenaer C.A. (Ed.). Histoire Naturelle des Insectes. Aptères. III. Paris: Roret. p. 132-288.

Gobble J. 2022. Oral Fluralaner (Bravecto®) use in the control of mites in 20 Ball Pythons (*Python regius*). Journal of Herpetological Medicine and Surgery, 32: 111-112. https://doi.org/10.5818/JHMS-D-21-00003

Goldberg S.R., Bursey C.R. 1991. Integumental lesions caused by ectoparasites in a wild population of the Side-blotched lizard *Uta stansburiana*. Journal of Wildlife Diseases, 27: 68-73. https://doi.org/10.7589/0090-3558-27.1.68

Gomes-Almeida, B.K., Pepato, A.R. 2021. A new genus and new species of macronyssid mite (Mesostigmata: Gamasina: Macronyssidae) from Brazilian caves including molecular data and key for genera occurring in Brazil. Acarologia, 61: 501-526. https://doi.org/10.24349/acarologia/2021447

Hallas T.E. 1978. Fortegnelse over Danske mider (Acari). Entomologiske Meddelelser, 46: 27-45.

Hassl A.R. 2016. Ticks and mites parasitizing free-ranging reptiles in Austria - with an identification key to Central European herpetophagous Acarina. Herpetozoa, 29: 77-83.

Heath A.C.G. 1986. First occurrence of the reptile mite, *Ophionyssus natricis* (Acari: Dermanyssidae) in New Zealand. New Zealand Veterinary Journal, 34: 78. https://doi.org/10.1080/00480169.1986.35304

Huchzermeyer E.W. 2003. Crocodiles. Biology, Husbandry and Diseases. Wallingford: CAB Publishing. pp. 353. https://doi.org/10.1079/9780851996561.0000

Jacobson E.R. 1984. Immobilization, blood sampling, necropsy techniques and diseases of crocodilians: a review. Journal of Zoo Animal Medicine, 15: 38-45. https://doi.org/10.2307/20094678

Keegan, H.L. 1956. Ectoparasitic lealaptid and dermanyssid mites of Egypt, Kenya and: the Sudan, primarily based on Namru 3 Collections, 1948 - 1953. Journal of the Egyptian Public Health Association, 31: 199-272 + Plates I-XIII.

Keskin A. 2021. Occurrence of *Ophionyssus natricis* (Acari: Macronyssidae) on the captive corn snake, *Pantherophis guttatus*, (Squamata: Colubridae) in Turkey. Acarological Studies, 3: 89-95.
https://doi.org/10.47121/acarolstud.907114

Khaire A., Khaire N., Joshi P.V., Bhat, H.R. 1987. An outbreak of infestation of *Ophionyssus natricis* (Gervais, 1844) in the snake park at Poona, India. The Snake, 19: 70-71.

Leslie A.J., Lovely C.J., Pittman J.M. 2011. A preliminary disease survey in the wild Nile crocodile (*Crocodylus niloticus*) population in the Okavango Delta, Botswana. Journal of the South African Veterinary Association, 82: 155-159. https://doi.org/10.4102/jsava.v82i3.54

Ma L.-M., Bai X.-L. 2012. Discovery of the genus *Ophionyssus* Mégnin in China and description of a new species (Acari: Mesostigmata: Macronyssidae). Acta Archnologica Sinica, 21: 83-86.

Magnusson W.E. 1985. Habitat selection, parasites and injuries in Amazonian crocodilians. Amazoniana, 9: 193-204.

Manoj R.R.S., Latrofa M.S., Mendoza-Roldan J.A., Otranto D. 2021. Molecular detection of *Wolbachia* endosymbiont in reptiles and their ectoparasites. Parasitology Research, 120: 3255-3261. https://doi.org/10.1007/s00436-021-07237-1

Mariana A., Vellayan S., Halimaton I., Ho T.M. 2011. Acariasis on pet Burmese python, *Python molurus bivittatus* in Malaysia. Asian Pacific Journal of Tropical Medicine, 4: 227-228. https://doi.org/10.1016/S1995-7645(11)60075-8

Markov G.S., Bogdanov O.P. 1960. Helminths and mites – parasites of snakes in Central Asia. Uzbek biological journal, 1: 35-41.

Markov G.S., Ivanov V.P., Kryuchkov G.P., Lukianova Zh.F., Nikulin V.P., Chernobay V.F. 1964. Protozoan and parasitic mites of reptiles from the Caspian region. Scientific notes of the Volgograd State Pedagogical Institute Named after A.S. Serafimovich, 16: 106-110. [In Russian]

Maslova I.V., Dochevoy Y.E. 2016. A case of protocooperation between the pitviper *Gloydius ussuriensis* and the ground beetle *Carabus granulatus telluris*. Russian Journal of Herpetology, 23: 231-234.
https://doi.org/10.30906/1026-2296-2016-23-3-231-234
Mauri R. 1967. Acaros Mesostigmata parásitos de vertebrados de la República Argentina. Segundas Jornadas Entomoepidemiológicas Argentinas, 1: 65-73.

Maxwell C. 2022. Snake Mites: What They Are and How to Treat. Available from: https://a-z-animals.com/blog/snake-mites-what-they-are-and-how-to-treat. [13 Oct 2023]

McClain D., Dana A.N., Goldenberg G. 2009. Mite infestations. Dermatologic Therapy, 22: 327-346.
https://doi.org/10.1111/j.1529-8019.2009.01245.x

Mégnin P. 1884a. Etude d'un acarien parasite des serpents, l'Ophionyssus natricis, P. Gervais, et de son action. Compte Rendus des Séances et Mémoires de la Société de Biologie, Paris, 35: 614-619.

Mégnin P. 1884b. Étude sur l'Ophionyssus natricis P. Gervais. Bulletin de la Societe Zoologique de France, 9: 107-13 + Plate II.

Mehl R. 1979. Checklist of Norwegian ticks and mites (Acari). Fauna Norvegica, Series B, 26, 31-45.

Mendoza-Roldan J.A.R. 2019 Acarofauna of Reptiles and Amphibians of Brazil: Morphological and Molecular Studies and Pathogens Research [Doctor Thesis], University of Sao Paulo. pp. 461.

Mendoza-Roldan J.A., Colella V., Lia R.P., Nguyen V.L., Barros-Battesti D.M., Iatta R., Dantas-Torres F., Otranto D. 2019. *Borrelia burgdorferi* (sensu lato) in ectoparasites and reptiles in southern Italy. *Parasites & Vectors*, 12; 35. https://doi.org/10.1186/s13071-019-3286-1

Mendoza-Roldan J.A., Modry D., Otranto D. 2020a. Zoonotic parasites of reptiles: A crawling threat. Trends in Parasitology, 36: 677-687. https://doi.org/10.1016/j.pt.2020.04.014

Mendoza-Roldan J., Ribeiro S.R. Castilho-Onofrio V., Grazziotin F.G., Rocha B., Ferreto-Fiorillo B., Pereira J.S., Benellig G., Otranto D., Barros-Battesti D.M. 2020b. Mites and ticks of reptiles and amphibians in Brazil. Acta Tropica, 208: 105515. https://doi.org/10.1016/j.actatropica.2020.105515

Mendoza-Roldan J.A., Mendoza-Roldan M.A., Otranto D. 2021. Reptile vector-borne diseases of zoonotic concern. International Journal for Parasitology: Parasites and Wildlife, 15: 132-142.
https://doi.org/10.1016/j.ijppaw.2021.04.007

Mendoza-Roldan J.A., Napoli E., Perles L., Marino M., Spadola F., Berny P., España B., Brianti E., Beugnet F., Otranto D. 2023. Afoxolaner (NexGard®) in pet snakes for the treatment and control of *Ophionyssus natricis* (Mesostigmata: Macronyssidae). Parasites & Vectors, 16: 6.
https://doi.org/10.1186/s13071-022-05611-1

Mendyk R.W. 2015. Preliminary notes on the use of the predatory soil mite *Stratiolaelaps scimitus* (Acari: Laelapidae) as a biological control agent for acariasis in lizards. *Journal of Herpetological Medicine and Surgery*, 25: 24-27. https://doi.org/10.5818/1529-9651.25.1.24

Metaxa L. 1823. Monographia de Serpenti di Roma e Suoi Contorni. Rome: Nella Stamperia de Romanis. pp. 48. https://doi.org/10.5962/bhl.title.5101

Micherdzinski W. 1980. Eine Taxonomische Analyse der Familie Macronyssidae Oudemans, 1936. I. Subfamilie Ornithonyssinae Lange, 1958 (Acarina, Mesostigmata). Warszawa: Polska Akademia Nauk. pp 263.

Miranda R.J., Cleghorn J.E., Bermúdez S.E., Perotti, M.A. 2017. Occurrence of the mite *Ophionyssus natricis* (Acari: Macronyssidae) on captive snakes from Panama. Acarologia, 57: 365-368.
https://doi.org/10.1051/acarologia/20164161

Miron L., Ivan O. 2003. O nouă species a genului *Ophionyssus* Mégnin (1844) (Acari, Gamasida, Macronyssidae), ectoparazit pe *Vipera ursinii*. Revista Scientia Parasitologica, 4: 172-174.

Moraza M.L., Irwin N.R., Godinho R., Baird S.J.E., de Bellocq J.G. 2009. A new species of *Ophionyssus* Mégnin (Acari: Mesostigmata: Macronyssidae) parasitic on *Lacerta schreiberi* Bedriaga (Reptilia: Lacertidae) from the Iberian Peninsula, and a world key to species. *Zootaxa*, 2007: 58-68.
https://doi.org/10.11646/zootaxa.2007.1.3

Nieri-Bastos F.A., Labruna M.B., Marcili A., Durden, L.A., Mendoza-Uribe, L., Barros-Battesti D.M. 2011. Morphological and molecular analysis of *Ornithonyssus* spp. (Acari: Macronyssidae) from small terrestrial mammals in Brazil. Experimental and Applied Acarology, 55: 305-327.
https://doi.org/10.1007/s10493-011-9475-z

doi 10.1007/s10493-011-9475-z

Norval G., Halliday B., Sih A., Sharrad R.D., Gardner M.G. 2020. Occurrence of the introduced snake mite, *Ophionyssus natricis* (Gervais, 1844), in the wild in Australia. Acarologia, 60: 559-565.
https://doi.org/10.24349/acarologia/20204385

doi: 10.24349/acarologia/20204385 https://doi.org/10.24349/acarologia/20204385

Norval G., Halliday B., Sharrad R.D., Gardner M.G. 2021. Additional instances of snake mite (*Ophionyssus natricis*) parasitism on sleepy lizards (*Tiliqua rugosa*) in South Australia. Transactions of the Royal Society of South Australia, 145: 183-193. https://doi.org/10.1080/03721426.2021.1934629

Oliver J.H. 1971. Parthenogenesis in mites and ticks (Arachnida: Acari). American Zoologist, 11: 283-299. https://doi.org/10.1093/icb/11.2.283

Oudemans A.C. 1901. Notes on Acari. Third Series. Tijdschrift der Nederlandsche Dierkundige Vereeniging, 7: 50-88 + Plates I-III.

Page L.A. 1966. Diseases and infections of snakes: a review. Bulletin of the Wildlife Disease Association, 2: 111-126. https://doi.org/10.7589/0090-3558-2.4.111

Paredes-León R., García-Prieto L., Guzmán-Cornejo C., León-Regagnon V., Pérez T.M. 2008. Metazoan parasites of Mexican amphibians and reptiles. Zootaxa, 1904: 1-166. https://doi.org/10.11646/zootaxa.1904.1.1

Partyka J.K. 2019. Parasites and People: Crocodile Parasite Interactions in Human Impacted Areas of Belize [Master's Thesis]. Ás: Norwegian Journal of Life Sciences, pp. 48.

Pawełczyk O., Pająk C., Solarz K. 2016. The risk of exposure to parasitic mites and insects occurring on pets in Southern Poland. Annals of Parasitology, 62: 337-344.

Pereira A.C., Colli G.R. 2023. Landscape features affect caiman body condition in the middle Araguaia River floodplain. Animal Conservation, 26: 531-545. https://doi.org/10.1111/acv.12841
Mauri R. 1967. Acaros Mesostigmata parásitos de vertebrados de la República Argentina. Segundas Jornadas Entomoepidemiológicas Argentinas, 1: 65-73.

Maxwell C. 2022. Snake Mites: What They Are and How to Treat. Available from: https://a-z-animals.com/blog/snake-mites-what-they-are-and-how-to-treat. [13 Oct 2023]

McClain D., Dana A.N., Goldenberg G. 2009. Mite infestations. Dermatologic Therapy, 22: 327-346.
https://doi.org/10.1111/j.1529-8019.2009.01245.x

Mégnin P. 1884a. Etude d'un acarien parasite des serpents, l'Ophionyssus natricis, P. Gervais, et de son action. Compte Rendus des Séances et Mémoires de la Société de Biologie, Paris, 35: 614-619.

Mégnin P. 1884b. Étude sur l'Ophionyssus natricis P. Gervais. Bulletin de la Societe Zoologique de France, 9: 107-13 + Plate II.

Mehl R. 1979. Checklist of Norwegian ticks and mites (Acari). Fauna Norvegica, Series B, 26, 31-45.

Mendoza-Roldan J.A.R. 2019 Acarofauna of Reptiles and Amphibians of Brazil: Morphological and Molecular Studies and Pathogens Research [Doctor Thesis], University of Sao Paulo. pp. 461.

Mendoza-Roldan J.A., Colella V., Lia R.P., Nguyen V.L., Barros-Battesti D.M., Iatta R., Dantas-Torres F., Otranto D. 2019. *Borrelia burgdorferi* (sensu lato) in ectoparasites and reptiles in southern Italy. *Parasites & Vectors*, 12; 35. https://doi.org/10.1186/s13071-019-3286-1

Mendoza-Roldan J.A., Modry D., Otranto D. 2020a. Zoonotic parasites of reptiles: A crawling threat. Trends in Parasitology, 36: 677-687. https://doi.org/10.1016/j.pt.2020.04.014

Mendoza-Roldan J., Ribeiro S.R. Castilho-Onofrio V., Grazziotin F.G., Rocha B., Ferreto-Fiorillo B., Pereira J.S., Benellig G., Otranto D., Barros-Battesti D.M. 2020b. Mites and ticks of reptiles and amphibians in Brazil. Acta Tropica, 208: 105515. https://doi.org/10.1016/j.actatropica.2020.105515

Mendoza-Roldan J.A., Mendoza-Roldan M.A., Otranto D. 2021. Reptile vector-borne diseases of zoonotic concern. International Journal for Parasitology: Parasites and Wildlife, 15: 132-142.
https://doi.org/10.1016/j.ijppaw.2021.04.007

Mendoza-Roldan J.A., Napoli E., Perles L., Marino M., Spadola F., Berny P., España B., Brianti E., Beugnet F., Otranto D. 2023. Afoxolaner (NexGard®) in pet snakes for the treatment and control of *Ophionyssus natricis* (Mesostigmata: Macronyssidae). *Parasites & Vectors*, 16: 6.
https://doi.org/10.1186/s13071-022-05611-1

Mendyk R.W. 2015. Preliminary notes on the use of the predatory soil mite *Stratiolaelaps scimitus* (Acari: Laelapidae) as a biological control agent for acariasis in lizards. *Journal of Herpetological Medicine and Surgery*, 25: 24-27. https://doi.org/10.5818/1529-9651.25.1.24

Metaxa L. 1823. Monographia de Serpenti di Roma e Suoi Contorni. Rome: Nella Stamperia de Romanis. pp. 48. https://doi.org/10.5962/bhl title.5101

Micherdzinski W. 1980. Eine Taxonomische Analyse der Familie Macronyssidae Oudemans, 1936. I. Subfamilie Ornithonyssinae Lange, 1958 (Acarina, Mesostigmata). Warszawa: Polska Akademia Nauk. pp 263.

Miranda R.J., Cleghorn J.E., Bermúdez S.E., Perotti, M.A. 2017. Occurrence of the mite *Ophionyssus natricis* (Acari: Macronyssidae) on captive snakes from Panama. Acarologia, 57: 365-368.
https://doi.org/10.1051/acarologia/20164161

Miron L., Ivan O. 2003. O nouă species a genului *Ophionyssus* Mégnin (1844) (Acari, Gamasida, Macronyssidae), ectoparazit pe *Vipera ursinii*. Revista Scientia Parasitologica, 4: 172-174.

Moraza M.L., Irwin N.R., Godinho R., Baird S.J.E., de Bellocq J.G. 2009. A new species of *Ophionyssus* Mégnin (Acari: Mesostigmata: Macronyssidae) parasitic on *Lacerta schreiberi* Bedriaga (Reptilia: Lacertidae) from the Iberian Peninsula, and a world key to species. *Zootaxa*, 2007: 58-68.
https://doi.org/10.11646/zootaxa.2007.1.3

Nieri-Bastos F.A., Labruna M.B., Marcili A., Durden, L.A., Mendoza-Uribe, L., Barros-Battesti D.M. 2011. Morphological and molecular analysis of *Ornithonyssus* spp. (Acari: Macronyssidae) from small terrestrial mammals in Brazil. Experimental and Applied Acarology, 55: 305-327.
https://doi.org/10.1007/s10493-011-9475-z

doi 10.1007/s10493-011-9475-z

Norval G., Halliday B., Sih A., Sharrad R.D., Gardner M.G. 2020. Occurrence of the introduced snake mite, *Ophionyssus natricis* (Gervais, 1844), in the wild in Australia. Acarologia, 60: 559-565.
https://doi.org/10.24349/acarologia/20204385

doi: 10.24349/acarologia/20204385 https://doi.org/10.24349/acarologia/20204385

Norval G., Halliday B., Sharrad R.D., Gardner M.G. 2021. Additional instances of snake mite (*Ophionyssus natricis*) parasitism on sleepy lizards (*Tiliqua rugosa*) in South Australia. Transactions of the Royal Society of South Australia, 145: 183-193. https://doi.org/10.1080/03721426.2021.1934629

Oliver J.H. 1971. Parthenogenesis in mites and ticks (Arachnida: Acari). American Zoologist, 11: 283-299. https://doi.org/10.1093/icb/11.2.283

Oudemans A.C. 1901. Notes on Acari. Third Series. Tijdschrift der Nederlandsche Dierkundige Vereeniging, 7: 50-88 + Plates I-III.

Page L.A. 1966. Diseases and infections of snakes: a review. Bulletin of the Wildlife Disease Association, 2: 111-126. https://doi.org/10.7589/0090-3558-2.4.111

Paredes-León R., García-Prieto L., Guzmán-Cornejo C., León-Regagnon V., Pérez T.M. 2008. Metazoan parasites of Mexican amphibians and reptiles. Zootaxa, 1904: 1-166. https://doi.org/10.11646/zootaxa.1904.1.1

Partyka J.K. 2019. Parasites and People: Crocodile Parasite Interactions in Human Impacted Areas of Belize [Master's Thesis]. Ás: Norwegian Journal of Life Sciences, pp. 48.

Pawełczyk O., Pająk C., Solarz K. 2016. The risk of exposure to parasitic mites and insects occurring on pets in Southern Poland. Annals of Parasitology, 62: 337-344.

Pereira A.C., Colli G.R. 2023. Landscape features affect caiman body condition in the middle Araguaia River floodplain. Animal Conservation, 26: 531-545. https://doi.org/10.1111/acv.12841
Phisalix M. 1914. Sur une Hémogrégarine nouvelle, parasite de *Boodon fuliginosus* Boïe, et ses forms de multiplication endogène. Bulletin de la Société de Pathologie Exotique, 7: 358-360.

Radford C.D. 1950. The mites (Acarina) parasitic on mammals, birds and reptiles. Parasitology, 40: 366-394. https://doi.org/10.1017/S0031182000018230

Radovsky F.J. 1971. *Ophionyssus natricis* (Gervais). Proceedings of the Hawaiian Entomological Society, 21: 13-14.

Radovsky F.J. 2010. Revision of Genera of the Parasitic Mite Family Macronyssidae (Mesostigmata: Dermanyssoidea) of the World. West Bloomfield: Indira Publishing House. pp. 170.

Rataj A.V., Lindtner-Knific R., Vlahović K., Mavri U., Dovč A. 2011. Parasites in pet reptiles. Acta Veterinaria Scandinavica, 53: 33. https://doi.org/10.1186/1751-0147-53-33

Reeves W.K., Dowling A.P.G., Dasch G.A. 2006. Rickettsial agents from parasitic Dermanysssoidea (Acari: Mesostigmata). Experimental and Applied Acarology, 38: 181-188. https://doi.org/10.1007/s10493-006-0007-1

doi 10.1007/s10493-006-0007-1

Reinert W., Brandstätter F. 1993. Zur Biologie der Milbe *Ophionyssus natricis*. Salamandra, 29: 248-257.

Reptiles Cove. 2022. Acariasis Caused by Snake Mites: Everything You Need to Know. Available from https://reptilescove.com/care/snakes/acariasis. [13 Oct 2023].

Rimbaud E., Pineda N., Luna L., Zepeda N., Rivera G. 2006. Primer reporte de *Ophionyssus natricis* (Arthropoda, Acarina, Macronyssidae, Gervais 1953) parasitando *Boa constrictor constrictor* en Nicaragua. Boletín de Parasitología Escuela de Medicina Veterinaria Universidad Nacional Costa Rica, 7 (2): 1.

Rodriguez M.L., Lazcano D. 1992. Primer reporte de ácaro *Ophionyssus natricis* (Acarina: Macronyssidae) para México. Southwestern Naturalist, 37: 426. https://doi.org/10.2307/3671798

Rotter J. 1963. Die Warane. Wittenberg: Ziemsen Verlag. pp. 76.

Sabu L., Devada K., Subramanian H., Cheeran J.V. 2002. *Diploscapter coronata* (Cobb, 1893) infestation in captive snakes. Zoos' Print Journal, 17: 954-956. https://doi.org/10.11609/JoTT.ZPJ.17.12.954-6

Salazar M.M. 2014. Determination of Control Parasitario en iguana verde (Iguana iguana) Mantenida Cautivero [Master's thesis]. Mexico: Universidad del Mar. pp. 104.

Santana-Hernández K.M., Orós J., Priestnall S.L., Monzón-Argüello C., Rodríguez-Ponce E. 2021. Parasitological findings in the invasive California kingsnake (*Lampropeltis californiae*) in Gran Canaria, Spain. Parasitology, 148: 1345-1352. https://doi.org/10.1017/S0031182021000871

Schilliger L.H., Morel D., Bonwitt J.H., Marquis O. 2013. *Cheyletus eruditus* (Taurrus®): An effective candidate for the biological control of the snake mite (*Ophionyssus natricis*). Journal of Zoo and Wildlife Medicine, 44: 654-659. https://doi.org/10.1638/2012-0239R.1

Schroeder C.R. 1934. The snake mite (*Ophionyssus serpentium* Hirst). Journal of Economic Entomology, 28: 1004-1014. https://doi.org/10.1093/jee/27.5.1004

Schultz H. 1975. Human infestation by *Ophionyssus natricis* snake mite. British Journal of Dermatology, 93: 695-697. https://doi.org/10.1111/j.1365-2133.1975.tb05120.x

Schumacher J., Jacobson E.R. Homer B.L., Gaskin J.M. 1994. Inclusion body disease in boid snakes. Journal of Zoo and Wildlife Medicine, 25: 511-524.

Schweizer J. 1961. Die Landmilben der Schweiz (Mittelland, Jura und Alpen). Parasitiformes Reuter. Denkschriften der Schweizerischen Naturfoorschenden Gesellschaft, 84: 1-207.

Shulov A. 1957. Additions to the fauna of Acarina of Israel (excluding ticks and gall mites). Bulletin of the Research Council of Israel, 6B: 233-238.

Simonov E., Zinchenko V. 2010. Intensive infestation of Siberian pit-viper, *Gloydius halys halys* by the common snake mite, *Ophionyssus natricis*. North-Western Journal of Zoology, 6: 134-137.

Šlapeta J., Modrý D., Johnson R. (2018) Reptile parasitology in health and disease. In: Doneley B., Monks D., Johnson R., Carmel B. (eds) Reptile Medicine and Surgery in Clinical Practice. John Wiley & Sons Ltd., p. 425-440. https://doi.org/10.1002/9781118977705.ch31

Su H.-Y., Gao X.-P., Bai X.-L. 2010. Investigation on parasitic gamasid on *Phrynocephalus przewalskii* (Strauch, 1876) in Ningxia. Endemic Diseases Bulletin, 25: 4-5.

Sumaira, Bukhari M.S., Iram M., Javid A., Hussain A., Ali W., Rehman K.U., Andleeb S., Baboo I., Khalid N. 2020. Seasonal variation in ecdysis process and parasitic prevalence in Indian Rock Python (*Python molurus*). Pakistan Journal of Zoology, 52: 2341-2346. https://doi.org/10.17582/journal.pjz/20190417060401

Tarshis I.B. 1960. Control of the snake mite (*Ophionyssus natricis*), other mites, and certain insects with the sorptive dust, SG 67. Journal of Economic Entomology, 53: 903-908. https://doi.org/10.1093/jee/53.5.903

Tellez M. 2013. A checklist of host-parasite interaction of the order Crocodylia. University of California Publications in Entomology, 136: 1-376. https://doi.org/10.1525/california/9780520098893.003.0001

Thaler R., Ortega Z., Ferreira V.L. (2022) Extrinsic traits consistently drive microhabitat decisions of an arboreal snake, independently of sex and personality. Behavioral Processes, 199 (104649): 1-10. https://doi.org/10.1016/j.beproc.2022.104649

Thomasson V., Blouin-Demers G. 2015. Using habitat suitability models considering biotic interactions to inform critical habitat delineation: An example with the eastern hog-nosed snake (*Heterodon platirhinos*) in Ontario, Canada. Canadian Wildlife Biology and Management, 4: 1-17.

Till W.M. 1957. Mesostigmatic mites living as parasites of reptiles in the Ethiopian region (Acarina: Laelaptidae). Journal of the Entomological Society of Southern Africa, 20: 120-143.

Uetz P., Freed P., Aguilar R., Reyes F., Hošek J. 2023. *The Reptile Database*. Available from http://www.reptile-database.org. [13 Oct 2023].

Vasiliev D.B., Balakina O.S. (undated) Hematological studies in infectious and parasitic diseases of a rare species of pythons, *Morelia boeleni*. Available from http://www.dompitomci.ru/doc/conf/2097.html [19 Oct 2023]
Phisalix M. 1914. Sur une Hémogrégarine nouvelle, parasite de *Boodon fuliginosus* Boïe, et ses forms de multiplication endogène. Bulletin de la Société de Pathologie Exotique, 7: 358-360.

Radford C.D. 1950. The mites (Acarina) parasitic on mammals, birds and reptiles. Parasitology, 40: 366-394. https://doi.org/10.1017/S0031182000018230

Radovsky F.J. 1971. *Ophionyssus natricis* (Gervais). Proceedings of the Hawaiian Entomological Society, 21: 13-14.

Radovsky F.J. 2010. Revision of Genera of the Parasitic Mite Family Macronyssidae (Mesostigmata: Dermanyssoidea) of the World. West Bloomfield: Indira Publishing House. pp. 170.

Rataj A.V., Lindtner-Knific R., Vlahović K., Mavri U., Dovč A. 2011. Parasites in pet reptiles. Acta Veterinaria Scandinavica, 53: 33. https://doi.org/10.1186/1751-0147-53-33

Reeves W.K., Dowling A.P.G., Dasch G.A. 2006. Rickettsial agents from parasitic Dermanysssoidea (Acari: Mesostigmata). Experimental and Applied Acarology, 38: 181-188. https://doi.org/10.1007/s10493-006-0007-1

doi 10.1007/s10493-006-0007-1

Reinert W., Brandstätter F. 1993. Zur Biologie der Milbe *Ophionyssus natricis*. Salamandra, 29: 248-257.

Reptiles Cove. 2022. Acariasis Caused by Snake Mites: Everything You Need to Know. Available from https://reptilescove.com/care/snakes/acariasis. [13 Oct 2023].

Rimbaud E., Pineda N., Luna L., Zepeda N., Rivera G. 2006. Primer reporte de *Ophionyssus natricis* (Arthropoda, Acarina, Macronyssidae, Gervais 1953) parasitando *Boa constrictor constrictor* en Nicaragua. Boletín de Parasitología Escuela de Medicina Veterinaria Universidad Nacional Costa Rica, 7 (2): 1.

Rodriguez M.L., Lazcano D. 1992. Primer reporte de ácaro *Ophionyssus natricis* (Acarina: Macronyssidae) para México. Southwestern Naturalist, 37: 426. https://doi.org/10.2307/3671798

Rotter J. 1963. Die Warane. Wittenberg: Ziemsen Verlag. pp. 76.

Sabu L., Devada K., Subramanian H., Cheeran J.V. 2002. *Diploscapter coronata* (Cobb, 1893) infestation in captive snakes. Zoos' Print Journal, 17: 954-956. https://doi.org/10.11609/JoTT.ZPJ.17.12.954-6

Salazar M.M. 2014. Determination of Control Parasitario en iguana verde (Iguana iguana) Mantenida Cautivero [Master's thesis]. Mexico: Universidad del Mar. pp. 104.

Santana-Hernández K.M., Orós J., Priestnall S.L., Monzón-Argüello C., Rodríguez-Ponce E. 2021. Parasitological findings in the invasive California kingsnake (*Lampropeltis californiae*) in Gran Canaria, Spain. Parasitology, 148: 1345-1352. https://doi.org/10.1017/S0031182021000871

Schilliger L.H., Morel D., Bonwitt J.H., Marquis O. 2013. *Cheyletus eruditus* (Taurrus®): An effective candidate for the biological control of the snake mite (*Ophionyssus natricis*). Journal of Zoo and Wildlife Medicine, 44: 654-659. https://doi.org/10.1638/2012-0239R.1

Schroeder C.R. 1934. The snake mite (*Ophionyssus serpentium* Hirst). Journal of Economic Entomology, 28: 1004-1014. https://doi.org/10.1093/jee/27.5.1004

Schultz H. 1975. Human infestation by *Ophionyssus natricis* snake mite. British Journal of Dermatology, 93: 695-697. https://doi.org/10.1111/j.1365-2133.1975.tb05120.x

Schumacher J., Jacobson E.R. Homer B.L., Gaskin J.M. 1994. Inclusion body disease in boid snakes. Journal of Zoo and Wildlife Medicine, 25: 511-524.

Schweizer J. 1961. Die Landmilben der Schweiz (Mittelland, Jura und Alpen). Parasitiformes Reuter. Denkschriften der Schweizerischen Naturfoorschenden Gesellschaft, 84: 1-207.

Shulov A. 1957. Additions to the fauna of Acarina of Israel (excluding ticks and gall mites). Bulletin of the Research Council of Israel, 6B: 233-238.

Simonov E., Zinchenko V. 2010. Intensive infestation of Siberian pit-viper, *Gloydius halys halys* by the common snake mite, *Ophionyssus natricis*. North-Western Journal of Zoology, 6: 134-137.

Šlapeta J., Modrý D., Johnson R. (2018) Reptile parasitology in health and disease. In: Doneley B., Monks D., Johnson R., Carmel B. (eds) Reptile Medicine and Surgery in Clinical Practice. John Wiley & Sons Ltd., p. 425-440. https://doi.org/10.1002/9781118977705.ch31

Su H.-Y., Gao X.-P., Bai X.-L. 2010. Investigation on parasitic gamasid on *Phrynocephalus przewalskii* (Strauch, 1876) in Ningxia. Endemic Diseases Bulletin, 25: 4-5.

Sumaira, Bukhari M.S., Iram M., Javid A., Hussain A., Ali W., Rehman K.U., Andleeb S., Baboo I., Khalid N. 2020. Seasonal variation in ecdysis process and parasitic prevalence in Indian Rock Python (*Python molurus*). Pakistan Journal of Zoology, 52: 2341-2346. https://doi.org/10.17582/journal.pjz/20190417060401

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https://doi.org/10.2300/acari.2.75
Vitzthum H. 1943. Dr. H. G. Bronns Klassen und Ordnungen des Tierreichs. 5 Band: Arthropoda. IV. Abteilung: Arachnoidea. 5. Buch. Acarina. Leipzig: Akademische Verlagsgesellschaft, Becker & Erler Kom. pp. 1011.

Whan S.-T. 1966. Studies on the fauna of Korean Mites (Gamasoidea) (I). Saengmulhak *Biology*, 5 (3): 27-33 (not seen, cited by Beron, 2014).

Wiechert J.M. 2007. Infection of Hermann's tortoises, *Testudo hermanni boettgeri*, with the Common Snake Mite, *Ophionyssus natricis*. Journal of Herpetological Medicine and Surgery, 17: 53-54.
https://doi.org/10.5818/1529-9651.17.2.53

Womersley H. 1956. On some new Acarina-Mesostigmata from Australia, New Zealand and New Guinea. Journal of the Linnean Society of London, Zoology, 42: 505-599. https://doi.org/10.1111/j.1096-3642.1956.tb02218.x

Wozniak E.J., DeNardo D.F. 2000. The biology, clinical significance and control of the common snake mite, *Ophionyssus natricis*, in captive reptiles. Journal of Herpetological Medicine and Surgery, 10: 4-10. https://doi.org/10.5818/1529-9651-10.3.4

Yunker C.E. 1956. Studies on the snake mite, *Ophionyssus natricis*, in nature. Science, 124: 979-980.
https://doi.org/10.1126/science.124.3229.979

Zemskaya A.A. 1951. Biology and development of the mites from the family Dermanyssidae parasitic on reptiles in the connection with the problem of their mode of parasitism. Bulletin de la Societe des Naturalistes Moscow Biologie, 56: 42-57. [In Russian].

Zhang M.-Y., Uchikawa K. 1993. Occurrence of *Ophionyssus natricis* on zoo snakes in Japan (Mesostigmata, Macronyssidae). Journal of the Acarological Society of Japan, 2: 75-78.
https://doi.org/10.2300/acari.2.75