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Article

The Biodiversity of Demodecid Mites (Acariformes: Prostigmata), Specific Parasites of Mammals with a Global Checklist and a New Finding for Demodex sciurinus

by
Joanna N. Izdebska
* and
Leszek Rolbiecki
Department of Invertebrate Zoology and Parasitology, Faculty of Biology, University of Gdańsk, Wita Stwosza 59, 80-308 Gdańsk, Poland
*
Author to whom correspondence should be addressed.
Diversity 2020, 12(7), 261; https://doi.org/10.3390/d12070261
Submission received: 31 May 2020 / Revised: 23 June 2020 / Accepted: 24 June 2020 / Published: 30 June 2020
(This article belongs to the Special Issue Biodiversity of Mites)
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Abstract

:
Demodecidae are the most specialized parasitic mites of mammals; they typically inhabit the skin, but they have been found in other tissues and organs. They can cause demodecosis (a disease which is hazardous and difficult to cure) in humans, domestic animals and livestock. They are parasites with high host and topical specificity. They have been found for most orders of mammals, and they are common in the populations of numerous host species. Therefore, they not only constitute an important subject of veterinary and medical study, but also comprise an excellent model for faunistic and parasitological analyses concerning different aspects of functioning and evolution of the host–parasite relationship. The current level or knowledge of demodecid mites is irregular and fragmentary, and numerous questions require elaboration and ordering, from the taxonomic diversity to geographic distribution and relations with hosts. Such data may be of use i.a. for the development of more efficient and reliable diagnostic methods, as well as understanding the etiology and pathogenesis mechanisms of demodecosis, currently a contentious issue. The present paper lists all formally-described valid species of demodecid mites, together with other functioning specific names, verified and with comments on their status. This is significant for correct species identification and demodecosis diagnostics. The list has been drawn up on the basis of data acquired in the period 1842−2020. It contains 122 valid species of parasite, including their hosts and geographic distribution, data on parasitism, as well as only the second record of Demodex sciurinus in Eurasian red squirrel Sciurus vulgaris in over 100 years since its initial discovery.

Graphical Abstract

1. Introduction

The members of the Demodecidae are specialized, typically monoxenic mammal parasites, and are likely abundant within host populations. They are stationary parasites, with their whole life cycle spent on the host; however, depending on the species, its topography, seasonal dynamics and transmission mechanism, they may exhibit a variable level of infestation prevalence, which may reach up to 100% [1,2]. Typically, the presence of demodecid mites does not produce disease symptoms, even at high infestation intensity and high density on the skin [3,4,5]. However, under favorable host circumstances, the high density of these mites may be linked to the development of demodecosis (formerly demodicosis, demodicidosis). Demodecosis often has a complicated course, depending on various factors including the species constituting the etiologic factor. Its symptoms typically include the presence of various skin lesions with different topography, hair loss, eyelid margin inflammation and conjunctivitis, and changes within gum mucous membranes; however, it also can occur in generalized form. Complications may also occur in the form of secondary bacterial or fungal infections. It is suspected that the possibility of developing the parasitosis may be increased by reduced host immunity, poor condition or incorrect diet, such as deficiencies in vitamins and micronutrients [6,7]. Despite the high infestation prevalence in wild mammal populations, demodecosis is observed relatively rarely. However, it constitutes a serious problem for humans, domestic animals and livestock. Demodecosis canina, a domestic dog disease, typically caused by D. canis is characterized by a particularly hazardous course, with a chronic or even fatal result [2,8]. It has a highly variable course and symptoms, which on the one hand may be linked to the vast diversity of its hosts, with different breeds demonstrating variable susceptibility, as well as etiological factors, as three new species of these mites have been identified in dogs relatively recently (Table 1). Another burdensome and resilient parasitosis is demodecosis hominis, caused in humans by the synhospital human demodecid mite D. brevis, associated with skin sebaceous glands, and D. folliculorum, found in hair follicles. It manifests there typically in the form of skin lesions (e.g., pityriasis folliculorum, rosacea-like demodecidosis, pustular folliculitis, papulo-pustular scalp eruptions, perioral dermatitis and hyperpigmented patches of the face) in the head area (skin and facial hair), hair loss and eyelid margin inflammation, as well as conjunctivitis. The nature of the pathogenesis depends on various factors, such as the etiological agent (i.e., demodecid species), and the symptoms are reminiscent of other dermatoses; therefore, diagnostics, including demodecid species identification, is important for efficient treatment [6]. A number of demodecosis variants, dictated by the host inhabited by the demodecid mite, have been described for cats and cattle, or laboratory animals such as mice and hamsters (Table 1).
Due to their morphological modifications, including miniaturization, or various adaptations to parasitism within the different microhabitats, offered by the skin, tissues and organs of the host, the Demodecidae can claim to be the most specialized parasitic mites of mammals. Moreover, they demonstrate high host and topical specificity, representing the majority of orders of mammals [9]. Their long evolutionary relationship with hosts makes them the perfect model for faunistic or parasitological analyses concerning different aspects of the functioning of a host–parasite relationship and the co-evolution of these mites and their hosts. Although the family includes species of high medical and veterinary importance, their level of recognition is insufficient, and the available information regarding them is fragmentary and dispersed. A series of issues need to be studied and ordered, from taxonomic diversity, to geographic distribution and relations with hosts. In particular, data on their distribution, including their occurrence in different areas of the host range, remains incomplete. The majority of records describe cases of these parasites found in relation to parasitosis (demodecosis) symptoms, and these are relatively rare, being restricted mostly to domestic mammals and humans, and are only sporadically observed in wild animals [2,4,10]. Detecting an asymptomatic infestation is complicated by the miniature size of the mite, with the smallest species reaching only 70–80 µm in length, and its secretive life history; in addition, the parasites inhabit a range of skin structures, including normal and sensory hair follicles and different types of glands, as well as a variety of organ tissue types, such as the tongue, gums, anterior segments of the digestive tract or auditory canals [11]. Therefore, certain species, despite inhabiting widely distributed and common hosts, are known only from singular case studies or in sparse records [4,12].
Furthermore, the literature, particularly parasitological and veterinarian studies, encompasses a series of unverified data, including species with an actual nomen nudum status, invalid-unauthorized synonyms or information based on doubtful identification [2,10]: The numbers of species classified into this family varies from several dozen to over one hundred depending on the source [10,13,14]. Originally, one comprehensive study existed, covering the 16 then-known species [15], including full data on their distribution and documented records. Only one global checklist, covering 100 species and their hosts, has been published in modern times [13], this being a part of a more extensive study of the Eleutherengona as obligatory mammal parasites. However, it does not include data on records or any validation of the functioning of the names in the literature, and Demodecidae checklists for the selected host group, i.e., rodents and soricomorphs, that have been published [16,17,18].
The present paper serves as a comprehensive study of all known species, and includes a verification of the unauthorized names, including the nomen nudum. Furthermore, it also includes a new record: The second finding of Demodex sciurinus globally, confirming the existence of this species close to one hundred years from its original discovery. Another significant objective of the revision is to organize currently-available Demodecidae records, not only for faunistic purposes, but also for parasitological, veterinary and medical research. A key value of such a summary of the current state of research is that it also highlights the absence of information from numerous countries where demodecid mites, and its relationship with demodecosis, are a significant area of study: In some of these areas, no information on the distribution of demodecid mite species has been published. Our global data also constitute a significant starting point for future, more comprehensive regional analyses, as well as the development of diversity models in the context of host–parasite relationships. These more specific findings would be of great value in the development of more efficient and reliable diagnostic methods, and in improving our understanding of the etiological mechanisms and pathogenesis of demodecosis, which is currently a contentious issue.

2. Materials and Methods

2.1. Detection of Demodecidae in Sciurus vulgaris

One squirrel, Sciurus vulgaris, originating from northern Poland (Gdynia 54°30′ N 18°32′ E), collected in 2017, was examined for demodecid mites.
Demodecid mites were isolated using skin digestion methods [19]. Skin fragments of 1 cm2 were collected from several body regions, including the head (around eyes, ear pinnae, nose, lips, chin, cheeks and vertex), neck, abdomen, back, limbs, tail and genital-anal area. Skin samples were preserved in 70% ethanol and digested in 10% potassium hydroxine solution. The obtained samples were decanted and analyzed using phase-contrast microscopy; an examination of 1 cm2 of skin was equal to the analysis of approximately 100 wet preparations. The mites were mounted in polyvinyl-lactophenol solution and photographed. The following measurements (µm) were taken as follows: Total body length equals length of gnathosoma, podosoma and opisthosoma; gnathosomal width equals width at base; and podosomal and opisthosomal width equals maximum width. The specimens were deposited in scientific collections within the framework of the Collection of Extant Invertebrates in Department of Invertebrate Zoology and Parasitology, University of Gdańsk, Gdańsk, Poland (UGDIZP).

2.2. Literature Review—The Checklist Structure, Biogeographic and Parasitological Data Analysis

The checklist has been drawn up based on manuscripts published during the period 1842–2020 (278 items). It also contains a new record, marked in the Table 1 as the present study. Demodecidae species have been listed in systematic order, and in alphabetical order within the genera. The list includes all formally described species and other functioning specific names; all of which are verified and provided with comments on their status. Information on dates of host species, as well as the occurrence have been also included. Wherein, for cosmopolitan demodecid mite species, selected records from various range regions were given. Host records related to unidentified Demodex spp. have not been included.
The scientific names, common names, and systematics of the hosts follow Wilson and Reeder [20] and the Taxonomic Information System [21].

3. Results

3.1. A New Record of Demodex sciurinus

The examined squirrel specimens were found to have D. sciurinus (Table 2, Figure 1). A total number of 13 females and 8 males were identified, as well as several specimens at nymphal stages; male and immature stages were demonstrated for the first time. All mites were found in the skin of the penis. The presence of demodecid mites was not associated with demodecosis symptoms.

3.2. Biodiversity and Geographic Distribution of Demodecidae Mites

A total of 122 demodecid mite species with verified systematic status are presently known, of which one represents Apodemodex, 106 Demodex, one Glossicodex, seven Ophthalmodex, one Pterodex, one Rhinodex, one Soricidex and four Stomatodex (Table 1). Representatives of the Demodecidae have been recorded on all continents outside of the polar regions (Figure 2), and their presence is typically dictated by the presence of a typical host; however, no studies of this group have been conducted in numerous areas of its range, even for common mammal species. Many species of hosts have wide distribution ranges and are also considered cosmopolitan.

3.3. Demodecidae Parasitism and Relationships with Hosts

The greatest diversity of Demodecidae has, thus far, been described for bats (five genera), followed by rodents (three genera) and soricomorphs (two genera) as hosts; only Demodex representatives have been recorded in the remaining mammal orders. In turn, the species diversity in the individual host groups typically corresponds to the species diversity of the host (Figure 3 and Figure 4). Hence the highest number of Demodecidae species have been described from the most abundant groups of mammals, e.g., 43 from rodents, 27 from bats, 17 from ungulates and 15 from carnivorans. In contrast, the Demodecidae from primates (six species) or marsupials (three species) have been especially poorly studied, which may be associated with limited access to the material. More than one species of Demodecidae has been recorded in 27 species of mammals, and the greatest number of synhospital species have been described among rodents and bats (Table 3).

4. Discussion

Although the Demodecidae are associated with almost all of the modern mammalian orders (Table 1), their distribution and occurrence in host populations have been poorly and unevenly studied. This paucity of information has been attributed to the difficulty in detecting asymptomatic infestation, the low number of faunistic studies and issues associated with correct species identification. This is confirmed by the high number of records for demodecid mites without species identification, listed as Demodex sp. e.g., [22,23,24,25,26,27,28,29], including valuable new records of hosts.

4.1. Demodecidae Biodiversity Analysis in the Light of Taxonomic Identification Problems

The number of demodecid mite species described thus far (122) does not seem to be very high when considering that over 5000 mammal species could act as potential hosts. This primarily stems from technical issues linked to their detection (e.g., miniature size, secretive life history with rare manifesting of their presence in the form of demodecosis), and the strict species description criteria used for the group: Representative series of specimens of both sexes are used, often with juvenile stages [9,11].
Demodecidae species associated with human, domestic and livestock mammals are an important issue in the comprehensive elaboration of this group. Although demodecid mites have been listed in handbooks and other overview papers, in lists with parasites with data on their pathogenic importance for hosts in different countries and continents, comparatively few published records exist. Therefore, verification of whether these purported Demodecidae indeed occur in these sources, or can be potentially detected in them, has proven difficult. A series of demodecosis descriptions, case studies or clinical studies exist indicating occurrence of individual demodecid mites in a given area; however, these descriptions typically lack the information needed to determine the geographic locality of the parasite or its frequency of occurrence, or whether earlier records exist. Often the identification of the demodecid mite is limited only to the genus e.g., [30,31,32,33,34,35].
In addition, a number of species with unverified status or nomen nudum exist in the parasitological or veterinary literature. As the Demodecidae are monoxenic parasites, hosts are often assigned species solely on the basis of an alleged host specificity. Recording the presence of a Demodex species in a new host species suggests a high probability that this species is new to Science; however, this requires an appropriate taxonomic analysis to be conducted and a description in accordance with the International Commission on Zoological Nomenclature (ICZN) requirements to be published [2,10]. Unfortunately, such newly-discovered taxa have been assigned unsupported names with suitable descriptions; these have also been copied in other publications, thus becoming established in the specialist literature. For instance, the lists of the parasitofauna of the gerbil, a commonly-used laboratory animal, frequently contain a reference to Demodex merioni [36,37]: An alleged species not supported with a description and not assigned to any concrete host species, since the term gerbil is used to refer to a multitude of taxa. A similar situation concerned D. cornei from the domestic dog. The name had functioned for many years to describe an alleged species referred to as “short form” from the dog epidermis. Although morphological and morphometric research [8,38] has confirmed the existence of such species, the lack of a formal, unambiguous description meant that it was impossible to verify records from different parts of the world, as these were mostly based on the criterion of length, possibly confirmed with topical distinctiveness. Although, eventually, the specific status has been explained and confirmed with an appropriate species description [2], this does not provide any possibility to verify earlier records not supported with morphological characteristics. The groundlessness of using size as a criterion to identify species within the same host has been further confirmed by the discovery of another “short” canine demodecid species, D. cyonis. Furthermore, literature data including the nomina nuda D. araneae and D. bonaparti has been published without an appropriate description [39], despite being correctly distinguished by the author: A specialist on Demodecidae research. Likewise, the description of D. myotidis, D. sciurei, D. sylvilagi or D. transitionalis were included in an unpublished dissertation, a procedure that does not meet the ICZN criterion on the publication of species descriptions (Table 1).
At present, numerous directions in Demodecidae research have employed molecular methods; however, their outcomes are not reliable given the lack of correlation between morphological taxonomy and molecular divergence. An example here would be D. “felis”, which was recorded for the domestic cat in a study solely based on molecular analyses [40]. The authors, who assigned it a temporary name, giving the impression of a species name, stipulate that it is only a working name proposal, intended for the purpose of distinguishing the alleged species, which, according to those authors, differs from other feline demodecid mites e.g., [41,42,43,44]. Such a study where cladistics is based solely on molecular data without confirming that the inferred genetic distance is a reliable evidence at the infraspecific level, cannot provide a sufficient basis to assign a species name for an identified demodicid mite; this is especially true as the results, in this case, have not been supported with any other evidence, including morphology, and it cannot be said which taxa they concern. However, despite this irregularity, the species already functions in the veterinary literature at the specific name level.
Another issue in the study of Demodecidae distribution concerns the existence of uncertainties with regards to correct identification. Individual species can differ with regard to sets of characters and small morphological elements which may be only several micrometers in size, sometimes less than 1 µm; such minute variation requires the use of suitable preparation methods, phase contrast techniques and immersion microscopy, as well as experience in such taxonomic analyses. At the same time, some studies use alleged host specificity or, sometimes, size as the basis for identification; however, the study methods described in the works do not leave any doubt that a correct identification had not been possible. As a host may be associated with different specific Demodecidae species with similar sizes and proportions, any application of the host specificity criterion in species identification is not only insufficient, but also groundless. An accidental transfer onto atypical hosts cannot be excluded, which may happen under favorable conditions, even in the case of highly-specialized parasites.

4.2. State and Perspectives for the Study on Geographic Distribution

In view of presented data, the highest number of species (confirmed records) have been recorded in Poland (51 species), the USA (23), Czech Republic (18) and Great Britain (18). Naturally, this does not stem from any special preferences of the Demodecidae for the hosts occurring in those countries, but it is consequence of sampling bias. A clear contrast can be seen between Demodecidae records obtained from wild, domestic and livestock mammals. Detections in wild animals are rare, because demodecid mites rarely manifest their presence in the form of demodecosis.
It is also possible that the occurrence of the known Demodecidae species is considerably wider than that indicated by the published data, and likely coincides with the ranges of their hosts. This has been confirmed by the latest records of D. chiropteralis, D. melesinus and, the present record of, D. sciurinus: Species formerly known only from individual records from England, and a single observation recorded a hundred years previously from distant Poland. The currently identified individuals of D. sciurinus exhibited traits complying with the description and figures published by Hirst [45], despite the description deviating from the modern standards assumed for the Demodecidae taxonomy. Therefore, a redescription will definitely be necessary in the future; this would include an initial description of the juvenile stages, which will be possible after collecting a wider range of material from a greater number of hosts. Current intensive research conducted within the area of Poland has further confirmed the presence of almost all species formerly described from the Czech Republic, the Netherlands or the USA, provided that typical hosts are to be found. It should be added that many species exhibit very high infestation prevalence in the host populations, reaching up to 100% e.g., [1,4,12,18,19].
It is currently important to organize the diverse body of data concerning the occurrence of species of medical and veterinary significance, i.e., to verify and correlate data on the occurrence of demodecosis in various host species with information on the occurrence of the agent species, as confirmed by taxonomic identification.

4.3. Host-Parasite Relationships

The diversity of Demodecidae in individual host groups is typically convergent with the species richness of the host group (Figure 2 and Figure 3). Moreover, the study of this group of species is also related to the incidence rate of demodecosis and the practical importance of hosts, thus the number of species described from carnivores is relatively higher (with regards to the biodiversity of this group) than for other mammals. Among the 17 demodecid mite species described for carnivorous mammals, six originate from domestic dogs and cats. In addition, the availability of material for study, and the technical issues related to detection, are other significant factors. The detection of asymptomatic infestation is labor-intensive, with an efficiency that is inversely proportionate to the host size. It is therefore unsurprising that the highest number of species have been described from small mammals. In contrast, the Demodecidae from primates (six species), or marsupials (three species) have been especially poorly studied, which may be associated with the limited access to the material, with only individual specimens of selected species obtained from zoological gardens being tested, and the fact that certain areas of the world, such as Asia and Australia, are absent from the body of data (Table 1 and Table 2). Therefore, it is possible that the number of existing species is considerably higher than presented herein.
The Demodecidae exhibit high specificity towards hosts (monoxeny). Only five species are considered oligoxenic, i.e., recorded from more than one host; however, all host species were closely related. Of this group, D. apodemi and S. corneti have been described according to insufficient criteria, based on the current state of knowledge, and hence require redescription. In turn, D. sabani and D. kutzeri also require taxonomic revision, as they may constitute aggregate species, with their taxa being difficult to distinguish according to morphological criteria; such revision should include additional criteria such as ontogeny, molecular characteristics and parasitological testing. Furthermore, the host status is not always clear, e.g., D. kutzeri has been recorded from various deer species, with ambiguous species status (Cervus canadensis or C. elaphus canadensis).
Demodecid mites are generally characterized by monoxeny and a strict co-evolutionary relationship with their hosts, which is linked with the development of advanced adaptations to parasitism [9]. The mites also tend to inhabit different microhabitats within the host species, and synhospital (co-occurring) species are known to occur [46]. Although such species have been described for rodents, soricomorphs, bats, carnivores, ruminants and primates, the amount of knowledge regarding these species corresponds more to the amount of research performed on their hosts, rather than their actual distribution in the environment (Table 3). Until recently, the highest number of Demodecidae species were known from C. perspicillata, a species of bat that has been thoroughly studied for the presence of these mites (Table 3); in contrast, until the end of the 20th Century, only two demodecid species were known for the house mouse: A cosmopolitan, synanthropic animal used as a laboratory subject, pet or as food for other animals and one of the best-studied mammals. Intensive research conducted in recent years has revealed five further species of parasite, including one from a new genus: Glossicodex (Figure 5) [11,47].
Table
Table 1. Checklist of demodecid mites.
Table 1. Checklist of demodecid mites.
Demodecid MitesHost Species (Ordo, Family)OccurrenceComments to the Status of Demodecid Mite Species
Apodemodex cornutus Bukva, 1996Neomys anomalus Cabrera, 1907 (Soricomorpha, Soricidae)Czech Republic loc. class. [48]Valid
Demodex acutipes Bukva and Preisler, 1988Cervus elaphus Linnaeus, 1758 (Artiodactyla, Cervidae)Czech Republic loc. class. [49], Poland [50]Valid
Demodex aelleni Fain, 1960Myotis daubentonii (Kuhl, 1817) (Chiroptera, Vespertilionidae)Switzerland loc. class. [51]Valid
Demodex agrarii Bukva, 1994Apodemus agrarius (Pallas, 1771) (Rodentia, Muridae)Poland [46,52], Slovak Republic loc. class. [53]Valid
Demodex ailuropodae Xu, Xie, Liu, Zhou and Shi, 1986Ailuropoda melanoleuca (David, 1869) (Carnivora, Ursidae)China, zoological garden ex situ [54]Valid
Demodex antechini Nutting and Sweatman, 1970Antechinus stuartii Macleay, 1841 (Dasyuromorphia, Dasyuridae)Australia loc. class. [55]Valid
Demodex apodemi Hirst, 1918Apodemus agrarius (Rodentia, Muridae)Poland [52], Russia [56]Valid;
described by Hirst [57], next considered as a subspecies D. arvicolae var. apodemi [15], and verified by Izdebska [16] as D. apodemi;
specimens from A. agrarius, probably belongs to the separate species
Apodemus sylvaticus (Linnaeus, 1758) (Linnaeus, 1758)Great Britain loc. class [15,57], Poland [58], Russia [56]
Demodex araneae Nutting, 1950Ateles sp. (Primates, Atelidae)Nutting [39] after Nutting [59]Nom. nud.;
description not published within the meaning of the ICZN
Demodex aries Desch, 1986Ovis aries Linnaeus, 1758 (Artiodactyla, Bovidae)New Zealand loc. class [60], Czech Republic [61]Valid
Demodex artibei Vargas, Bassols, Desch, Quintero and Polaco, 1995Artibeus aztecus K. Andersen, 1906 (Chiroptera, Phyllostomidae)Mexico loc. class. [62]Valid
Demodex arvicolae Zschokke, 1888Microtus agrestis (Linnaeus, 1761) (Rodentia, Cricetidae)Astrahan/Europe on the border with Asia [56], Europe loc. class. [15]Valid;
the record from the M. arvalis is questionable; maybe D. microti was found
Microtus arvalis (Pallas, 1778) (Rodentia, Cricetidae)Astrahan/Europe on the border with Asia [56]
Demodex aurati Nutting, 1961Mesocricetus auratus (Waterhouse, 1839) (Rodentia, Cricetidae)Described and finding in laboratory animals, e.g., ex situ [37,63,64,65,66,67,68,69]Valid
Demodex auricularis Izdebska, Rolbiecki and Fryderyk, 2014Apodemus sylvaticus (Rodentia, Muridae)Poland loc. class. [58]Valid
Demodex bandicotae Izdebska, Rolbiecki, Morand and Ribas, 2017Bandicota indica (Beschstein, 1800) (Rodentia, Muridae)Laos loc. class. [17]Valid
Demodex bantengi Firda, Nutting and Sweatman, 1987Bos javanicus d’Alton, 1823 (Artiodactyla, Bovidae)Bali loc. class. [70]Valid
Demodex bicaudatus Kniest and Lukoschus, 1981Macroglossus minimus (E. Geoffroy, 1810) (Chiroptera, Pteropodidae)Australia loc. class. [71]Valid
Demodex bisonianus Kadulski and Izdebska, 1996Bison bonasus (Linnaeus, 1758) (Artiodactyla, Bovidae)Poland loc. class. [72,73,74,75,76,77,78]Valid
Demodex bonapartei Nutting, 1950Mustela erminea cicognanii Bonaparte, 1838 (Carnivora, Mustelidae)Nutting [39] after Nutting [59]Nom. nud.; description not published within the meaning of the ICZN
Demodex bovis Stiles, 1892
(redescription Desch and Nutting 1971)		Bos taurus Linnaeus, 1758 (Artiodactyla, Bovidae)Probably cosmopolitan, e.g., s. loc. [79], Egypt [80], Ethiopia [81], Nigeria [82], Sudan [83,84,85], Canada [86], USA [87], Argentina [88], Colombia [89], India [90], Mongolia [91], New Zealand [92,93], Czech Republic [94], Germany [95], Hungary [96], Italy [97], Poland [78,98]Valid;
European bison - an accidental finding in a closed farm
Bos taurus indicus Linnaeus, 1758 (Artiodactyla, Bovidae)
[B. indicus given by the Authors [99] is actually B. t. indicus]		Brazil [99]
Bison bonasus (Artiodactyla, Bovidae)
Poland, in the breeding condition ex situ [74,77]
Demodex brevis Akbulatova, 1963
(redescription, Desch and Nutting 1972)		Homo sapiens Linnaeus, 1758 (Primates, Hominidae)Cosmopolitan, e.g., USA [100,101,102], China [103,104], Australia [105], New Zealand [93], Poland [106], Russia loc. class. [107], Turkey [108,109,110]Valid
Demodex buccalis Bukva, Vítovec and Vlček 1985Myodes glareolus (Schreber, 1780) (Rodentia, Cricetidae)Czech Republic loc. class [111], Poland [112]Valid
Demodex caballi (Railliet, 1895) (redescription, Desch and Nutting 1978)Equus caballus Linnaeus, 1758 (Perissodactyla, Equidae)Probably cosmopolitan, e.g., s. loc. [113], USA [114,115], New Zealand [116]Valid
Demodex cafferi Nutting and Guilfoy, 1979Syncerus caffer (Sparrman, 1779) (Artiodactyla, Bovidae)Botswana loc. class. [117], Republic of South Africa [118,119]Valid
Demodex canis (Leydig, 1859) (redescription, Nutting and Desch, 1978)Canis lupus familiaris Linnaeus, 1758 (Carnivora, Canidae)Probably cosmopolitan, e.g., s. loc. [120], USA [121], Colombia [122], Cuba [123], India [124,125,126], Nepal [127], Pakistan [128], Thailand [129], New Zealand [93,130], Bangladesh [131], Poland [2,8,38,132,133], Russia [134], Turkey [135]Valid
Demodex caprae Railliet, 1895Capra hircus Linnaeus, 1758 (Artiodactyla, Bovidae)Probably cosmopolitan, e.g., Ethiopia [136], China [137], New Zealand [93], Czech Republic [138], France loc. class. [113], Poland [139], Switzerland [113,140]Valid
Demodex carolliae Desch, Lebel, Nutting and Lukoschus, 1971Carollia perspicillata (Linnaeus, 1758) (Chiroptera, Phyllostomidae)Republic of Suriname loc. class. [141]Valid
Demodex castoris Izdebska, Fryderyk and Rolbiecki, 2016Castor fiber Linnaeus, 1758 (Rodentia, Castoridae)Poland loc. class. [142]Valid
Demodex cati Megnin, 1877 (redescription, Desch and Nutting, 1979)Felis catus Linnaeus, 1758 (Carnivora, Felidae)Probably cosmopolitan, e.g., s. loc. [143], USA [144,145], Brazil [146], New Zealand [116], Bulgaria [147], Germany [148], Great Britain [15], Italy [149], Poland [132], Spain [150]Valid
Demodex caviae Bacigalupo and Roveda, 1954Cavia porcellus (Linnaeus, 1758) (Rodentia, Caviidae)Described and finding in laboratory animals, e.g., ex situ [151,152,153]Valid
Demodex cervi Prietsch, 1886Rusa unicolor (Kerr, 1792) (Artiodactyla, Cervidae)Germany ex situ [154]Valid;
hom. for D. cervi sensu Kutzer and Grünberg, 1972 (see D. kutzeri)
Demodex chiropteralis Hirst, 1921Plecotus auritus (Linnaeus, 1758) (Chiroptera, Vespertilionidae)Great Britain loc. class. [155], Poland [12]Valid
Demodex conicus Izdebska and Rolbiecki, 2015Mus musculus Linnaeus, 1758 (Rodentia, Muridae)Poland loc. class. [156]Valid
Demodex cornei: Izdebska and Rolbiecki, 2018Canis lupus familiaris Linnaeus, 1758 (Carnivora, Canidae)Probably cosmopolitan, Poland loc. class. [2]Valid;
records before 2018 have uncertain status, except Izdebska [8] and Izdebska and Fryderyk [38]
Demodex corniculatus Izdebska, 2012Apodemus flavicollis (Melchior, 1834) (Rodentia, Muridae)Poland loc. class. [1,16]Valid
Demodex criceti Nutting and Rauch, 1958Mesocricetus auratus (Waterhouse, 1839) (Rodentia, Cricetidae)Described and finding in laboratory animals, e.g., ex situ [37,65,66,68,69,157]Valid
Demodex cricetuli Hurley and Desch, 1994Cricetulus migratorius (Rodentia, Cricetidae)Described in laboratory animals ex situ [158]Valid
Demodex cuniculi Pfeiffer, 1903Oryctolagus cuniculus (Linnaeus, 1758) (Lagomorpha, Leporidae)In the breeding condition, e.g., China [15], Great Britain loc. class. [15,159]Valid
Demodex cyonis Morita, Ohmi, Kiwaki, Ike and Nagata, 2018Canis lupus familiaris Linnaeus, 1758 (Carnivora, Canidae)Japan loc. class. [160]Valid
Demodex dasypodi Desch and Stewart, 2002Dasypus novemcinctus Linnaeus, 1758 (Cingulata, Dasypodidae)USA loc. class. [161]Valid
Demodex desmodi Desch, 1994Desmodus rotundus (E. Geoffroy, 1810) (Chiroptera, Phyllostomidae)Republic of Suriname loc. class. [162]Valid
Demodex erinacei Hirst, 1917Erinaceus europaeus Linnaeus, 1758 (Erinaceomorpha, Erinaceidae)Great Britain loc. class. [163]Valid
Demodex erminae Hirst, 1919Mustela erminea Linnaeus, 1758 (Carnivora, Mustelidae)New Zealand [93], Great Britain loc. class. [15]Valid
Demodex equi Railliet, 1895Equus caballus Linnaeus, 1758 (Perissodactyla, Equidae)Probably cosmopolitan, e.g., s. loc. [113], USA [164], Great Britain [15], Poland [165]Valid
Demodex felisFelis catus Linnaeus, 1758 (Carnivora, Felidae)[40]Nom. nud.
Demodex flagellurus Bukva, 1985Mus musculus (Rodentia, Muridae)Czech Republic loc. class. [166], Poland [167,168,169]Valid
Demodex folliculorum (Simon, 1842) (redescription Desch and Nutting, 1972)Homo sapiens (Primates, Hominidae)Cosmopolitan, e.g., Algeria [170], Egypt [171], USA [100,101], China [103], India [172,173], Australia [105], New Zealand [93,130], Belgium [174], Croatia [175], Germany loc. class. [176], Great Britain [177], Greece [178], Iceland [179], Ireland [180], Poland [106], Turkey [108,109,110,181,182]Valid
Demodex folliculorum sinensis Xie, Liu, Hsu and Hsu, 1982Homo sapiens (Primates, Hominidae)China [183]
Demodex foveolator Bukva, 1984Crocidura suaveolens (Pallas, 1811) (Soricomorpha, Soricidae)Czech Republic loc. class. [184], Poland [18]Valid
Demodex fusiformis Izdebska and Rolbiecki, 2015Mus musculus (Rodentia, Muridae)Poland loc. class. [169]Valid
Demodex gapperi Nutting, Emejuaiwe and Tisolel, 1971Myodes gapperi (Rodentia, Cricetidae)USA loc. class. [185]Valid
Demodex gatoi Desch and Stewart, 1999Felis catus Linnaeus, 1758 (Carnivora, Felidae)USA loc. class. [186,187], Austria [188], Finland [189], Poland [44,132], Spain [150])Valid
Demodex ghanensis Oppong, Lee and Yasin, 1975Bos taurus (Artiodactyla, Bovidae)Ghana loc. class. [190], Sudan [83]Valid
Demodex glareoli Hirst, 1919Myodes glareolus (Rodentia, Cricetidae)Great Britain loc. class. [15], Poland [112]Valid;
described by Hirst [15] as a subspecies D. arvicolae var. glareoli, than verified by Izdebska [112] as
D. glareoli
Demodex gliricolens Hirst, 1921Arvicola amphibius (Linnaeus, 1758) (Rodentia, Cricetidae)Great Britain loc. class. [155]Valid
Demodex gracilentus Izdebska and Rolbiecki, 2013Apodemus agrarius (Rodentia, Muridae)Poland loc. class. [46]Valid
Demodex huttereri Mertens, Lukoschus and Nutting, 1983Apodemus agrarius (Rodentia, Muridae)Germany loc. class. [191], Poland [192]Valid
Demodex injai Desch and Hillier, 2003Canis lupus familiaris Linnaeus, 1758 (Carnivora, Canidae)Probably cosmopolitan, USA loc. class. [193], Brazil [194], Spain [195], Poland [8,38]Valid
Demodex intermedius Lukoschus, Mertesn, Nutting and Nadchatram, 1984Tupaia glis (Diard, 1820) (Scandentia, Tupaiidae)Malaysia loc. class. [196]Valid
Demodex kutzeri Bukva, 1987Alces alces (Linnaeus, 1758) (Atriodactyla, Cervidae)Poland [197]Valid;
(=D. cervi sensu Kutzer and Grünberg, 1972; hom. for D. cervi Prietsch, 1886)
Capreolus capreolus (Linnaeus, 1758) (Artiodactyla, Cervidae)Poland [198,199]
Cervus elaphus Linnaeus, 1758 (Artiodactyla, Cervidae)Austria [200], Czech Republic loc. class. [154], Poland [198,201]
Cervus elaphus nelsoni Nelson, 1902 (Artiodactyla, Cervidae)USA [202,203]
Cervus nippon pseudaxis Gervais, 1841 (Artiodactyla, Cervidae)Berlin, zoological garden ex situ [154]
Dama dama (Linnaeus, 1758)Poland [204]
Odocoileus hemionus hemionus (Rafinesque, 1817) (Artiodactyla, Cervidae)USA [202,203]
Odocoileus virginianus (Zimmermann, 1780) (Artiodactyla, Cervidae)USA [202]
Demodex lacrimalis Lukoschus and Jongman, 1974Apodemus sylvaticus (Rodentia, Muridae)Italy [205], Netherlands loc. class. [205], Poland [206]Valid
Demodex leucogasteri Hughes and Nutting, 1981Onychomys leucogaster (Wied-Neuwied, 1841) (Rodentia, Cricetidae)USA loc. class. [207]Valid
Demodex longior Hirst, 1918Apodemus sylvaticus (Rodentia, Muridae)Great Britain loc. class. [15,57], Poland [46], Russia [56]Valid
Demodex longissimus Desch, Nutting and Lukoschus, 1972Carollia perspicillata (Chiroptera, Phyllostomidae)Republic of Suriname loc. class. [208]Valid
Demodex lutrae Izdebska and Rolbiecki, 2014Lutra lutra (Linnaeus, 1758) (Carnivora, Mustelidae)Poland loc. class. [3]Valid
Demodex macaci Karjala, Desch and Starost, 2005Macaca mulatta (Zimmermann, 1780) (Primates, Cercopithecidae)USA, laboratory colony ex situ [209]Valid
Demodex macroglossi Desch, 1981Macroglossus minimus (Chiroptera, Pteropodidae)Australia loc. class. [210]Valid
Demodex marculus Izdebska and Rolbiecki, 2015Mus musculus (Rodentia, Muridae)Poland loc. class. [169]Valid
Demodex marsupiali Nutting, Lukoschus and Desch, 1980Didelphis marsupialis Linnaeus, 1758 (Didelphimorphia, Didelphidae)Republic of Surinam loc. class. [211]Valid
Demodex melanopteri Lukoschus, Jongman and Nutting, 1972Eptesicus brasiliensis melanopterus (Jentink, 1904) (Chiroptera, Vespertilionidae)
[E. melanopterus given by the Authors [212] is actually E. b. melanopterus]		Republic of Suriname loc. class. [212]Valid
Demodex melesinus Hirst, 1921Meles meles (Linnaeus, 1758) (Carnivora, Mustelidae)Great Britain loc. class. [213], Poland [4]Valid
Demodex merioni (=meriones)Meriones spp. (Rodentia, Muridae)Finding in laboratory animals, e.g., [36,37]Nom. nud.
Demodex mexicanus Vargas, Bassols, Desch, Quintero and Polaco, 1995Corynorhinus mexicanus G. M. Allen, 1916 (Chiroptera, Vespertilionidae)Mexico loc. class. [62]Valid
Demodex microti Izdebska and Rolbiecki, 2013Microtus arvalis (Rodentia, Cricetidae)Poland loc. class. [214]Valid
Demodex mollis Izdebska, Rolbiecki, Fryderyk and Mierzyński, 2017Apodemus flavicollis (Rodentia, Muridae)Poland loc. class. [1]Valid
Demodex molossi Desch, Nutting and Lukoschus, 1972Molossus molossus (Pallas, 1766) (Chiroptera, Molossidae)Republic of Suriname loc. class. [208]Valid
Demodex muscardini Hirst, 1917Muscardinus avellanarius (Linnaeus, 1758) (Rodentia, Gliridae)Armenia [56], Great Britain loc. class. [15,163]Valid
Demodex musculi Oudemans, 1897 (redescription, Izdebska and Rolbiecki, 2015)Mus musculus (Rodentia, Muridae)Europe [15], ds loc. class. [215], Poland [167,169], Russia [56], Spain [216]; laboratory animals, e.g., ex situ [7,217,218]Valid
Demodex myotidisMyotis lucifugus lucifugus (Le Conte, 1831) (Chiroptera, Vespertilionidae)Nutting [39] after Nutting [59]Nom. nud.; description not published within the meaning of the ICZN
Myotis septentrionalis Trouessart, 1897 (Chiroptera, Vespertilionidae)Nutting [39] after Di Benedetto [219]
Eptesicus fuscus (Beauvois, 1796) (Chiroptera, Vespertilionidae)Nutting [39] after Di Benedetto [219]
Demodex mystacina Desch, 1989Mystacina tuberculata Gray, 1843 (Chiroptera, Mystacinidae)New Zealand loc. class. [220]Valid
Demodex nanus Hirst, 1918 (redescription Desch, 1987)Rattus norvegicus (Berkenhout, 1769) (Rodentia, Muridae)Great Britain [57], Poland [19,221,222], Russia [56]; laboratory animals ex situ [223]Valid
Rattus rattus (Linnaeus, 1758) (Rodentia, Muridae)New Zealand [223], Great Britain loc. class. [57], Russia [56]
Demodex neomydis Bukva, 1995Neomys anomalus (Soricomorpha, Soricidae)Czech Republic loc. class. [224]Valid
Demodex neoopisthosomae Desch, Lukoschus and Nadchatram, 1986Eonycteris spelaea (Dobson, 1871) (Chiroptera, Pteropodidae)Malaysia loc. class. [225]Valid
Demodex norvegicus Bukva, 1995Rattus norvegicus (Rodentia, Muridae)Czech Republic loc. class. [226], Poland [19,221,222]Valid
Demodex novazelandica Desch, 1989Mystacina tuberculata (Chiroptera, Mystacinidae)New Zealand loc. class. [220]Valid
Demodex nycticeii Desch, 1996Nycticeius humeralis (Rafinesque, 1818) (Chiroptera, Vespertilionidae)USA loc. class. [227]Valid
Demodex odocoilei Desch and Nutting, 1974Odocoileus virginianus (Artiodactyla, Cervidae)USA loc. class. [228]Valid
Odocoileus hemionus columbianus (Richardson, 1829) (Artiodactyla, Cervidae)USA [229]
Demodex ovis Railliet, 1895 (redescription, Desch 1986)Ovis aries (Artiodactyla, Bovidae)s. loc. [113], Australia [60], New Zeland [60], Czech Republic [61], Israel [230], Poland [231]Valid
Demodex peromysci Lambert, Lukoschus and Whitaker, 1983Peromyscus leucopus (Rafinesque, 1818) (Rodentia, Cricetidae)USA loc. class. [232]Valid
Demodex phocidi Desch, Dailey and Tuomi, 2003Phoca vitulina Linnaeus, 1758 (Carnivora, Phocidae)USA, sealife center ex situ [233], Poland in situ [5]Valid
Demodex phodopi Desch, Davis and Klompen, 2006Phodopus sungorus (Pallas, 1773) (Rodentia, Cricetidae)Described in laboratory animals ex situ [234]Valid
Demodex phylloides Csokor, 1879Sus scrofa scrofa Linnaeus, 1758 (Artiodactyla, Suidae)Poland [235,236,237,238]Valid
Sus scrofa domesticus Erxleben, 1777 (Artiodactyla, Suidae)Probably cosmopolitan, e.g., Tanzania [239], Canada [240], USA [87], Brasil [241,242], New Zealand [93,130], historical Galicia loc. class. [243], Italy [244]
Demodex phyllostomatis Leydig, 1859Phyllostomus hastatus (Pallas, 1767) (Chiroptera, Phyllostomidae)Republic of Suriname loc. class. [51,120]Valid
Demodex plecoti Izdebska Rolbiecki, Mierzyński and Bidziński, 2019Plecotus auritus (Chiroptera, Vespertilionidae)Poland loc. class. [10]Valid
Demodex ponderosus Izdebska and Rolbiecki, 2014Rattus norvegicus (Rodentia, Muridae)Poland loc. class. [222]Valid
Demodex pseudaxis Schpringol’ts-Schmidt, 1937Cervus nippon hortulorum Swinhoe, 1864 (Artiodactyla, Cervidae)Russia/Far east [245]Valid;
need verification
Demodex ratti Hirst, 1917 (redescription Bukva, 1995)Rattus norvegicus (Rodentia, Muridae)Czech Republic [226], Europe, s. loc. [15,246], Poland [19,221,222,246,247,248], Russia [56]Valid
Demodex ratticola Bukva, 1995Rattus norvegicus (Rodentia, Muridae)Czech Republic loc. class. [226], Poland [222,247,248]Valid
Demodex rosus Bukva, Vítovec and Vlček, 1985Apodemus flavicollis (Rodentia, Muridae)Czech Republic loc. class. [111], Poland [249]Valid
Demodex sabani Desch, Lukoschus and Nadchatram, 1984Leopoldamys edwardsi (Thomas, 1882) (Rodentia, Muridae)Malaysia [250]Valid
Leopoldamys sabanus (Thomas, 1887) (Rodentia, Muridae)Malaysia loc. class. [250]
Niviventer cremoriventer (Miller, 1900) (Rodentia, Muridae)Malaysia [250]
Niviventer rapit (Bonhote, 1903) (Rodentia, Muridae)Malaysia [250]
Rattus annandalei (Bonhote, 1903) (Rodentia, Muridae)Malaysia [250]
Rattus tiomanicus (Miller, 1900) (Rodentia, Muridae)Malaysia [250]
Sundamys muelleri (Jentink, 1879) (Rodentia, Muridae)Malaysia [250]
Demodex saimiri Lebel and Nutting 1973Saimiri sciureus (Linnaeus, 1758) (Primates, Cebidae)Biological supply houses ex situ [251]Valid
Demodex sciurei Lebel, 1970Saimiri sciureus (Primates, Cebidae)[252]Nom. nud.; description not published within the meaning of the ICZN
Demodex sciurinus Hirst, 1923Sciurus vulgaris Linnaeus, 1758 (Rodentia, Sciuridae)Great Britain loc. class. [45], Poland [present study]Valid
Demodex sinocricetuli Desch and Hurley, 1997Cricetulus barabensis (Pallas, 1773) (Rodentia, Cricetidae)laboratory animals ex situ [253]Valid
Demodex soricinus Hirst, 1918 (redescription Bukva, 1993Plecotus auritus (Chiroptera, Vespertilionidae)
Great Britain [57,155]Valid; P. auritus -probably wrong host record
Sorex araneus Linnaeus, 1758
(Soricomorpha, Soricidae)
[S. vulgaris given by the Author [57] is actually S. araneus]		Czech Republic [254], Great Britain loc. class. [57], Poland [255]
Demodex spelaea Desch, Lukoschus and Nadchatram, 1986Eonycteris spelaea (Chiroptera, Pteropodidae)Malaysia loc. class. [225]Valid
Demodex suis (Kadlec, 1975)Sus scrofa domesticus (Artiodactyla, Suidae)s. loc. [256], s. loc. [257], Czech Republic [258]Nom abort.; syn. D. phylloides
Demodex sungori Desch, Davis and Klompen, 2006Phodopus sungorus (Rodentia, Cricetidae)Described in laboratory animals ex situ [234]Valid
Demodex sylvilagi Maravelas, 1962Sylvilagus transitionalis (Bangs, 1895) (Lagomorpha, Leporidae)Nutting [39] after Maravelas [259]Nom. nud.; description not published within the meaning of the ICZN
Demodex talpae Hirst, 1921Talpa europaea Linnaeus, 1758 (Soricomorpha, Talpidae)Great Britain loc. class. [155], Poland [260]Valid
Demodex tauri Bukva, 1986Bos taurus (Artiodactyla, Bovidae)Czech Republic loc. class. [261]Valid
Demodex tigris Shi, Xie and Hsu, 1985Panthera tigris amoyensis (Hilzheimer, 1905) (Carnivora, Felidae)China, zoological garden ex situ [262]
Valid
Demodex tortellinioides Desch and Holz, 2006Antechinus agilis Dickman, Parnaby, Crowther and King, 1998 (Dasyuromorphia, Dasyuridae)Australia loc. class. [263]Valid
Demodex transitionalis Moravelas, 1962Sylvilagus transitionalis (Bangs, 1895) (Lagomorpha, Leporidae)Nutting [39] after Maravelas [259]Nom. nud.; description not published within the meaning of the ICZN
Demodex uncii Desch, 1993Uncia uncia (Schreber, 1775) (Carnivora, Felidae)USA, zoological garden ex situ Desch [264]Valid
Demodex ursi Desch, 1995Ursus americanus Pallas, 1780 (Carnivora, Ursidae)USA loc. class. [265,266]Valid
Demodex vibrissae Izdebska, Rolbiecki and Fryderyk, 2016Mus musculus (Rodentia, Muridae)Poland loc. class. [47]Valid
Demodex zalophi Dailey and Nutting, 1979Zalophus californianus (Lesson, 1828) (Carnivora, Otariidae)USA, Australia loc. class. [267]Valid
Glossicodex musculi Izdebska and Rolbiecki, 2016Mus musculus (Rodentia, Muridae)Poland loc. class. [11]Valid
Ophthalmodex apodemi Bukva, Nutting and Desch, 1992Apodemus sylvaticus (Rodentia, Muridae)Czech Republic loc. class. [268]Valid
Ophthalmodex artibei Lukoschus and Nutting, 1979Artibeus lituratus (Chiroptera, Phyllostomidae)Republic of Surinam loc. class. [269]Valid
Ophthalmodex australiensis Woeltjes and Lukoschus, 1981Rhinonicteris aurantia (Gray, 1845) (Chiroptera, Hipposideridae)Australia loc. class. [270]Valid
Ophthalmodex carolliae Lukoschus, Woeltjes, Desch and Nutting, 1980Carollia perspicillata (Chiroptera, Phyllostomidae)Republic of Surinam loc. class. [271]Valid
Ophthalmodex juniatae Veal, Giesen and Whitaker, 1984Myotis lucifugus (Le Conte, 1831) (Chiroptera, Vespertilionidae)USA loc. class. [272]Valid
Ophthalmodex molossi Lukoschus, Woeltjes, Desch and Nutting, 1980Molossus molossus (Chiroptera, Molossidae)Republic of Surinam loc. class. [271]Valid
Ophthalmodex wilsoni Woeltjes and Lukoschus, 1981Vespadelus pumilus (Gray, 1841 (Chiroptera, Vespertilionidae)Australia loc. class. [270]Valid
Pterodex carolliae Lukoschus, Woeltjes, Desch and Nutting, 1980Carollia perspicillata (Chiroptera, Phyllostomidae)Republic of Suriname loc. class. [273]Valid
Rhinodex baeri Fain, 1959Galago moholi A. Smith, 1836
(Primates, Galagidae)		Rwanda loc. class. [274]Valid
Soricidex dimorphus Bukva, 1982Sorex araneus Linnaeus, 1758 (Soricomorpha, Soricidae)Czech Republic loc. class. [275,276], Poland [255]Valid
Stomatodex cercarteti Desch, 1991Cercartetus nanus (Desmarest, 1818) (Diprotodontia, Burramyidae)Australia loc. class. [277]Valid
Stomatodex corneti Fain, 1960 Valid
Stomatodex corneti corneti Fain, 1960Barbastella barbastellus (Schreber, 1774) (Chiroptera, Vespertilionidae)Belgium loc. class. [51], Great Britain [278]
Nycteris sp. (Chiroptera, Nycteridae)Rwanda [51]
Stomatodex corneti myotis Fain, 1960Myotis dasycneme (Boie, 1825) (Chiroptera, Vespertilionidae)Belgium [51]
Myotis myotis (Borkhausen, 1797) (Chiroptera, Vespertilionidae)Belgium [51]
Stomatodex galagoensis Fain, 1959Galago moholi A. Smith, 1836
(Primates, Galagidae)		Rwanda loc. class. [274]Valid
Stomatodex rousetti Fain, 1960Rousettus aegyptiacus (Geoffroy, 1810)
(Chiroptera, Pteropodidae)		Democratic Republic of the Congo loc. class. [51]Valid
Hom.: homonym, loc. class.: locus classicus, nom. abort.: nomen abortium, nom. nud.: nomen nudum, s. loco: sine loco, syn.: synonymum.
Table
Table 2. Body size (micrometers) for adults of Demodex sciurinus.
Table 2. Body size (micrometers) for adults of Demodex sciurinus.
Character♂ (n = 8)♀ (n = 13)
Length of gnathosoma15 (13–18), SD 217 (15–18), SD 1
Width of gnathosoma (at base)18 (15–23), SD 218 (14–20), SD 2
Length of podosoma49 (43–55), SD 554 (50–60), SD 3
Width of podosoma31 (28–35), SD 333 (30–38), SD 2
Length of opisthosoma72 (55–90), SD 1197 (75–125), SD 16
Width of opisthosoma31 (25–36), SD 433 (30–38), SD 2
Aedeagus20 (16–23), SD 3–
Vulva–6 (4–8), SD 1
Total length of body135 (120–158), SD 14168 (143–193), SD 17
Table
Table 3. Synhospital (co-occurring) demodecid mites in the same host species.
Table 3. Synhospital (co-occurring) demodecid mites in the same host species.
Mammals OrdoMammals SpeciesDemodecid Mites
PRIMATESGalago moholiRhinodex baeri
Stomatodex galagoensis
Homo sapiensDemodex brevis
Demodex folliculorum
RODENTIAApodemus agrariusDemodex agrarii
Demodex apodemi
Demodex gracilentus
Demodex huttereri
Apodemus flavicollisDemodex corniculatus
Demodex mollis
Demodex rosus
Apodemus sylvaticusDemodex apodemi
Demodex auricularis
Demodex lacrimalis
Demodex longior
Ophthalmodex apodemi
Mus musculusDemodex conicus
Demodex flagellurus
Demodex fusiformis
Demodex marculus
Demodex musculi
Demodex vibrissae
Glossicodex musculi
Rattus norvegicusDemodex nanus
Demodex norvegicus
Demodex ponderosus
Demodex ratti
Demodex ratticola
Mesocricetus auratusDemodex aurati
Demodex criceti
Myodes glareolusDemodex buccalis
Demodex glareoli
Phodopus sungorusDemodex phodopi
Demodex sungori
SORICOMORPHANeomys anomalusApodemodex cornutus
Demodex neomydis
Sorex araneusDemodex soricinus
Soricidex dimorphus
CHIROPTERACarollia perspicillataDemodex carolliae
Demodex longissimus
Ophthalmodex carolliae
Pterodex carolliae
Eonycteris spelaeaDemodex neoopisthosomae
Demodex spelaea
Macroglossus minimusDemodex bicaudatus
Demodex macroglossi
Molossus molossusDemodex molossi
Ophthalmodex molossi
Mystacina tuberculataDemodex mystacina
Demodex novazelandica
Plecotus auritusDemodex chiropteralis
Demodex plecoti
CARNIVORACanis lupus familiarisDemodex canis
Demodex cornei
Demodex cyonis
Demodex injai
Felis catusDemodex cati
Demodex gatoi
PERISSODACTYLAEquus caballusDemodex caballi
Demodex equi
ARTIODACTYLACervus elaphusDemodex acutipes
Demodex kutzeri
Cervus nipponDemodex kutzeri
Demodex pseudaxis
Odocoileus hemionusDemodex kutzeri
Demodex odocoilei
Odocoileus virginianusDemodex kutzeri
Demodex odocoilei
Bos taurusDemodex bovis
Demodex ghanensis
Demodex tauri
Ovis ariesDemodex aries
Demodex ovis

5. Conclusions

The Demodecidae have high veterinary and medical importance, and these aspects have directed the majority of research into the family. Despite the fact that such research stretches back to the 19th Century, appropriate zoological studies (taxonomy, fauna) are scarce and limited to species descriptions, typically based on singular records from one locality and a single host. More detailed biodiversity studies, based on the analysis of the occurrence of the parasite in different populations of individual host species, or the co-occurrence of different Demodecidae in one host species, have been published only in Poland. Even those studies have been limited by the availability of the host or the need to conduct comprehensive studies of its entire parasitofauna.
Unfortunately, no comparable data on Demodecidae are available from other regions; in addition, the group has never been included in holistic parasitofauna studies, or even studies of parasitic arthropods. The greatest obstacles to such studies are associated with parasite detection, particularly since most infestations are asymptomatic, and problems with species identification. This stands in contrast with the multitude of global medical and veterinary reports on demodecosis, or routine testing for this disease conducted at diagnostic laboratories.
There is a clear need to integrate zoological and parasitological research with medical and veterinary studies, or to perform further interdisciplinary studies. Only, such, broader approaches will provide a greater understanding of the key issues concerning Demodecidae parasitism, allow the development of efficient diagnostic methods and deepen our understanding of the causes and mechanisms of demodecosis.

Author Contributions

Conceptualization, J.N.I. and L.R.; Sampling, J.N.I. and L.R.; Microscopy Analysis, J.N.I. and L.R.; Data Analysis, J.N.I. and L.R.; Writing-Original Draft Preparation, J.N.I. and L.R.; Writing Review and Editing J.N.I. and L.R. All authors have read and agreed to the published version of the manuscript.

Funding

This research received no external funding.

Conflicts of Interest

The authors declare no conflict of interest.

References

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Figure 1. Demodex sciurinus: (A) female. (B) male.
Figure 1. Demodex sciurinus: (A) female. (B) male.
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Figure 2. Geographis distribution (green dots) of Demodecidae in the world.
Figure 2. Geographis distribution (green dots) of Demodecidae in the world.
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Figure 3. Share (%) of species from particular orders within mammals in comparison with the participation (%) of related demodecid mites.
Figure 3. Share (%) of species from particular orders within mammals in comparison with the participation (%) of related demodecid mites.
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Figure 4. Mammals—hosts of Demodecidae (species number and % of mammals from different orders).
Figure 4. Mammals—hosts of Demodecidae (species number and % of mammals from different orders).
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Figure 5. Topographic preferences of demodecid mites in house mouse: (A) Demodex conicus. (B) Demodex musculi. (C) Demodex marculus. (D). Demodex flagellurus. (E) Demodex fusiformis. (F) Demodex vibrissae. (G) Glossicodex musculi.
Figure 5. Topographic preferences of demodecid mites in house mouse: (A) Demodex conicus. (B) Demodex musculi. (C) Demodex marculus. (D). Demodex flagellurus. (E) Demodex fusiformis. (F) Demodex vibrissae. (G) Glossicodex musculi.
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Izdebska, J.N.; Rolbiecki, L. The Biodiversity of Demodecid Mites (Acariformes: Prostigmata), Specific Parasites of Mammals with a Global Checklist and a New Finding for Demodex sciurinus. Diversity 2020, 12, 261. https://doi.org/10.3390/d12070261

AMA Style

Izdebska JN, Rolbiecki L. The Biodiversity of Demodecid Mites (Acariformes: Prostigmata), Specific Parasites of Mammals with a Global Checklist and a New Finding for Demodex sciurinus. Diversity. 2020; 12(7):261. https://doi.org/10.3390/d12070261

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Izdebska, Joanna N., and Leszek Rolbiecki. 2020. "The Biodiversity of Demodecid Mites (Acariformes: Prostigmata), Specific Parasites of Mammals with a Global Checklist and a New Finding for Demodex sciurinus" Diversity 12, no. 7: 261. https://doi.org/10.3390/d12070261

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