# Taxonomic Catalog of the Brazilian Fauna: The Brazilian acarofauna (Arachnida: Holothyrida, Ixodida, Mesostigmata, Opilioacarida, Sarcoptiformes, and Trombidiformes)

Fernando C. Jacinavicius, Michel P. Valim, Leopoldo F.O. Bernardi, Juliana

C. Almeida, Gabrielle R. Andrade, Marcel S. Araújo, Marina F.C. Barbosa, Carlos Barreto, Ávyla R.A. Barros, Ricardo Bassini-Silva, et al.

► To cite this version:

Fernando C. Jacinavicius, Michel P. Valim, Leopoldo F.O. Bernardi, Juliana C. Almeida, Gabrielle R. Andrade, et al. Taxonomic Catalog of the Brazilian Fauna: The Brazilian acarofauna (Arachnida: Holothyrida, Ixodida, Mesostigmata, Opilioacarida, Sarcoptiformes, and Trombidiformes). Zoologia (Curitiba), 2025, 42, pp.e24081. 10.1590/s1984-4689.v42.e24081. hal-05490087

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RESEARCH ARTICLE
TAXONOMIC CATALOG OF THE BRAZILIAN FAUNA

Taxonomic Catalog of the Brazilian Fauna: The Brazilian acarofauna
(Arachnida: Holothyrida, Ixodida, Mesostigmata, Opilioacarida, Sarcoptiformes, and Trombidiformes)

Fernando C. Jacinavicius<fnref n="1" />, Michel P. Valim<fnref n="2" />,<fnref n="3" />, Leopoldo F.O. Bernardi<fnref n="4" />, Juliana C. Almeida<fnref n="5" />, Gabrielle R. Andrade<fnref n="6" />, Marcel S. Araújo<fnref n="7" />, Marina F.C. Barbosa<fnref n="8" />, Carlos Barreto<fnref n="9" />, Ávyla R.A. Barros<fnref n="8" />, Ricardo Bassini-Silva<fnref n="6" />, Darci M. Barros-Battesti<fnref n="10" />, Gabriel L. Bizarro<fnref n="11" />, Raphael C. Castilho<fnref n="8" />, Elizeu B. Castro<fnref n="12" />,<fnref n="13" />, Luiz A.S. Castro<fnref n="13" />, Peterson R. Demite<fnref n="13" />,<fnref n="14" />, Mércia E. Duarte<fnref n="15" />, Noeli J. Ferla<fnref n="16" />, Carlos H.W. Flechtmann<fnref n="8" />, Elizabeth Franklin<fnref n="17" />, Donald Gettinger<fnref n="18" />, Ryszard Haitlinger<fnref n="19" />, Fabio A. Hernandes<fnref n="20" />, Liana Johann<fnref n="21" />, Katarzyna Kaszewska<fnref n="22" />, Marcelo B. Labruna<fnref n="23" />, Antônio C. Lofego<fnref n="13" />, Natalia Marciniak<fnref n="22" />, João P.I. Martin<fnref n="8" />, Thiago F. Martins<fnref n="23" />,<fnref n="24" />, Carolina S. Mascarenhas²⁵, Andre L. Matioli²⁶, Gilberto J. Moraes<fnref n="8" />, Sebastián A. Muñoz-Leal²⁷, João P. Narita<fnref n="8" />, Denise Navia²⁸,²⁹, Ronald Ochoa³⁰, Heather C. Proctor³¹, Diana Rueda-Ramírez³², José M. Rezende<fnref n="13" />, Matheus S. Rocha<fnref n="16" />, Jandir C. Santos³³, Darliane E. Silva<fnref n="16" />, Guilherme L. Silva³⁴, Rafaela A. Silva³⁵, Vinícius B. Silva<fnref n="8" />, Maciej Skoracki<fnref n="21" />, Aline D. Tassi³⁶, Maria M. Vázquez-Gonzalez³⁷, Cal Welbourn³⁸

<footnote n="1">¹Departamento de Biologia Animal, Instituto de Biologia, Universidade Estadual de Campinas. Campinas, SP, Brazil.</footnote>

<footnote n="2">²Universidade Iguaçu. Nova Iguaçu, RJ, Brazil.</footnote>

<footnote n="3">³Universidade Santa Úrsula. Rio de Janeiro, RJ, Brazil.</footnote>

<footnote n="4">⁴Departamento de Entomologia, Universidade Federal de Lavras. Lavras, MG, Brazil.</footnote>

<footnote n="5">⁵Faculdade Idor de Ciências Médicas, Instituto D'or Pesquisa e Ensino. Rio de Janeiro, RJ, Brazil.</footnote>

<footnote n="6">⁶Laboratório de Coleções Zoológicas, Instituto Butantan. São Paulo, SP, Brazil.</footnote>

<footnote n="7">⁷Programa de Pós-Graduação em Ecologia e Recursos Naturais, Universidade Federal de São Carlos. São Carlos, SP, Brazil.</footnote>

<footnote n="8">⁸Departamento de Entomologia e Acarologia, Escola Superior de Agricultura “Luiz de Queiroz”, Universidade de São Paulo. Piracicaba, SP, Brazil.</footnote>

<footnote n="9">⁹Department of Biology, Algoma University. Sault Ste. Marie, Canada.</footnote>

<footnote n="10">¹⁰Departamento de Patologia, Reprodução e Saúde Única, Faculdade de Ciências Agrárias e Veterinárias, Universidade Estadual Paulista. Jaboticabal, SP, Brazil.</footnote>

<footnote n="11">¹¹Museu de Ciências Univates, Universidade do Vale do Taquari. Lajeado, RS, Brazil.</footnote>

<footnote n="12">¹²Instituto de Biociências, Universidade Estadual Paulista. Rio Claro, SP, Brazil.</footnote>

<footnote n="13">¹³Laboratório de Acarologia, Instituto de Biociências, Letras e Ciências Exatas, Universidade Estadual Paulista. São José do Rio Preto, SP, Brazil.</footnote>

<footnote n="14">¹⁴Instituto Federal de Roraima, Campos Bonfim. Bonfim, RR, Brazil.</footnote>

<footnote n="15">¹⁵Laboratório de Biologia, Ecologia e Sistemática de Artrópodes, Universidade Federal do Piauí. Floriano, PI, Brazil.</footnote>

<footnote n="16">¹⁶Laboratório de Acarologia, Universidade do Vale do Taquari. Lajeado, RS, Brazil.</footnote>

<footnote n="17">¹⁷Instituto Nacional de Pesquisas da Amazônia. Manaus, AM, Brazil.</footnote>

<footnote n="18">¹⁸Harold W. Manter Laboratory of Parasitology, University of Nebraska. Lincoln, Nebraska, USA.</footnote>

<footnote n="19">¹⁹Institute of Environmental Biology, Department of Invertebrate Systematics and Ecology, Wrocław University of Environmental and Life Sciences. Wrocław, Lower Silesian, Poland.</footnote>

<footnote n="20">²⁰Departamento de Ecologia e Zoologia, Universidade Federal de Santa Catarina. Florianópolis, SC, Brazil.</footnote>

<footnote n="21">²¹Programa de Pós-graduação em Biotecnologia, Universidade do Vale do Taquari. Lajeado, RS, Brazil.</footnote>

<footnote n="22">²²Department of Animal Morphology, Faculty of Biology, Adam Mickiewicz University. Poznan, Poland.</footnote>

<footnote n="23">²³Departamento de Medicina Veterinária Preventiva e Saúde Animal, Faculdade de Medicina Veterinária e Zootecnia, Universidade de São Paulo. São Paulo, SP, Brazil.</footnote>

<footnote n="24">²⁴Área Técnica de Doenças Vinculadas a Vetores e Hospedeiros Intermediários, Instituto Pasteur, Secretaria de Estado da Saúde de São Paulo. São Paulo, SP, Brazil.</footnote>
**ABSTRACT.** The diversity of organisms is one of the most fundamental questions in sciences. Here, we inventory species distributed in 273 families of mites from Brazil. Taxonomic information is provided for all 88 superfamiliies in Brazil, accompanied by relevant publications for the families, including notes on their identification and importance. However, despite recent taxonomic improvement in a few groups, particularly in Mesostigmata (Dermanysssoidea, Phytoseioidea and Uropodoidea), Ixodida (Ixodoidea), Sarcoptiformes (Analgoidea, Galumnoidea, Oppioidea, Oripodoidea, Phthiracaroidea, Pterlichioidea and Sarcoptoidea) and Trombidiformes (Arrenuroidea, Bdelloidea, Cheyletoidea, Eriophyoidea, Hygrobatoidea, Pygmephoroidea, Tarsonemoidea, Tetranychoidea, Trombiculoidea and Tydeoidea), knowledge remains limited for most other taxa. This limitation underscores the urgent need for taxonomic revisions and improvements in describing new species. Based on groups of host-specificity, we estimate nearly 50,000 species of mites occurring in Brazil, but the diversity could be greater. This enormous gap in our taxonomic knowledge of mites suggests that greater investment in training and hiring acarologists, is essential. In this context, the Taxonomic Catalog of the Brazilian Fauna (TCBF) should serve as a key platform for compiling, updating, and disseminating taxonomic data, ensuring that mite diversity in Brazil is systematically documented and accessible for future research and conservation efforts.

**KEY WORDS.** Acari, Mites, ticks, taxonomy, biodiversity, Neotropics, Brazil, checklist.

## INTRODUCTION

The traditional group Acari Leach, 1817, in class Arachnida Lamarck, 1801, was proposed to accommodate mites and ticks. It is organized into six orders: Holothyrida Thon, 1909; Ixodida Leach, 1815; Mesostigmata Canestrini, 1891; Opilioacarida With, 1902; Sarcoptiformes Reuter, 1909 and Trombidiformes Reuter, 1909. In a recent study by Zhang (2011), the number of described mite species worldwide was estimated to be about 55,000. This number shows that mites and ticks, along with spiders, are the most diverse subclasses of the class Arachnida (Zhang 2011).

Brief historical accounts of the development of Acarology in Brazil and its importance were presented by Flechtmann et al. (2006) and Moraes and Flechtmann (2008). As stated in those publications, the first Brazilian Symposium of Acarology, bringing together 180 professionals and students interested in this topic, was held in May, 2006, at Universidade Federal de Viçosa, Minas Gerais State. One of the main themes selected for discussion was the state of knowledge on mite diversity in the country, in an attempt to establish a list of priorities for studies, given the assumed importance of this type of information for the general development of acarology (Pallini et al. 2007).

<footnote n="25"><sup>25</sup>Instituto Federal Sul-rio-grandense, Campus Pelotas. Pelotas, RS, Brazil.</footnote>

<footnote n="26"><sup>26</sup>Laboratório de Acarologia, Instituto Biológico. Campinas, SP, Brazil.</footnote>

<footnote n="27"><sup>27</sup>Departamento de Ciencia Animal, Facultad de Ciencias Veterinarias, Universidad de Concepción. Chillán, Chile.</footnote>

<footnote n="28"><sup>28</sup>CBGP, INRAE, CIRAD, Institut Agro, IRD, Université de Montpellier. Montpellier, France.</footnote>

<footnote n="29"><sup>29</sup>UMR CBGP, Centre de Biologie pour la Gestion des Populations, Montferrier sur Lez cedex. France.</footnote>

<footnote n="30"><sup>30</sup>Systematic Entomology Laboratory, United States Department of Agriculture, Agricultural Research Service. Beltsville, MD, USA.</footnote>

<footnote n="31"><sup>31</sup>Department of Biological Sciences, University of Alberta. Edmonton, AB, Canada.</footnote>

<footnote n="32"><sup>32</sup>Ecology group, Institute of Biology, Humboldt-Universität zu Berlin. Berlin, Germany.</footnote>

<footnote n="33"><sup>33</sup>Independent Researcher. Ontario, Canada.</footnote>

<footnote n="34"><sup>34</sup>Programa de Pós-graduação em Ciências Médicas, Universidade do Vale do Taquari. Lajeado, RS, Brazil.</footnote>

<footnote n="35"><sup>35</sup>Universidade Estadual de Santa Cruz. Ilhéus, BA, Brazil.</footnote>

<footnote n="36"><sup>36</sup>University of Florida Tropical Research and Education Center. Homestead, Florida, USA.</footnote>

<footnote n="37"><sup>37</sup>Departamento de Ciencias, División de Ciencias e Ingeniería, Universidad de Quintana Roo. Chetumal, Quintana Roo, Mexico.</footnote>

<footnote n="38"><sup>38</sup>National Insect and Mite Collection, National Museum of Natural History, Smithsonian Institution. Washington, D.C., USA.</footnote>

<footnote>Corresponding author: Fernando de Castro Jacinavicius (fcjacinavicius@gmail.com)</footnote>

<footnote>https://zoobank.org/0B885B33-5AF3-42B2-9748-57F7900A8AD2</footnote>
That symposium generated interest in conducting a series of works in Brazilian research institutions, such as of biodiversity studies, including the description of new taxa and the preparation of catalogs and checklists dealing specifically with the Brazilian mite fauna of the following groups: Analgoidea Trouessart & Mégnin, 1884 and Pterolichoidea Trouessart & Mégnin, 1884 (Valim et al. 2011), Sarcoptoidea Murray, 1877 (Bochkov and Valim 2016), Tetranychidae Donnadieu, 1875 (Flechtmann and Moraes 2017), Ixodida (Barros-Battesti et al. 2006), Oribatida Dugès, 1834 (Oliveira et al. 2017), Trombiculidae Ewing, 1929, and Leeuwenhoekiidae Womersley, 1944 (Jacinavicius et al. 2018), EriophyoideaALEPA, 1898 (Navia et al. 2021), and Macronyssidae Oudemans, 1936 (Bassini-Silva et al. 2021).

In 2015, the “Taxonomic Catalog of the Brazilian Fauna (TCBF)”, an interactive tool, was created to host information gathered by over 500 specialists to constitute an online catalogue of the animal species reported from Brazil, presently accounting for over 125,138 valid species (Boeger et al. 2024). Concerning the Acari, the present status of that catalog was constructed by the effort of about 50 researchers, who accepted the invitation and became committed to establishing a list of the mite species reported to be present in the country. The maintenance of that catalog is a never-ending task, given the need for its continuous updating, as new data become available.

The objective of this publication is to present an ini-
tial account of the Brazilian Acari fauna within the scope
of TCBF, including references to about 3,678 species. This
publication is not intended to provide a taxonomic review
of mite groups but rather to compile literature information
on extant Brazilian mite biodiversity, with brief diagnosis of
their suprageneric features. Fossil and unpublished species
records will be treated in a different publication.

MATERIAL AND METHODS

In the “Taxonomic Catalog of the Brazilian Fauna”, an
effort has been made to place each species in the most up
to date classification. Thus, as in any database of this type,
changes in generic or family placement are expected to
occur, according to the publication of new findings.

Mites were classified into six orders based on Krantz
and Walter (2009), and the number of species considered in
this manuscript is based on the latest survey proposed by
Zhang (2011), although these numbers are nearly 15 years
out of date. We use this data as an indicator of diversity
rather than an estimate of the total number of currently

known species. Thus, based on the cited works, this study
includes six orders: Holothyrida, Ixodida, Mesostigmata,
Opilioacarida, Sarcoptiformes, and Trombidiformes.

Mesostigmata is divided into three suborders, while Trombidiformes is divided into two suborders (Krantz and Walter 2009, Beaulieu et al. (2011), Zhang et al. (2011). Although Krantz and Walter (2009) divided Sarcoptiformes into just two suborders (Endeostigmata and Oribatida, with the latter including Astigmata), some molecular studies (e.g., Dabert et al. 2010, Bolton et al. 2017, 2023) have placed Astigmata within Desmonomata (Oribatida), while others do not support its origin within Oribatida (Domes et al. 2007, Ban et al. 2022). Until the classification of Astigmata is better elucidated, we follow Krantz (1978) in maintaining Sarcoptiformes divided into three suborders (Astigmata, Endeostigmata, and Oribatida) as a matter of convenience, without reflecting phylogenetic relationships among these groups.

In the online system of TCBF (http://fauna.jbrj.gov.br/), we are limited to using the following taxonomic groups above family: superfamily, infraorder, suborder, order, superorder, infraclass, subclass, up to the class Arachnida. Taxonomic groups not used in TCBF are marked with an asterisk (*) in the list below.

RESULTS

The data presented include publications up to July 19, 2024. The numbers of valid genera and species at the supra-generic level reported in Brazil are provided in Appendix 1, compared with the respective global numbers. Other details about each species (distribution, hosts/substrates, type data, and references of mite records) are still being uploaded to the system by each expert. The entire dataset is updated periodically.

So far, 3,678 named mite species (about 6.5% of the
total for the world) in 273 families of 88 superfamilies have
been reported from Brazil. Twenty-two of the superfamilies
have been reported to contain more than 50 species in the
country. The accumulation curve of species per year shows
a significant rise in rates of acarofauna records in Brazil
from 1758 for Trombidiformes and Sarcoptiformes, 1772 for
Mesostigmata, 1787 for Ixodida (Ixodidae), 1844 for Ixodida
(Argasidae), 1905 for Opilioacarida and only 1999 for Holo-
thyrida onwards. (Figs 1, 2).

In the Parasitiformes Reuter, 1909, the most diverse
superfamilies in Mesostigmata are the Dermanyssoidea Ko-
lenati, 1859 (292 species), Phytoseioidea Berlese, 1916 (267),
and Uropodoidea Kramer, 1881 (221), followed by Ixodoidea
<figure><img src="image_3.png" /><figcaption>Figures 1–2. Accumulation curves of description year (1758–2024) of (1) Opilioacarida, Ixodida, and Holothyrida; and (2) Mesostigmata, Sarcoptiformes, and Trombidiformes reported in Brazil.</figcaption></figure>

Dugès, 1834 (80) in Ixodida; each of the other Parasitiformes superfamilies have less than 50 species (Fig. 3). Within this superorder, the superfamilies best represented in the country are Opilioacaroidea Redikorzev, 1937 (51% of the world species), followed by Heterozerconoidea Berlese, 1892 (38%), each of the other superfamilies represents at most 20% of the world diversity (Fig. 4).

Concerning the Acariformes Zakhvatkin, 1952, in Sarcoptiformes the most diverse superfamilies are the feather mites Analgoidea (227) and Pterolichoidea (104), followed by the Sarcoptoidea (85) and the oribatidid Oripodoidea Jacot, 1925 (83), Phthiracaroidea Perty, 1841 (77), Oppioidea Grandjean, 1951 (70), and Galumnoidea Jacot, 1925 (50); each of the other superfamilies have at most 50
<figure><img src="image_4.png" /></figure>

<figure><img src="image_5.png" /></figure>

Figures 3–4. (3) Number of Brazilian species per superfamily of the superorder Parasitiformes; and (4) proportion of Parasitiformes species per superfamily occurring in Brazil compared with world diversity. Red bar: Holothyrida, orange bar: Ixodida, yellow bars: Mesostigmata (Monogynaspida: Gamasina), green bars: Mesostigmata (Monogynaspida: Uropodina), light blue bars: Mesostigmata (Sejida), dark blue bars: Mesostigmata (Trigynaspida), and purple bar: Opilioacarida.
<figure><img src="image_6.png" /><figcaption>Figure 5. Number of Brazilian species per superfamily of the order Sarcoptiformes. Red bars: Astigmata, yellow bar: Endeostigmata, and blue bars: Oribatida.</figcaption></figure>

species (Fig. 5). In comparison with the world diversity of the order, the Ctenacaroidea Grandjean, 1954 accounts for 45% of the species, followed by the Caleremaeoidea Grandjean, 1965 (43%) and Nematalycoidea Strenzke, 1954 (33%) (Fig. 6). In Trombidiformes, the best represented superfamilies are Eriophyoidea (265), Hygrobatoidea Koch, 1842 (245), Tetranychoidea Donnadieu, 1875 (173), Arrenuroidea Thor, 1900 (151), Trombiculoidea Ewing, 1929 (89), Pygme-

phoroidea Cross, 1965 (67), Tydeoidea Kramer, 1877 (66), Cheyletoidea Leach, 1815 (60), Bdelloidea Dugès, 1834 (51), and Tarsonemoidea Canestrini and Fanzago, 1877 (50) (Fig. 7). In comparison with the world diversity of groups in the order, the Trochometridioidea Mahunka, 1970 accounts for 29%, followed by Arrenuroidea Thor, 1900 (11%), Labi-dostomatoidea Ewing, 1934 (12%), and Hygrobatoidea Koch, 1842 (10%) (Fig. 8).
<figure><img src="image_7.png" /><figcaption>Figure 6. Proportion of Sarcoptiformes species per superfamily occurring in Brazil in comparison with the world diversity. Red bars: Astigmata, yellow bar: Endeostigmata, and blue bars: Oribatida.</figcaption></figure>
<figure><img src="image_8.png" /></figure>

Figure 7. Number of Brazilian species per superfamily of the order Trombidiformes. Red bar: Labidostommatina, yellow bars: Eupodina, blue bars: Anystina, and green bars: Eleutherengona.

<figure><img src="image_9.png" /></figure>

Figure 8. Proportion of Trombidiformes species per superfamily occurring in Brazil compared with the world diversity. Red bar: Labidostommatina, yellow bars: Eupodina, blue bars: Anystina, and green bars: Eleutherengona.
### Class Arachnida Lamarck, 1801
### Superorder Parasitiformes Reuter, 1909*

This superorder comprises the orders Holothyrida, Ixodida, Mesostigmata and Opilioacarida.

### Order Holothyrida Thon, 1905

Holothyrida is a small order. The free-living edaphic mites of this order are found in forests and comprise only the superfamily Holothyroidea Thorell, 1882.

### Holothyroidea Thorell, 1882

Holothyroidea is divided into Allothyridae Hammen, 1972, Holothyridae Thorell, 1882; and Neothyridae Lehtinen, 1981 (Lehtinen 1991). In Brazil, only Neothyridae is presented, represented by two species of the genus *Diplothyrus* Lehtinen, 1999 (Lehtinen 1999, Vázquez et al. 2016).

### Order Ixodida Leach, 1815

Ixodida is the order comprising the mites commonly known as ticks. They are cosmopolitan ectoparasites of terrestrial vertebrates (amphibians, reptiles, birds and mammals) and some species accidentally attack humans (Guglielmone and Robbins 2018, Nogueira et al. 2022). Ixodida comprises a single superfamily, Ixodoidea. In Brazil, 77 species from two families have been reported.

### Ixodoidea Dugès, 1834

Ixodoidea is organized into three families, of which Argasidae Murray, 1877 and Ixodidae Murray, 1877 are represented in Brazil (Nava et al. 2017, Guglielmone et al. 2014). Argasidae comprises a group commonly referred to as soft ticks, containing 222 species (Nava et al. 2017, Muñoz-Leal et al. 2023). Ixodidae, commonly called hard ticks, contains 760 species (Guglielmone et al. 2014, 2021). The Brazilian tick fauna currently comprises 24 species of Argasidae and 53 species of Ixodidae (Guglielmone et al. 2021, Muñoz-Leal et al. 2023, Soares et al. 2023).

In Brazil, soft and hard ticks are the main arachnid vectors of pathogenic agents to humans and domesticated and wild animals. Therefore, some species are highly relevant to animal welfare and public health (Guglielmone and Robbins 2018, Nogueira et al. 2022). They vector bacteria such as *Ana-plasma* Theiler, *Borrelia* Swellengrebel, *Ehrlichia* Moroshkowsky and *Rickettsia* da Rocha-Lima, as well as protozoa, such as *Babesia* Starcovici, *Cytauxzoon* Neitz & Thomas, *Hepatozoon* Miller, *Rangelia* Carini & Maciel and *Theileria* Bettencourt, Franca & Borges (Barros-Battesti et al. 2006, Nava et al. 2017).

### Order Mesostigmata Canestrini, 1891

Mites of this order have long been known as potential biological control agents of agricultural pests; species of Phytoseiidae have been most widely studied for that purpose. Other mesostigmatids are known as important parasites of vertebrates and invertebrates. In Brazil, 990 species of 55 families have been reported. Species of the three suborders of Mesostigmata (Monogynaspida, Sejida and Trigynaspida) have been reported from Brazil.

### Suborder Monogynaspida Camin & Gorirossi, 1955

This is the most diverse mesostigmatid suborder, in terms of morphology, biology and ecology. They are commonly found on plants, in the soil and litter, in stored products, or associated with insects or vertebrates. Although mostly known as predators, many feed on fungi, pollen, and sweet substrates or are parasites of invertebrates and vertebrates. The suborder comprises two infraorders: Gamasina Kramer, 1881 and Uropodina Kramer, 1881 (Lindquist et al. 2009). Representatives of both infraorders have been reported from Brazil, comprising 724 and 238 species, respectively.

### Infraorder Gamasina Kramer, 1881

Gamasina are organized into four hyporders and, 11 superfamilies, of which seven superfamilies have been known to occur in Brazil. One of these belongs to the hyporder Parasitiae (Parasitoidea Oudemans, 1901) while the remaining belong to the hyporder Dermanyssiae (Ascoidea Voigts & Oudemans, 1905; Eviphidoidea Berlese, 1913; Dermanyssoidea Kolenati, 1859; Phytoseioidea Berlese, 1916; Rhodacaroidea Oudemans, 1902; and Veigaioidea Oudemans, 1939). Dermanyssoidea contains many parasitic mite groups and some predators, whereas other superfamilies consist predominantly of predators (except Ameroseiidae Evans, 1961, mostly fungivorous and pollinivorous). Representatives of the remaining hyporders, Arctacariae and Epicriiae, have not been reported from Brazil. Among other aspects, Gamasina are characterized by having the epigynal shield not surrounded by fused sternal, endopodal and ventral shields, usually bearing *st*5.

### Ascoidea Voigts & Oudemans, 1905

This superfamily comprises four families, three of which have been reported from Brazil, namely Ameroseiidae, Ascidae Voigts and Oudemans, 1905, and Melicharidae Hirschmann, 1962 (Lindquist et al. 2009, Lindquist and Moraza 2014). In addition to the predatory habit, they are also known to be fungivorous and pollinivorous.
Eight species of Ameroseiidae have been reported from the country. A world catalog of mites of this family was published by Mašán (2017), and included 138 species in 12 genera. One new genus and three new species were described since that publication (Abo-Shnaf et al., 2023), when these authors also recharacterized the family. Biological studies and observations have suggested that some species exhibit a fungivorous habit (Flechtmann 1985, Mohamed et al. 1985, Moustafa and El-Hady 2006), and less often a predacious habit (Elhalawany et al. 2021). In contrast, other species are nectar and pollen feeders (Seeman and Walter 1995, Seeman 1996, Kar et al. 2015).

Ascidae species have been described from a wide range of habitats mostly from edaphic environments but also from plants or associated with and phoretic upon insects and mammals. Most species have been cited as predators, feeding on other mites or small arthropods. In Brazil, 22 species have been reported (Moraes et al. 2016, Santos et al. 2023a).

Most Melicharidae species have been cited as predators of small arthropods or consuming sugary substrates or pollen (Moraes et al. 2016). In Brazil, 24 species have been reported (Moraes et al. 2016, Santos et al. 2023b). Morphological details of each genus of both Ascidae and Melicharidae can be found in the catalog published by Moraes et al. (2016), which refers respectively to 372 and 206 species. Databases concerning world distribution and substrates of mites of these families are available on line (Santos et al. 2023a, 2023b, 2023c).

## Dermanyssoidea Kolenati, 1859

Mites of this group have diverse feeding habits, and many species are associated with vertebrates or invertebrates (parasitically or phoretically), while many others are free-living (Dowling and O'Connor 2010). In Brazil, representatives of 12 of the 17 families listed by Beaulieu et al. (2011) have been reported: Dasyponyssidae Fonseca, 1940, Dermanyssidae Kolenati, 1859, Entonyssidae Ewing, 1923, Haemogamasidae Oudemans, 1926, Halarachnidae Oudemans, 1906, Ixodorhynchidae Ewing, 1923, Laelapidae Berlese, 1892, Macronyssidae Oudemans, 1936, Rhinonyssidae Trouessart, 1895, Spelaeorhynchidae Oudemans, 1902, Spinturnicidae Oudemans, 1901 and Varroidae Delfinado & Baker, 1974. Representatives of Hystriconyssidae Keegan, Yunker & Baker, 1960, Larvamimidae Elzinga, 1993, Manitheronyssidae Radovsky & Yunker, 1971, and Omentolaelapidae Fain, 1961, cited by Beaulieu et al. (2011) as members of Dermanyssoidea have not been reported from Brazil. Iphiopsidiae Kramer, 1886, also included by Beaulieu et al. (2011) is in this work included in Laelapidae, as done in Moraes et al. (2022).

Dasyponyssidae is a small family consisting of only three species in two genera (*Dasyponyssus* Fonseca, 1940 and *Xenarthronyssus* Radovsky & Yunker, 1971), which has been little studied (Radovsky and Yunker 1971). This family is only found in the Neotropical region, exclusively as armadillo parasites (Dowling 2006). One of the three species of this family has been reported from Brazil.

Dermanyssidae is a slightly larger family, consisting of about 26 species also in two genera: *Dermanyssus* Dùges, 1834 (bird parasites), and *Liponyssoides* Hirst, 1913 (mainly rodent parasites) (Roy and Chauve 2007, 2009). Only two species of this family have been reported from Brazil, including *Dermanyssus gallinae* (DeGeer, 1778), and *Liponysoides sangineus* (Hirst, 1914). The first species has a global distribution and is commonly reported as chicken mite or poultry red mite, a serious parasite worldwide (Tucci 1997).

Entonyssidae is a group of parasites inhabiting the respiratory system of snakes, including the bronchi, trachea and lungs. Fain (1961b) divided the family into two subfamilies, namely Entonyssinae Ewing, 1923, with representatives in Africa, Asia, Europe and North America, and Pneumophonyssinae Fonseca, 1940, which has been reported from South America. This family contains eight genera and 27 species (Fajfer 2012). Two of these species belong to Pneumophionyssinae and are the only entonyssids reported from Brazil: *Entophioptes liophis* Fain, 1960, and *Pneumophionyssus aristoterisi* Fonseca, 1940.

Halarachnidae consists of species commonly reported from the respiratory system of marine and terrestrial mammals. They are known to feed on blood as well as on other tissues, causing heavy infections that can lead to the host's death. Of the seven genera listed in Beaulieu et al. (2011), four have been reported from Brazil, *Orthohalarachne* Newell, 1947; *Pneumonyssoides* Fain, 1955; *Pneumonyssus* Banks, 1901; and *Raillietia* Trouessart, 1902. *Orthohalarachne* is represented in Brazil by *O. attenuata* (Banks, 1910) and *O. diminuata* (Doetschman, 1944), parasites of marine mammals of the order Pinnipedia (Gastal et al. 2016). *Pneumonyssus* Fonseca, 1940 and *Pneumonyssoides* Fain, 1955 are represented by one species each, *Pneumonyssus simicola* Banks, 1901, a parasite of rhesus monkeys, *Macaca mulatta* (Zimmermann, 1780), collected from a research breeding colony of Institute Oswaldo Cruz, Rio de Janeiro, and *Pneumonyssoides caninum* (Chandler & Ruhe, 1940), associated with domestic dogs (Andrade and Marchevsky 2007, Parraz et al. 2010). *Raillietia* species, namely *Raillietia auris* (Leidy, 1872), *R. caprae* Quintero, Bassols & Acebedo, 1980, and *R. flechtmanni* Faccini, Leite & Costa, 1992 (Faccini et al. 1992,
Faccini and Ribeiro 2008). Unlike, Beaulieu et al. (2011), some researchers consider *Raillietia* to belong to Raillietidae Vitzthum, 1943.

Ixodorhynchidae is represented by approximately 30 species of six genera, all snake ectoparasites. They are fixed under the host's scales, often attached to the head, around the eyes or in the gular region. Four of the six world genera are represented in Brazil, *Chironobius* Lombert & Moss, 1983, *Ixobioides* Fonseca, 1934, *Ophiogongylus* Lombert & Moss, 1983, and *Strandtibbettsia* Fain, 1961 (Fonseca 1934, Fain 1961a, Lizaso 1983).

Laelapidae is one of the most numerous and spe-
cies-rich mesostigmatid families, has highly variable feed-
ing habits. Many species are free-living while others are
associated with invertebrates, birds and mammals. Moraes
et al. (2022) published a catalog of the free-living and ar-
thropod-associated laelapids, some of which also contained
some species of parasitic habits. The vertebrate parasites
are commonly found on the hosts and in nests and other
microhabitats where they live (Strandtmann and Wharton
1958, Radovsky 1985, Dowling 2006, Dowling and O'Connor
2010). Among the laelapids recorded from Brazil, *Laelaps
echidninus* Berlese, 1887; *L. nuttalli* Hirst, 1915 and *Andro-
*laelaps fahrenholzi* (Berlese, 1911) are cosmopolitan species
(Linardi et al. 1984), while species of *Gigantolaelaps* Fonseca,
1939; *Mysolaelaps* Fonseca 1936 and *Tur* Baker & Wharton,
1952, have an almost exclusively Neotropical distribution
(Strandtmann and Wharton 1958, Furman 1972). In Brazil,
a total of 109 species of Laelapidae have been recorded.

Macronyssidae constitute a large group of over 230
parasitic species organized into 35 genera (Radovsky 2010,
Bassini-Silva et al. 2021). The most ancestral species are
apparently bat (Chiroptera) associated, whereas the most
derived species are associated with other vertebrates (Ra-
dovsky 2010). More than 30 macronyssid species have been
reported from Brazil (Bassini-Silva et al. 2021), including
six of *Ornithonyssus* Sambon, 1928, four of *Parichoronyssus*
Radovsky, 1967, three of *Macronyssus* Kolenati, 1858, two each
of *Chiroptonyssus* Augustson, 1945, *Lepronyssoides* Fonseca,
1941, *Pellonyssus* Clark & Yunker, 1956 and *Radfordiella*
Fonseca, 1948, and one each of *Acanthonyssus* Yunker &
Radovsky, 1966, *Chiasmanyssus* Gomes-Almeida & Pepato,
2021, *Macronyssoides* Radovsky, 1966, *Mitonyssoides* Yunker,
Lukoschus & Giesen, 1990, *Mitonyssus* Yunker & Radovsky,
1980, *Ophionyssus* Mégnin, 1884, *Steatonyssus* Kolenati, 1858
and *Thigmonyssus* Radovsky, 2010. In addition to their impor-
tance as parasites, Bassini-Silva et al. (2019a, 2021) considered
the possibility that some species could cause dermatitis to

their definitive and accidental hosts, for example, *Ornithonys-
sus bursa* (Berlese, 1888) lives in nests and parasitizes birds
and bites human, such as when these nests are abandoned.
Some species are also been suspected to be involved in the
epidemiological cycle of pathogens affecting their usual
hosts (Fonseca 1935, Reeves et al. 2007, Hornok et al. 2012,
Amaral et al. 2018).

Rhinonyssidae is a large family of parasitic mites that inhabit bird respiratory systems, being found preferentially on the membrane covering the nasal turbinates. There are approximately 600 species described worldwide (Domrow 1969, Dimov 2020), of which 123 species have been reported from Brazil. *Ptilonyssus* Berlese & Trouessart, 1889 is the genus with the highest number of species recorded in the country (70 species), followed by *Sternostoma* Berlese & Trouessart, 1889 (16 species), *Tinaminyssus* Strandtmann & Wharton, 1958 (15 species), *Rhinonyssus* Trouessart, 1894 (seven species), *Passeronyssus* Fain, 1960, *Rallinyssus* Strandtmann, 1948, *Rhinoecius* Cooreman, 1946 (two species each), *Larinyssus* Strandtmann, 1948, *Pipronyssus* Fain & Aitken, 1967 and *Sternoecius* Fain & Aitken, 1967 (one species each). Although the majority of the hosts are native birds, some mites have been reported parasitizing captive exotic birds, and introduced species (Santos et al. 2018, Mascarenhas et al. 2018, 2022).

Spelaeorhynchidae is a small family consisting of just seven species of the single genus *Spelaeorhychus* Neuman, 1902, all parasites of Neotropical chiropterans (Peracchi 1990). Three of the species of this family have been reported from Brazil: *Spelaeorhynchus hutsoni* Martyn, 1988; *S. jimi* Peracchi, 1990; and *S. praecursor* Neumann, 1902. Spinturnicidae Oudemans, 1902 are also obligate parasites of bats of all groups, usually living mainly on the wings or tail membranes of their hosts (Rudnick 1960); rarely the adult females of some species are found on the face, ears, and anal orifices of their hosts (Rudnick 1960, Herrin and Tipton 1975). Spinturnicids have a worldwide distribution. This family comprises about 110 species of 12 genera (Beron 2020a). In Brazil, 17 spinturnicid species of four genera have been reported, including 13 in *Periglischrus* Kolenati, 1857, two in *Spinturnix* Von Heyden, 1826 and one each in *Cameronieta* Machado-Allison, 1965 and *Mesoperiglischrus* Dusbábek, 1968.

Varroidae Delfinado & Baker, 1974 are also known as
bee mites or varroa mites. This is a small family of only six
species in two genera, *Euvarroa* Delfinado and Baker, 1974
and *Varroa* Oudemans 1904 (Delfinado and Baker 1974). A
single species was recorded from Brazil, *Varroa destructor*
Anderson & Trueman, 2000, formerly identified as *Varroa jacobsoni* Oudemans, 1904 (Anderson and Trueman 2000), and is the species that causes economic impact to honey bees (Rosenkranz et al. 1988, De Jong 1997, Sammataro et al. 2000). It is worth noting that the validity of this family has been recently questioned based on advances in molecular technology, with it being considered a subfamily within *Laelapidae* (Oh et al. 2024).

## Eviphidoidea Berlese, 1913

Mites of this superfamily are mainly edaphic predators. However, many species inhabit ephemeral habitats such as dung and decaying dead organisms and therefore have phoretic relationships with various arthropods for dispersal from deteriorating habitats. This superfamily comprises *Eviphididae* Berlese, 1913, *Leptolaelapidae* Karg, 1978, *Macrochelidae* Vitzthum, 1930, *Pachylaelapidae* Berlese, 1913 and *Parholaspididae* Evans, 1956 (Beaulieu et al. 2011). Although not explicitly stated, the comparison by Fonseca (1946) and Cómbita-Heredia et al. (2018) of *Pachylaelapidae* with *Megalolaelapidae* Fonseca, 1946 suggests that the latter is best placed within *Eviphidoidea*.

*Eviphididae* includes both non-phoretic forms inhabiting relatively stable substrates and phoretic forms inhabiting ephemeral habitats. Members of this family are predators of nematodes and other organisms such as insect larvae and eggs (Castilho et al. 2015, Rueda-Ramírez et al. 2023). Of the 108 world species of this family, only one has been reported from Brazil.

*Macrochelidae* is by far the largest family of this superfamily. It comprises mites that inhabit most humus or temporary habitats such as excrement and decaying animals. Several species of this family are found phoretic upon other animals, mainly insects, but also mammals and occasionally birds (Lindquist et al. 2009, Azevedo et al. 2015). Several studies have suggested that adults of macrochelid species prefer feeding on fly larvae or eggs, while immatures of the same species prefer feeding on nematodes (Rodriguez et al. 1962, Axtell 1969, Ito 1977, Rueda-Ramírez et al. 2023). About 470 species are placed in this family, with *Macrocheles* Latreille, 1829 being the most diverse genus (Azevedo et al. 2015). In Brazil, 46 species have been reported, of which *Macrocheles* is the most diverse, with 27 reported species.

*Megalolaelapidae* is a group of large mites commonly associated with dung beetles. Studies with *Megalolaelaps colossus* Cómbita-Heredia & Quintero-Gutierrez, 2018 established that its association with Coleoptera is necessary for the survival of the mite, although it seems not to be parasitic

on insects, as no feeding punctures have been observed (Cómbita-Heredia et al. 2020). The same authors suggested that this species feeds on nematodes as well as on fly larvae and eggs. Currently, this family comprises a single genus with seven species, three of which have been reported from Brazil (Mašán and Halliday 2014, Cómbita-Heredia et al. 2018).

*Parholaspididae* includes predominantly edaphic predatory mites, but some species have been reported from moss or tree holes. In contrast with other *Eviphidoidea*, phoresis has not been reported in *parholaspidids* (Lindquist et al. 2009). This family is reported to comprise 163 species, two of which have been reported from Brazil. Morphological details of each genus can be found in the catalog published by Quintero-Gutiérrez and Halliday (2021).

*Leptolaelapidae* is a group of morphologically very diverse mites found in soil, associated with insects and bird nests (Costa 1975, Karg 1997, Beaulieu and Walter 2007). *Pachylaelapidae* includes edaphic species and species found on mosses and in nests of social insects, mammals and birds. Mites of this family are known as predators of the microfauna in their habitats (Lindquist et al. 2009, Mašán and Halliday 2014, Castilho et al. 2015). A review of the *Pachylaelapidae* was published by Mašán and Halliday (2014). No species of these last two families have been reported from Brazil.

## Parasitoidea Oudemans, 1901

This superfamily contains a single family, *Parasitidae* Oudemans, 19101. This family is divided into two subfamilies, *Parasitinae* and *Pergamasinae* (Krantz and Walter 2009). Altogether, the family contains 420 species. These mites are most numerous in temperate habitats, and they have not been extensively collected in Brazil. *Parasitids* are found mostly in soils and mosses, whereas some live in freshwater, and saltwater marshes or beaches. They prey on microarthropods and nematodes, but *parasitines* are frequently found as deutonymphs phoretically on insects. No species of this family has been nominally reported from Brazil, although unidentified *parasitid* species have occasionally been reported.

## Phytoseioidea Berlese, 1916

This group comprises the families *Blattisociidae* Garman, 1948, *Otopheidomenidae* Treat, 1955, *Phytoseiidae* Berlese, 1916, and *Podocinidae* Berlese, 1913. It includes predatory, parasitic, fungivorous and pollinivorous species from plants, animals and edaphic habitats (Lindquist et al. 2009).

*Blattisociidae* constitutes a diverse group of mites associated with plants, fungi, soil and other animals (inver-
tebrates and vertebrates) (Moraes et al. 2015). It includes two subfamilies: Blattisociinae and Platyseiinae (Lindquist et al. 2009). This family presently contains more than 400 described species, about half of them from a single genus, *Lasioseius* Berlese, 1916 (Moraes et al. 2015, 2016, Santos et al. 2016, 2023a, 2023b, 2023c). Due to the large number of endemic species (approximately 80%), it seems that this family originated from the Neotropical region (Santos et al. 2016). In Brazil, 33 species of five genera have been reported, with *Lasioseius* being the most diverse genus (17 species) (Santos et al. 2023c). Morphological details of each genus of Blattisociidae can be found in the catalog published by Moraes et al. (2016). A database concerning world distribution and substrates of blattisociid mites is available on-line (Santos et al. 2023c).

Otopheidomenidae comprises a distinct group of insect parasites found in temperate and tropical regions of the world (Lindquist et al. 2009). This problematically phytoseioid family comprises 28 species (Beaulieu et al. 2011). However, some species are revised by Lindquist et al. (2020) and are actually replaced in other families (eg., Laelapidae: Dermanyssoidea), also suggesting that Otopheidomenidae be included within Dermanyssoidea. We will maintain the classification proposed by Krantz and Walter (2009), including Otopheidomenidae in Phytoseioidea, until the placement of the Phytoseioidea families, including molecular data, is carried out for validation of the placement of these groups of mites inside the gamisine. In Brazil, only two species are reported.

Podocinidae consists of species that inhabit humus and litter habitats in temperate and tropical regions (Lindquist et al. 2009), with one report on a rodent (Yan et al. 2011) and a small genus, *Africoseius* Krantz, 1962, found on plants (Rueda-Ramirez et al. 2019). Some species have been found to feed on springtails, nematodes and other mites (Wong 1967, Lindquist et al. 2009, Rueda-Ramírez et al. 2019). Podocinidae is divided into two subfamilies (Rueda-Ramírez et al. 2019), Africoseiinae and Podocinae. Currently the family has 38 species, 31 of which are in *Podicinum* Berlese, 1882 (Barros et al. 2020). In Brazil, six species have been reported (Santos et al. 2017, 2019, Rueda-Ramirez et al. 2019).

Phytoseiidae is the largest family of Phytoseioidea. It is of major economic importance, composed primarily of predatory mites found most commonly on aerial plant parts (McMurtry et al. 2015). Despite their practical importance as natural enemies of small arthropods (mostly mites and insects), species of this group are known to consume pollen, plant exudates and fungi as part of their diets or as alternative food in the absence of the prey (McMurtry et al. 2013). Some phytoseiid species have been produced and commercialized worldwide

(including Brazil) for the biological control of phytophagous mites and small insects (McMurtry et al. 2013, 2015, Knapp et al. 2018). This family is currently composed of approximately 2,650 valid species (Demite et al. 2023) organized in three subfamilies: Amblyseiinae, Phytoseiinae and Typhlodromiinae (Chant and McMurtry, 2007). Amblyseiinae is the largest, with approximately 65% of the described species (Demite et al. 2014). Almost 250 species have been reported from Brazil. *Amblyseius* Berlese 1914, with 47 species, is the most diverse genus recorded in this country (Lofego et al. 2024).

## Rhodacaroidea Oudemans, 1902

This superfamily is composed of the families Digamasellidae Evans, 1957, Halolaelapidae Karg, 1965, Laelaptonyssidae Womersley, 1956, Ologamasidae Ryke, 1962, Rhodacaridae Oudemans, 1902, and Teranyssidae Halliday, 2006 (Castilho et al. 2012). These are free-living and cosmopolitan mites, generally reported as predators (Lindquist et al. 2009, Castilho et al. 2012, 2015, 2016). They are widespread in soil, litter, mosses, lichens, manure, decaying wood, galleries of bark beetles, associated with termites, in nests of birds and small mammals, and other types of organic matter (Castilho et al. 2012, 2015, 2016). According to Beaulieu et al. (2011), the three largest families are Digamasellidae, Ologamasidae and Rhodacaridae. Only representatives of these families have been reported from Brazil.

Digamasellidae includes about 277 species worldwide, of which well over half belongs to *Dendrolaelaps* Halbert, 1915 (Castilho 2012). About 65% of the digamasellids were described from the Palearctic region. Ologamasidae is a much larger family, including 470 species worldwide (Castilho et al. 2016). Most ologamasids were described from the Australasian and Neotropical regions (about 32% each) (Castilho et al. 2015). Rhodacaridae includes about 160 species worldwide. Most rhodacarids were described from the Afrotropical (about 42%) and Palearctic (about 27%) regions. Some species have been reported as potential biological control agents of soil pests (Castilho et al. 2009, Rossini et al. 2022). In Brazil, these three families comprise 3, 25, and 11 species, respectively. World catalogs of the Rhodacaridae and Ologamasidae were published by Castilho et al. (2012, 2016).

## Veigaioidea Oudemans, 1939

This superfamily consists of a single family, Veigaiidae Oudemans, 1939, containing about 100 species. They are predatory soil-inhabitants, common in temperate forests. They are not common in Brazil, from where a single species, *Gamasolaelaps whartoni* (Farrier, 1957), has been reported.
## Infraorder Uropodina Kramer, 1881

The second infraorder of Monogynaspida, Uropodina, is more homogeneous than the, Gamasina, and it is not divided into hyporders. Among other aspects, these mites are characterized by having the epigynal shield surrounded by the fused sternal, endopodal and ventral shields, usually bearing no setae. It is divided into four superfamilies (Diarthrophalloidea Trägårdh, 1946, Microgynioidea Trägårdh, 1942, Thinozerconoidea Halbert, 1915, and Uropodoidea Kramer, 1881). Only representatives of the first and the last of these have been reported from Brazil.

## Diarthrophalloidea Trägårdh, 1946

This is a small superfamily consisting of a single family, Diarthrophallidae, which contains about 63 species worldwide. All developmental stages of these mites are found in association with beetles of Passalidae, suggesting that the beetles provide them some nourishment. In Brazil, six species have been reported.

## Uropodoidea Kramer, 1881

This is a much more diverse superfamily, consisting of 30 families, with species richness varying from three species in Baloghjkaszabiidae Hirschmann, 1979 to 401 species in Trematuridae Berlese, 1917, for a total of 2,209 species. Commonly found in decaying organic substrates, but also occasionally in nests or different animals, especially as phoretic deutonymphs on arthropods, to which each deutonymph attaches by a pedicel secreted through the anal opening. They are predators of nematodes, fly larvae, bacteria, fungi, and organic debris. The Uropodoidea is rather well represented in Brazil, with 232 mite species in 16 families.

## Suborder Sejida Kramer, 1885

Mites of this group are characterized, among other aspects, by having the oviporus covered by a single plate usually bearing at least six setae (rarely 2-4), tarsus IV with 20 pairs of setae (with setae av4 and pv4 on an intercalary sclerite between basitarsus and telotarsus) and movable cheliceral digit without excrescences. A phylogenetic study of this group, based on morphological and molecular data, was published by Lekveishvili and Klompen (2004). Species of this group have been reported as predators of small arthropods and nematodes, especially in litter of moist tropical and subtropical forests, under bark, in tree holes and in mammal nests. This suborder comprises two superfamilies Heterozerconoidea Berlese, 1892 and Sejoidea Berlese, 1885, with a total of about 80 species.

## Heterozerconoidea Berlese, 1892

Heterozerconoidea consists of only two families, Discozerconidae Berlese, 1910 and Heterozerconidae Berlese, 1892, the latter being commonly associated with smooth and cylindrical hosts of very different groups, millipedes, snakes and worm-lizards (Klompen and Gerdeman 2023). Discozerconidae has not been reported from Brazil. Worldwide, Heterozerconidae consists of 13 species, five of which have been reported from Brazil: two species each of *Amhet-erozercon* Fain, 1989 and *Heterozercon* Berlese, 1892, and one species of *Maracozercon* Fain, 1989 (Fain 1989).

## Sejoidea Berlese, 1885

Lindquist et al. (2009) included three families in this superfamily, namely Ichthyostomatogasteridae Sellnick, 1953, Sejidae Berlese, 1885 and Uropodellidae Berlese, 1888. Walter (2013) described a new family, Reginacharlottiidae Walter, 2013, which was included in this superfamily. Currently, this latter family contains a single species, described from Brazil, whereas Uropodellidae Camin, 1955 contains six species, one of which was reported from Brazil. According to H. Klompen (pers. communication, november 2023), no sejid species has been reported from Brazil, probably because of insufficient efforts in looking for them, given their known common occurrence on Cerambycidae in tropical areas. A key to the species of *Uropodella* Berlese, 1888 was presented by Lindquist and Vorontsov (2023). A diagnosis of Sejidae Berlese, 1885 and of the genera included in it was published by Lekveishvili and Klompen (2006).

## Suborder Trigynaspida Camin & Gorirossi, 1955

This suborder is divided in two infraorders (Lindquist et al. 2009): Antennophorina Camin & Gorirossi, 1955 (seven superfamilies) and Cercomegistina Camin & Gorirossi, 1955 (one superfamily). Members of the following superfamilies of Antennophorina were reported from Brazil: Antennophoroidea Camin & Gorirossi, 1955, Celaenopsoidea Berlese, 1892, Fedrizioidea Trägårdh, 1937, Megisthanoidea Berlese, 1914 and Parantennuloidea Willmann, 1941. Aenictequoidea Kethley, 1977 and Paramegistoidea Trägårdh, 1946, of Antennophorina, as well as the single superfamily of Cercomegistina (Cercomegistoidea Trägårdh, 1937) have not been reported from Brazil.

## Antennophoroidea Camin & Gorirossi, 1955

The superfamily contains a single family, Antennophoridae Berlese, 1892, containing 19 species, two of which are reported from Brazil.
### Celaenopsoidea Berlese, 1892

This superfamily contains eight families, of which Euzerconidae Trägårdh, 1938, Diplogyniidae Trägårdh, 1941, Celaenopsidae Berlese, 1892 and Triplogyniidae Funk, 1977 have been reported from Brazil, comprising 15 species.

### Fedrizzioidea Trägårdh, 1937

This superfamily contains two families, of which only two species of Klinckowstroemiidae Camin & Gorirossi, 1955, *Antennurella trouessarti* Berlese, 1904 and *Klinckowstroemiella helleri* (Oudemans, 1929), have been reported from Brazil.

### Megistanoidea Berlese, 1914

This superfamily contains two families, of which only Hoplomegistidae Camin and Gorirossi, 1955 have been reported from Brazil, with only one representative, *Stenosternum armiger* (Berlese, 1888).

### Parantennuloidea Willmann, 1941

This superfamily contains three families, of which only Parantennulidae Willmann, 1941 have been reported from Brazil, also with only one representative, *Micromegistus bakeri* Tragardh, 1948.

### Order Opilioacarida Zakhvatkin, 1952

Opilioacarida is represented by a single superfamily, Opilioacaroidea Redikorzev, 1937, with one family of free-living edaphic mites, Opilioacaridae With, 1904. The group consists of 13 genera and 65 described species (Araújo et al. 2020, Bernardi et al. 2023).

They are found in forests (Araújo et al. 2022), deserts (Vázquez et al. 2022) and caves (Bernardi et al. 2023), with records from most zoogeographic zones, except of the Antarctica, Pacific Islands, and Eastern Palearctic regions (Araújo et al. 2020). Many of its taxa are considered endemic (Bernardi et al. 2012, Vázquez and Klompen 2015), with restricted distribution and sparse occurrences (Vázquez and Klompen 2010, Bernardi et al. 2014).

In Brazil, Opilioacaridae is known from 19 described species, distributed across the North, Northeast, South and Southeast regions, with no species described for the Central-West region. These account for about 28% of the world known species, belonging to four genera: *Neocarus* Chamberlin & Mulaik, 1942 (15 species), *Amazonacarus* Vázquez, De Araújo & Feres, 2014 (two species) and one species each of *Brazilacarus* Vázquez, De Araújo & Feres, 2015 and *Caribeacarus* Vázquez & Klompen, 2009.

### Superorder Acariformes Zakhvatkin, 1952*

This superorder comprises the orders Sarcoptiformes and Trombidiformes.

### Order Sarcoptiformes Reuter, 1909

The Sarcoptiformes are here organized into three groups to comply the TCBF system and does not reflect the current phylogenetic relationships of these groups: Astigma-ta, the greatest economic and health importance to humans and other animals, the soil mites (Oribatida), and the small and enigmatic Endeostigmata. This group is mostly found in terrestrial environments, although less commonly also found in aquatic environments. In Brazil, 1,122 species belonging to 138 families have been reported.

### Suborder Astigmata Canestrini, 1891

This group is organized into ten superfamilies: Acaroidea Latreille, 1802, Analgoidea Trouessart & Mégnin, 1884, Canestrinioidea Berlese, 1884, Glycyphagoidea Berlese, 1897, Hemisarcoptoidea Oudemans, 1904, Histiostomatoidea Berlese, 1897, Hypoderatoidea Murray, 1877, Pterolichoidea Trouessart & Mégnin, 1884, Sarcoptoidea Murray, 1877 and Schizoglyphoidea Mahunka, 1978. All except the last super-family have been reported from Brazil.

### Acaroidea Latreille, 1802

Members of Acaroidea infest stored food products, decaying organic matter, plant pests and inhabitants of vertebrate or insect nests (Khaing et al. 2014). This super-family comprises six families, of which Acaridae Latreille, 1802, Gaudiellidae Atyeo, Baker & Delfinado, 1974 and Suidasiidae Hughes, 1948 have been reported from Brazil, with respectively 14, four and two species.

### Analgoidea Trouessart & Mégnin, 1884

Together with the Pterolichoidea, part of these mites constitutes what has been termed the "true feather mites". In Brazil, 15 of the 19 analgoidean families of feather mites have been reported: Alloptidae Gaud, 1957, Analgidae Trouessart & Mégnin, 1884, Avenzoariidae Oudemans, 1905, Dermationidae Fain, 1965, Dermoglyphidae Mégnin & Trouessart, 1884, Epidermoptidae Trouessart, 1892, Proctophyllodidae Trouessart & Mégnin, 1884, Psoroptoididae Gaud, 1958, Pteronyssidae Oudemans, 1941, Ptyssalgidae Atyeo & Gaud, 1979, Pyroglyphidae Cunliffe, 1958, Thysanocercidae Atyeo & Peterson, 1972, Trouessartiidae Gaud, 1957 and Xolalgidae Dubinin, 1953. Although feather mites from various families
have been described on passerine birds (Hernandes and Valim 2015, Mironov et al. 2016), recent years have seen notable contributions to the improved understanding of mites commonly collected on the wings of Passeriformes birds, particularly the Proctophyllodidae (Hernandes and Valim 2005, 2006, 2012, Hernandes et al. 2007, Valim and Hernandes 2006, 2008, 2009, 2010, Hernandes 2021). These studies have revealed that dozens of new Brazilian species await description. The main references reporting anal-goidean mites in Brazil can be found in Valim et al. (2011), Pedroso and Hernandes (2016), Hernandes (2020, 2021) and Hernandes and Flechtmann (2020).

Besides the feather mites, some families are endoparasites. The species in Cytoditidae Oudemans, 1908 have been found in the air sacs and respiratory tracts of birds, as well as in the coelomic cavity (Fain and Bafort 1964). Large infestations can be responsible for granulomatous pneumonia, aerosaculitis and peritonitis (Herpich et al. 2012). Of three species in Brazil, two were described from native birds, the white-bellied parrot and the grey-breasted martin (Mascarenhas et al. 2018) and one from domestic chicken (Smaniotto et al. 2014). Mites of Laminosioptidae Vitzthum, 1931 parasitize the connective tissues of birds. They invade the subcutaneous tissue causing the formation of small nodules (Fain 1981). They have occasionally been reported to cause lesions in the abdomen, lungs and peritoneum. After the death of the mite, a nodule of calcareous infiltration is formed. In Brazil, only the worldwide important fowl cyst mite, *Laminosioptes (Laminosioptes) cysticola* (Vizioli, 1870) has been reported, on domesticated chickens (Martins et al. 2010, Herpich et al. 2012). The family Turbinoptidae Fain, 1957 is composed exclusively of mites living in the nasal passages of birds worldwide. In Brazil, only two species have been reported, one parasitizing the waved woodpecker and another described from domestic chicken (Castro and Pereira 1951, Fain and Aitken 1970).

## Canestrinioidea Berlese, 1884

Canestrinioidea comprises three families, Canestrinidae Berlese, 1884, Heterocoptidae Fain, 1967, and Lemanniellidae Wurst, 2001, with the first being the most representative of the three (Haitlinger 2012). Members of this family are beetle associates or parasites. In Brazil, 46 species have been reported. The Brazilian Canestrinioidea are mostly unknown, considering the few species described and the many (35,699) species of Coleoptera reported from the country (Caron et al. 2024).

## Glycyphagoidea Berlese, 1897

Members of Glycyphagoidea comprises seven families, of which Aeroglyphidae Zakhvatkin, 1941, Chortoglyphidae Berlese, 1897, Echimyopodidae Fain, 1967, Glycyphagidae Berlese, 1897, and Rosensteiniidae Cooreman, 1954 have been reported from Brazil by one, one, five, seven and three species, respectively.

As typical glycyphagoids, species of Rosensteiniidae and Echimyopodidae are inhabitants of vertebrate nests that share a suite of characters common to early nidicolous taxa (Acaroidea and Hypoderatoidea) suggesting a common ancestry (Krantz and Walter 2009). In Brazil, three species of Rosensteiniidae have been reported, directly associated with bats or with their guano (Fain 1963, Fain and Flechtmann 1978). Echimyopodidae Fain, 1967 are also mostly nest-associated, but all genera – with the exception of *Blomia* – have follicular or subcutaneous deutonymphs. In Brazil, three of these have been reported, two from rodents' tails and the other from a rodent's stomach, and other two species from house dust and often related with allergic symptoms in humans (Galvão and Guitton 1986, Binotti et al. 2001). Glycyphagidae, Aeroglyphidae and Chortoglyphidae have been reported from house dust and stored products and some species are known to be the sources of allergens (Binotti et al. 2001, Arlian et al. 2009).

## Hemisarcoptoidea Oudemans, 1904

Hemisarcoptoidea comprises seven families, Algophagidae Fain, 1975, Carpoglyphidae Oudemans, 1923, Chaetodactylidae Zachvatkin, 1941, Hemisarcoptidae Oudemans, 1904, Hyadesiidae Halbert, 1915, Meliponocoptidae Fain & Rosa, 1983, and Winterschmidtiidae Oudemans, 1923. In Brazil, Carpoglyphidae, Chaetodactylidae, Hyadesiidae, Meliponocoptidae, and Winterschmidtiidae are recorded with 33 species.

## Histiostomatoidea Berlese, 1897

Members of Histiostomatoidea comprise two families: Guanolichidae Fain, 1968 and Histiostomatidae Berlese, 1897. In Brazil only Histiostomatidae has been reported with seven species listed.

## Hypoderatoidea Murray, 1877

This superfamily consists of a single family, Hypoderatidae Murray, 1877, reported to contain a total of 77 species (Seeman and Walter 2023), and divided in two subfamilies. One of these, Hypoderatinae Murray, 1877, has been found
on birds and in their nests (O'Connor 1985, Mironov and O'Connor 2013). The other, Muridectinae Fain, 1968, consists of five species distributed in two genera and are associated with desert rodents (Fain and Lukoschus 1986). Most species of this family are known only by their parasitic deutonymphal stage (hypopus) found in the subcutaneous tissues of the hosts, while the other stages (larva, protonymph, tritonymph and adults) live in the host's nest (Mironov and Kivganov 2010). The only species reported from Brazil is *Hypodectes propus* (Nitzsch, 1861), in the nest of *Zenaida auriculata* (Aves: Columbiformes: Columbidae) (Boas Filho et al. 2006).

## Pterolichoidea Trouessart & Mégnin, 1884

Together with the Analgoidea, mites of this group constitute what has been called the “true feather mites”, which live permanently on birds. In their monograph about the feather mites of the world, Gaud and Atyeo (1996) included a third superfamily, Freyanoidea Dubinin, 1953, but later authors have demonstrated this taxon to be part of the Pterolichoidea (Ehrnsberger et al. 2001, Klimov and O'Connor 2008). Feather mites are associated with all major bird orders worldwide and from the 33 orders in Brazil (Pacheco et al. 2021), feather mites have been reported from at least 27 orders (Valim et al. 2011, Enout et al. 2012, Pedroso et al. 2015, Hernandes et al. 2015, Hernandes and O'Connor 2015, Hernandes 2019).

Members of 10 of the 18 pterolichoidean families have been reported from Brazil, namely: Ascouracaridae Gaud & Atyeo, 1976 (four species in Brazil), Crypturoptidae Gaud, Atyeo & Berla, 1973 (11 species), Eustathiidae Oudemans, 1905 (six species), Falculiferidae Oudemans, 1905 (eight species), Freyanidae Dubinin, 1953 (six species), Gabuciniidae Gaud and Atyeo, 1975 (17 species), Kramerellidae Gaud & Mouchet, 1961 (one species), Pterolichidae Trouessart & Mégnin, 1884 (50 species), Ptiloxenidae Gaud, 1982 (one species), and Syringobiidae Trouessart, 1897 (one species).

## Sarcoptoidea Murray, 1877

This superfamily contains obligate and permanent symbionts of therian mammals, as commensals or parasites. These mites and the bird-associated symbionts (Analgoidea and Pterolichoidea) constitute three major phylogenetic branches within the psoroptidian Astigmata (O'Connor 1982, 2009). Sarcoptoidea currently includes about 1,100 species in 12 families (Bochkov 2010, Bochkov and Mironov 2011). In Brazil, 87 species of the following 11 families have been reported: Atopomelidae Gunther, 1942, Chirodiscidae Trouessart, 1892, Listrophoridae Mégnin & Trouessart,

1884, Psoroptidae Canestrini, 1892, Sarcoptidae Murray, 1877, Gastronyssidae Fain, 1956, Lemurnyssidae Fain, 1957, Listropsoralgidae Fain, 1965, Lobalgidae Fain, 1965, Myocoptidae Gunther, 1942, and Rhyncoptidae Lawrence, 1956 (Bochkov and Valim 2016).

Sarcoptoidea comprises parasitic species belonging to five main ecological groupings: (I) the fur-mites or pilicolous mites, including Atopomelidae and Listrophoridae on therian mammals, and Lobalgidae on placental mammals, are adapted to live on hairs of their diverse small mammal hosts; (II) mites inhabiting the hairs of therian mammals including Chirodiscidae with species exclusively occurring on bats; (III) mite inhabitants of the skin of therian mammals, including Myocoptidae, Listropsoralgidae, Psoroptidae and Sarcoptidae, with species specialized to live on superficial skin layers (stratum spinosum and stratum corneum); (IV) intracutaneous mites inhabitants of hair follicles of therian mammals including Rhyncoptidae; and (V) the endoparasites, Gastronyssidae and Lemurnyssidae, inhabitants of the nasal passages of Primates (Bochkov et al. 2008, Bochkov 2010).

Infestation of domestic animals with one of the several genera of parasitic mites causes mange, a skin condition. Some species of this superfamily are mange agents of medical and/or veterinary importance. The species reported from Brazil include: Sarcoptidae – *Sarcoptes scabiei* (Linnaeus, 1758) (sarcoptic mange, zoonotic); *Notoedres (Notoedres) cati* (Hering, 1838) (notoedric mange); *Trixacarus caviae* Fain, Howell and Hyatt, 1972 (guinea pig mange); Psoroptidae – *Psoroptes ovis* (von Hering, 1838) (psoroptic mange); *Otodectes cynotis* (von Hering, 1838) (otodectic mange); *Chorioptes bovis* (von Hering, 1845) (chorioptic mange); Atopomelidae – *Chirodiscoides caviae* Hirst, 1917 (guinea pig fur mite); Myocoptidae – *Myocoptes (Myocoptes) musculinus* (Koch, 1844) (fur mite of mice).

## Suborder Endeostigmata Reuter, 1909

Endeostigmata is probably a paraphyletic group presumed to include the most primitive Acariformes. They are organized in four infraorders, namely Bimichaeliida O'Connor, 1984, Nematalycina Lindquist, Krantz & Walter, 2009, Terpnacarida O'Connor, 1984 and Alicorhagiida O'Connor, 1984 (Walter et al. 2011). In Brazil, only members of the Nematalycina have been recently reported. This infraorder comprises the single superfamily Nematalycoidea Strenke, 1954.

## Nematalycoidea Strenke, 1954

Nematalycoidea comprises three families, Micropsamnidae Coineau & Theron, 1983, Nematalycidae Strenke,
1954 and Proteonematalycidae Kethley, 1989. In Brazil, only Nematalycidae has been reported. This is an unusual family of soil mites with vermiform body adapted for living in small interstitial spaces in the mineral soil fraction (Haupt and Coineau 1999). Two species of this family have been reported from Brazil, namely *Gordialycus tuzetae* Coineau, Fize and Delamare Deboutteville, 1967 and *Psammolycus delamarei* Schubart, 1973 (Schubart 1973, Norton at al. 2008). Recent molecular phylogenetic evidence, suggests that Eriophyoidea is a sister group of Nematalycidae (Klimov et al. 2018), instead of a member of the infraorder Eupodina Krantz, 1978.

## Suborder Oribatida van der Hammen, 1968

Members of this suborder are often well-sclerotized and dark brown, however, colorless, lightly sclerotized, and yellowish species also occur (Behan-Pelletier and Lindo 2023). They exhibit K-style life history traits, including low reproductive output and long-life spans (on average 1–2 years) (Norton and Behan-Pelletier 2009). Evolutionarily, their long life-span partly resulted in selection for defenses, especially camouflage, cuticular hardening and defensive structures like glands and diverse body forms (Norton and Behan-Pelletier 2009, Peschel et al. 2006).

Oribatid mites inhabit most terrestrial environments and are often the dominant or most abundant arthropod group. In natural ecosystems, they often occur in high densities (Norton and Behan-Pelletier 2009). They are very diverse, with >11,500 named species in 163 families (Subías 2023) in most soils and other organic-rich detrital systems, like moss-dominated habitats (Barreto et al. 2021, 2023).

Most oribatid mite species are particle-feeding saprophages (i.e., they consume dead plants and animals) and mycrophages (i.e., they consume fungi) (Norton 2007). However, some species have been shown to feed on mosses, protozoans and nematodes (Lehmitz and Maraun 2016). Members of all five infraorders of Oribatida are present in Brazil. In total, 582 species have been reported, as summarized by Oliveira et al. (2017).

## Infraorder Desmonomata Woolley, 1973

This is the most diverse infraorder, with two hyporders (Nothrina van der Hammen, 1982 and Brachypylina Hull, 1918). Brachypylina (colloquially 'higher oribatids') are the oribatid mites that have the traditional aspect of the Oribatida, and comprise the bulk of species (Behan-Pelletier and Lindo 2023). In Brazil, 430 species of Nothrina and Brachypylina have been reported, classified into 24 superfamilies and 67 families as summarized below in the Appendix 1.

## Achipterioidea Thor, 1929

Achipterioidea is composed of three families worldwide: Epactozetidae Grandjean, 1930, Tegoribatidae Grandjean, 1954, Achipteriidae Thor, 1929 (Subías 2023). In Brazil, the superfamily is represented by the family Epactozetidae, with the species *Epactozetes setosus* Balogh and Mahunka 1969 and *Truncozetes mucronatus* Balogh and Mahunka 1969 recorded from the Amazon (Franklin et al. 2006).

## Ameroidea Bulanova-Zachvatkina 1957

Of the 14 families known in this superfamily, Damaeolidae Grandjean, 1965, Eremobelbidae Balogh, 1961, Eremulidae Grandjean, 1965, Heterobelbidae Balogh, 1961, Rhynchoribatidae Balogh, 1961, and Staurobatidae Grandjean, 1966 have been reported from Brazil, comprising 25 species. Among them, six are believed to be endemic, including *Suctoribates crassisetosus* Franklin & Woas, 1992, described from specimens collected in the Amazon (Franklin and Woas 1992). Many Ameroidea species have been recorded by de Moraes et al. (2011), Ferreira et al. (2012) and Oliveira et al. (2005).

## Ameronothroidea Vitzthum 1943

This superfamily is composed of five families, Ameronothroidea which also includes Ameronothridae Vitzthum, 1943, Podacaridae Grandjean, 1955, Selenoribatidae Schuster, 1963 and Tegeocranellidae Balogh & Balogh, 1988 (Behan-Pelletier and Lindo 2023). Only members of Selenoribatidae are known to occur in Brazil, represented by *Schusteria littorea* Grandjean, 1968 (Grandjean 1968, Oliveira et al. 2017).

## Caleremaeoidea Grandjean 1965

The monofamilial Caleremaeoidea Grandjean 1965 is part of a paraphyletic cluster of superfamilies in Oribatida requiring revision (Norton and Behan-Pelletier 2020). In Brazil, *Anderemaeus hidasii* Balogh, 1995, *Epieremulus braziliensis* Balogh & Mahunka, 1969 and *Epieremulus longicarinatus* Balogh & Mahunka, 1978 represent this superfamily.

## Carabodoidea Koch 1837

This is a species-rich superfamily requiring revision due to phylogenetic uncertainty which families belong within the superfamily (e.g., Subías 2023, Behan-Pelletier and Lindo 2023, Norton and Behan-Pelletier 2009). In Brazil, members of Carabodidae Koch, 1837, Dampfiellidae Balogh, 1961 and Otocepheidae Balogh, 1961 have been reported, with a total of 24 species.
## Ceratozetoidea Jacot, 1925

This is also a highly widespread and diverse superfamily, estimated to contain 600 species (Subías 2023), although its taxonomy also needs revision. Some species in this superfamily are of economic importance, including *Humerobates rostrolamellatus* Grandjean, 1936 (De Giosa and Barreto 2022) for its association with crops, and others for being intermediate hosts of anoplocephalid tapeworms (Denegri 1993). In Brazil, *Ceratokalummidae* Balogh, 1970, *Ceratozetidae* Jacot, 1925, *Chamobatidae* Grandjean, 1954, *Humerobatidae* Grandjean, 1971, and *Punctoribatidae* Thor, 1937 are represented by 12 species.

## Crotonioidea Thorell, 1876

The taxonomic relationships within Crotonioidea are not entirely resolved (see also Woas 2002), but Behan-Pelletier and Lindo (2023) list in it Crotoniidae Thorell, 1876 (incl. Camisiidae auct.), *Hermanniidae* Sellnick, 1928, *Malaconothridae* Berlese, 1916, *Nanhermanniidae* Sellnick, 1928, *Nothridae* Berlese, 1896, and *Trhypochthoniidae* Willmann, 1931. Members of all of these families have been found in Brazil, for a total of 36 species. It is important to note that *Archegozetes longisetosus* Aoki, 1965 (*Trhypochtoniidae*), is among the best studied oribatids due to its high fecundity and short generation time (Norton and Palmer 1991), which makes it a model species for studies on development (Brückner et al. 2018), ecotoxicology (Seniczak and Seniczak 2002) and morphology (Alberti et al. 2003).

## Cymbaeremaeoidea Sellnick, 1928

This superfamily includes a single family, *Cymbaeremaeidae* Sellnick, 1928 (Norton and Behan-Pelletier 2009). It is represented in Brazil by a single species.

## Eremaeozetoidea Piffl, 1972

The taxonomic relationships within this superfamily are inconclusive, and some authors do not recognize it as a valid taxon (e.g., Subías 2023), while Norton and Behan-Pelletier (2009) included *Eremaeozetidae* Piffl, 1972 and *Idiozetidae* Aoki, 1976 in it, pending on the discovery of idiozetid imatures. *Eremaeozetes arboreus* Nübel-Reidelbach & Woas, 1992, *Eremaeozetes lineatus* Mahunka, 1985, and *Eremaeozetes woelkei* Piffl, 1972 (*Eremaeozetidae*) are found in Brazil.

## Eutegaeoidea Balogh, 1965

The complicated taxonomic relationships in this superfamily have been recently reviewed by Colloff (2023),

who listed 84 species worldwide. However, we still follow Oliveira et al. (2017) in this publication. In Brazil, six species are recorded in the families *Brazilobatidae* Koçak & Kemal, 2008, *Compactozetidae* Luxton, 1988 and *Microtegeidae* Balogh, 1972.

## Galumnoidea Jacot, 1925

Subías (2023) recognized about 700 species of Galumnoidea in the families *Galumnidae* Jacot, 1925 and *Galumnellidae* Piffl, 1970. Mites of this superfamily have as a distinct feature the notogaster with hinged and auriculate pteromorphs with alary furrow (Norton and Behan-Pelletier 2009). In Brazil, *Galumnoidea* is represented by 50 species in both *Galumnidae* and *Galumnellidae*.

## Gustavioidea Oudemans, 1900

Of the nine families containing about 400 species included in *Gustavioidea* by Subías (2023), members of Astegistidae Balogh, 1961, *Liacaridae* Sellnick, 1928 and *Peloppiidae* Balogh, 1943 are represented in Brazil by 25 species, 19 of which belong to *Xenillus* Robineau-Desvoidy, 1839 (*Liacaridae*). Species of this genus as well as of *Ceratoppia* Berlese, 1908 (*Peloppiidae*) and *Liacarus* Michael, 1898 (*Liacaridae*) have been shown to host tapeworms (Denegri 1993).

## Hermannielloidea Grandjean, 1934

*Hermannielloidea* are ornamented mites with distinct paired opisthonotal glands opening on funnel-shaped tubes or large apophyses of their notogaster (Norton and Behan-Pelletier 2009). Subías (2023) listed 85 valid species in this superfamily, which includes the families *Hermanniellidae* Grandjean, 1934 and *Plasmobatidae* Grandjean, 1961, both of which have been reported from Brazil from a total of nine species.

## Licneremaeoidea Grandjean, 1931

This superfamily includes five families containing in total about 175 species (Subías 2023). Of these, 10 species representing the following families have been reported from Brazil: *Charassobatidae* Grandjean, 1958, *Eremellidae* Balogh, 1961, *Licneremaeidae* Balogh, 1943 and *Scutoverticidae* Grandjean, 1954.

## Limnozetoidea Thor, 1937

This superfamily contains two families: *Hydrozetidae* Grandjean, 1954 and *Limnozetidae* Grandjean, 1954 (Oliveira et al. 2017, Behan-Pelletier and Lindo 2023). Both are repre-
sented in Brazil by the following species: *Hydrozetes dimorphus* Hammer, 1962, *Hydrozetes paulista* Pérez-Íñigo & Baggio, 1989, *Hydrozetes uberabensis* Pérez-Íñigo & Baggio, 1996, and *Limnozetes similis* Pérez-Íñigo & Baggio, 1989. The latter three are considered endemic from to the Atlantic Forest.

### Microzetoidea Grandjean, 1936

This superfamily comprises a single family, Microzetiidae Grandjean, 1936, with over 200 species worldwide (Subías 2023). This family is represented in Brazil by 23 species all of which have only been collected from forested areas. Woas (2002) suggested that mites of this family are particularly diverse in Neotropical rain forests.

### Neoliodoidea Sellnick, 1928

These are heavily sclerotized and large mites (1,000–2000 µm) belonging to the single family Neoliodidae Sellnick, 1928 (Norton and Behan-Pelletier 2009). Subías (2023) listed 52 species worldwide, three of which have been reported from Brazil: *Neoliodes terrestris* Wallwork, 1963, *Teleioliodes ghanensis* Wallwork, 1963, and *Teleioliodes zikani* Sellnick, 1930.

### Oppioidea Grandjean, 1951

This is the largest oribatid superfamily (Norton and Behan-Pelletier 2009), consisting of over 1,300 species worldwide (Subías 2023). In Brazil, the families Machadobelbidae Balogh, 1972, Machuellidae Balogh, 1983, Oppiidae Sellnick, 1937, Quadroppiidae Balogh, 1983, Sternoppiidae Balogh & Mahunka, 1969, Teratoppiidae Balogh, 1983 are represented by 70 species. Importantly, the cosmopolitan species *Oppiella nova* (Oudemans, 1902), of the family Oppiidae, is considered to be the most intensively studied oribatid species (Marshall et al 1987, Kaneko 1988), occurring from remote islands such as Svalbard (Seniczak et al 2014) to large urban areas (Barreto et al. 2020).

### Oribatelloidea Jacot, 1925

This superfamily presently comprises only Oribatellidae Jacot, 1925, but in the past was a repository for a mixture of unrelated taxa (Norton and Behan-Pelletier 2009). *Oribatella serrata* Balogh & Mahunka, 1969 and *Oribatella serrula* Pérez-Íñigo & Baggio, 1985 are the only representatives of this group in Brazil.

### Oripodoidea Jacot, 1925

According to different authors, this superfamily comprises between 15 and 18 families (e.g., Norton & Behan-Pel-

letier 2009, Subías 2023). In Brazil, a rich diversity of these mites is found, with 83 species of nine families recorded. This research not only provides a comprehensive understanding of mite diversity but also highlights the importance of *Rosgrozetes ovulum* (Berlese, 1908) being used as a model species for studies of temperature-size relationships (Pequeno et al. 2018), as well as null models (Pequeno and Franklin 2014) and population dynamics (Pequeno et al. 2020). Alongside *R. ovulum*, *Scheloribates curvialatus* Hammer, 1961, also present in Brazil, is known to damage roots of pineapple (Sanyal and Das 1989).

### Plateremaeoidea Trägårdh, 1926

In Brazil, the families Aleurodamaeidae Paschoal & Johnston, 1985, Pheroliodidae Paschoal, 1987 and Plateremaeidae Trägårdh, 1926 are represented by 13 species.

### Tectocepheoidea Grandjean, 1954

This group consists of a single family, Tectocepheidae Grandjean, 1954 (Norton and Behan-Pelletier 2009, Behan-Pelletier and Lindo 2023). Subías (2023) recognized 17 species in this family, but only *Tectocepheus americanus* Pérez-Íñigo & Baggio, 1989, *Tectocepheus velatus* Michael, 1880 and *Tegeozetes tunicatus* Berlese, 1913 have been reported from Brazil. Interestingly, *T. velatus* is a cosmopolitan species present in a variety of ecosystems, but also often found in the plumage of birds (Lebedeva and Krivolutsky 2003) and in disturbed or early successional habitats (Norton and Behan-Pelletier 2009). This species has also been reported to be infected by anoplocephalid cysticercoids (Fritz 1995).

### Trizetoidea Ewing, 1917

This superfamily consists of three families. Suctobelbidae Jacot, 1938 is the only trizetoid family represented in Brazil, by 16 species of *Neosuctobelba* Balogh & Mahunka, 1969, *Parasuctobelba* Hammer, 1977, *Suctobelbella* Jacot, 1937, and *Suctobelbila* Jacot, 1937. Súbias (2023) lists over 300 species for this family worldwide.

### Zetorchestoidea Michael, 1898

This superfamily is recognized by Subías (2023) and Behan-Pelletier and Lindo (2023), although Lienhard et al. (2013) considered it paraphyletic. In Brazil, it is represented by species of Arceremaeidae Balogh, 1972, Eremaeidae Oudemans, 1900 and Zetorchestidae Michael, 1898. *Arceremaeus cubanus* Balogh & Mahunka, 1980, *Tecteremaeus anoporous* Balogh & Mahunka, 1969, *Tecteremaeus cachoeirensis* Franklin
& Woas, 1992, *Tecteremaeus cristatus* Balogh & Mahunka, 1969, *Eremaeus translamellatus* Hammer, 1952, and *Zetorchestes schusteri* Krisper, 1984 have been recorded from this country.

## Infraorder Enarthronota Grandjean, 1969

This infraorder is a highly plastic group in terms of body form (Norton 2001). They range from pale to vibrant yellow, orange or red species, all with different degrees of mineralization (Behan-Pelletier and Lindo 2023). In Brazil, this group is represented by 30 species in eight families.

### Brachychthonioidea Thor, 1934

This superfamily only includes Brachychthoniidae Thor, 1934. These small mites (about 250 µm in length) are considered the most geographically and ecologically widespread family of Oribatida (Norton and Behan-Pelletier 2009), occurring in a multitude of habitats including savanna (Santos et al. 2008), forests (Meehan et al. 2020), woodlands (Ashwood et al. 2022), peatlands (Barreto and Lindo 2021) and other habitats. In Brazil, it is represented by *Liochtho-nius unilateralis* Hammer, 1962 and *Sellnickochthonius oliates* Hammer, 1958.

### Hypochthonioidea Berlese, 1910

This superfamily is composed by Eniochthoniidae Grandjean, 1947, Hypochthoniidae Berlese, 1910, Lohmaniidae Berlese, 1916, Mesoplophoridae Ewing, 1917 and Psammochthoniidae Fuangarworn and Norton, 2013. This superfamily has both dichoid (free articulation of prodosoma and hysterosoma) and ptychoid (mite body can close like a seed when disturbed; box mites) body forms, with different levels of cuticle mineralization (Fuangarworn and Norton 2013). For instance, *Hypochthonius rufulus* Koch, 1835 has calcium phosphate whereas species of Eniochthoniidae and Mesoplophoridae have calcium oxalate in their cuticle (Norton and Behan-Pelletier 1991). Species of *Mesoplophora* Berlese, 1904 are known to be phoretic on beetles and cockroaches (Norton 1980). With the exception of Eniochthoniidae, all families are present in Brazil, where the superfamily is represented by 22 species.

### Protoplophoroidea Ewing, 1917

This is another group in need of taxonomic revision (Norton and Behan-Pelletier 2009). Six species of Cosmochthoniidae Grandjean, 1947, Haplochthoniidae van der Hammen, 1959 and Sphaerochthoniidae Grandjean, 1947 have been reported from Brazil.

## Infraorder Mixonomata Grandjean, 1969

This group is a paraphyletic infraorder comprising both dichoid and ptychoid forms (Behan-Pelletier and Lindo 2023). In Brazil, it is represented by 116 species in the families Epilohmanniidae Oudemans, 1923, Euphthiracaridae Jacot, 1930, Oribotritiidae Balogh, 1943, and Phthiracaridae Perty, 1841, of which only Epilohmanniidae does not consist of box mites.

### Epilohmannioidea Oudemans, 1923

Epilohmanniidae is the single family in the widespread Epilohmannioidea, with 56 species listed worldwide (Subías 2023). *Epilohmannia minuta* Berlese, 1920, *Neoepilohmannia dolosa Pérez-Íñigo* & Baggio, 1985 and *Neoepilohmannia lenkoi* Balogh & Mahunka, 1977 are recorded for Brazil, but Franklin et al. (2013) list other *Epilohmannia* species morphotyped at the genus level.

### Euphthiracaroidea Jacot, 1930

This speciose superfamily comprises the families Euphthiracaridae Jacot, 1930, Oribotritiidae Grandjean, 1954, and Synichotritiidae Walker, 1965, which together have about 400 species worldwide (Subías 2023). In Brazil, Euphthiracaroidea is represented by 36 species of Euphthiracaridae and Oribotritiidae. Niedbała (2004) has improved the knowledge of the fauna of ptyctimous mites for the Neotropical region.

### Phthiracaroidea Perty, 1841

This superfamily of ptyctimous mites consists of almost 1,000 species worldwide, all in the family Phthiracaridae Perty, 1841. In Brazil, 77 species of mites of this group have been recorded. Niedbała (2004) has improved the knowledge of the fauna of ptyctimous mites for the Neotropical region. Immature mites of this superfamily are endophages in spruce needles (Edsberg and Hagvar 1999).

## Infraorder Palaeosomata Grandjean, 1969

This group consists of two superfamilies, Acaronychoidea Grandjean, 1932 and Palaeacaroidea Grandjean, 1932 of pale and lightly sclerotized mites (Behan-Pelletier and Lindo 2023). These mites are represented in Brazil by six species in the two barely studied superfamilies.

### Acaronychoidea Grandjean, 1932

This superfamily comprises a single family, Acaronychidae Grandjean, 1932, and 23 species worldwide (Subías
2023). A single species of this group (*Acaronychus proximus* Schubart, 1968) has been recorded from Brazil; some authors place this species in the superfamily Archeonothroidea Grandjean, 1932 (e.g., Behan-Pelletier and Lindo 2023). Norton and Behan-Pelletier (2009) consider mites of this superfamily to be one of the most plesiomorphic oribatids.

### Ctenacaroidea Grandjean, 1954

Adelphacaridae Grandjean, 1954 and Ctenacaridae Grandjean, 1954 compose the Ctenacaroidea (Behan-Pelletier and Lindo 2023). Both of these families are represented in Brazil by five species: *Aphelacarus acarinus* Berlese, 1910, *Aphelacarus sellnicki* Grandjean, 1952, *Beklemishevia barbata* Schubart, 1968, *Ctenacarus araneola* Grandjean, 1932.

### Infraorder Parhyposomata Grandjean, 1969

Parhyposomata comprises a single superfamily (Parhyposchthonioidea Grandjean, 1932). These mites are weakly sclerotized and white or pale yellow (Behan-Pelletier and Lindo 2023).

### Parhypochthonioidea Grandjean, 1932

Parhypochthonioidea comprises two families, only one of which, Parhypochthoniidae Grandjean, 1932, is present in Brazil. In this family, only one species has been recorded in the country, *Parhypochthonius aphidinus* Berlese, 1904, although its record requires confirmation (Ferreira et al. 2012).

### Order Trombidiformes Reuter, 1909

This order is organized into two suborders, Prostigmata Kramer, 1877 and Sphaerolichida O'Connor, 1984, but the latter has not been reported from Brazil. Prostigmata contains most of the serious agricultural pests (e.g., Eriophyoidea, Tetranychoidea, and Tarsonemoidea), but many species are predominantly predatory, attacking small plant feeding arthropods. Some of these species have the potential or have been demonstrated as effective biological control agents (e.g., Cheyletidae Leach, 1815, Iolinidae Pritchard 1956, and Stigmaeidae Oudemans, 1931). In addition, some are parasites of invertebrates (e.g., Tarsonemidae Canestrini & Fanzago, 1877 and Trombidioidea Leach, 1815) or vertebrates (e.g., Myobiidae Mégnin, 1877 and Trombiculidae), while others are fungivorous (e.g., Pygmephoridae Cross, 1965) or pollinivorous. Many Trombidiformes are aquatic, acting as predators or parasites, mostly of arthropods. In Brazil, 1,469 species of 73 families have been reported.

### Suborder Prostigmata Kramer, 1877

This is a large group consisting of four infraorders: Anystina van der Hammen, 1972, Eleutherengona Oudemans, 1909, Eupodina Krantz, 1978, and Labidostommatina Krantz, 1978. It has a wide variation in morphological characteristics, inhabiting terrestrial and aquatic environments, and having diverse feeding behavior, such as predators, parasites, phytophages, and fungivores, among others (Krantz and Walter 2009).

### Infraorder Anystina van der Hammen, 1972

Anystina is organized in 21 superfamilies, with approximately 11,500 species (Zhang 2011). They are traditionally separated into three groups according to their biology and ecology: Anystae, aquatic Parasitengona and terrestrial Parasitengona. As these groups are not used in TCBF, the superfamilies are listed in alphabetical order within the infraorder.

Anystae, include terrestrial mites thought to be predatory in all life stages, with 255 species. These are divided into five superfamilies Adamystoidea Cunliffe, 1957, Anystoidea Oudemans, 1936, Caeculoidea Berlese, 1883, Paratydeoidea Baker, 1949, and Pomerantzioidea Baker, 1949. Among these, Adamystoidea and Pomerantzioidea have not been reported from Brazil.

In aquatic and terrestrial Parasitengona, the larvae are parasites and the active post-larval stages (deutonymphs and adults) are predators. Larvae of aquatic Parasitengona attach to immature and adult insects that spend at least part of their life cycle in the water, while, the active post-larval stages are predators of aquatic invertebrates. In few species, larvae act as commensals in freshwater sponges and mussels. The "true water-mites" form a monophyletic group, known as Hydrachnidia, comprising seven superfamilies: Arrenuroidea Thor, 1900, Eylaoidea Leach, 1815, Hydrachnoidea Leach, 1815, Hydrovolzioidea Thor, 1905, Hydryphantoidea Piersig, 1896, Hygrobatoidea Koch, 1842, and Lebertioidea Thor, 1900. Although representatives of Stygothrombidioidea Thor, 1935 share an aquatic lifestyle with the water mites, they are more phylogenetically related to the terrestrial Parasitengona (Dabert et al. 2016). Hydrovolzioidea and Stygothrombidioidea have not been reported from Brazil.

Terrestrial Parasitengona are parasites of terrestrial arthropods and tetrapods in the larval stage and predators of small arthropods in the post-larval stages. Additionally, some species, such as *Balaustium*, are pollenophagous and
have free-living larvae, which may represent an evolutionary
reversion from parasitism. They are again divided into two
groups. The first, Erythraeina, is organized in two superfami-
lies, Calyptostomatoidea Oudemans, 1923 and Erythraeoidea
Robineau-Desvoidy, 1828, and only the latter is represented
in Brazil. The second, Trombidiina, consists of seven super-
families, Amphotrombioidea Zhang, 1998, Allotanaupo-
doidea Zhang & Fan, 2007, Chyzerioidea Womersley, 1954,
Tanaupodoidea Thor, 1935, Trombiculoidea Ewing, 1929,
Trombidioidea Leach, 1815, and Yurebilloidea Southcott,
1996. Of these, only Trombiculoidea and Trombidioidea,
have been reported from Brazil.

Anystoidea Oudemans, 1936

These are long-legged and agile predators found on
vegetation, litter or on the soil surface. They are divided
into five families: Anystidae Oudemans, 1936, Chulacaridae
Fuangarworn et al., 2016, Erythracaridae Oudemans, 1936,
Pseudocheylidae Oudemans, 1909, and Teneriffiidae Thor,
1911. The Brazilian Anystoidea are mostly unknown and
are represented by five species distributed in three families,
Anystidae Oudemans, 1902, Pseudocheylidae Oudemans,
1909, and Teneriffiidae Thor, 1911.

Arrenuroidea Thor, 1900

These mites are organized in 19 families worldwide (Smit 2020), of which only four have been recorded from Brazil: Arrenuridae Thor, 1900, Athienemanniidae Viets, 1922, Krendowskiidae Viets, 1926, and Mideopsidae Koenike, 1910.

Arrenuridae is a cosmopolitan family whose members
are found in aquatic water bodies and phytotelmata (Walter
and Proctor 2013, Proctor et al. 2015). Larvae are active and
swim rapidly, searching their hosts beneath the water surface
and parasitizing insects of the orders Diptera, Coleoptera,
and Odonata (Zawal 2006, Proctor et al. 2015). This is the
most diverse family of water mites in Brazil, with 99 valid
species, followed by Krendowskiidae (24 species), Mideop-
sidae (24) and Athienemanniidae (two).

Caeculoidea Berlese, 1883

This superfamily contains a single family, Caeculidae
Berlese, 1883, consisting of seven genera worldwide (Zhang
et al. 2011). These are known as rake-legged mites; they have
been recorded from the Cenomanian aged Burmese amber
(Porta et al. 2019). Only *Andocaeculus* (*Andocaeculus*) *caioi*
Ott & Ott, 2014 and *Neocaeculus setecidades* Ott & Ott, 2018
were reported from Brazil.

Erythraeoidea Robineau-Desvoidy, 1828

Erythraeoidea is organized into two living families, Erythraeidae Robineau-Desvoidy, 1828 and Smarididae Kraemer, 1878, both of which are reported from Brazil. Species of these families can be found on plants, caves, humus and on open ground, walking quickly. The larvae of some species are potential vectors of pathogens, while the larvae of others are biological control agents of harmful insects (Welbourn 1983, DiBlasi et al. 2011, González-Moraga et al. 2015, Jacinavicius et al. 2019). In Brazil, 32 species of Erythraeidae have been recorded attacking different arachnids and insects. However, only four species of Smarididae are represented in Brazil.

Eylaoidea Leach, 1815

Eylaoidea is the superfamily of ancient water mites
comprising the families Eylaidae Leach, 1815, Limnocharidae
Grube, 1859, Piersigiidae Oudemans, 1902, and Aphevideruli-
cidae Gerecke, Smith & Cook, 1999. Only Eylaidae and Lim-
nocharidae, with seven species each, were reported in Brazil.

Hydrachnoidea Leach, 1815

Hydrachnoidea comprises the monotypic family
Hydrachnidae Leach, 1815. Seven species of this family have
been reported from Brazil, all from *Hydrachna* Müller, 1776.
Larvae of species of this genus are known as ectoparasites
of aquatic Hemiptera and Coleoptera (Davids 1973, Proctor
et al. 2015).

Hydryphantoidea Piersig, 1896

Members of Hydryphantoidea are found in all types
of freshwater environments (Smit 2020). This superfamily
comprises seven families, four of which are represented in
South America (Castro LAS et al. 2023). Three of them have
been recorded in Brazil: Hydryphantidae Piersig, 1896 (five
species), Rhynchohydracaridae Lundblad, 1936 (seven spe-
cies) and Hydrodromidae Viets, 1936 (five species).

Hygrobatoidea Koch, 1842

The water mites of this superfamily constitute 12
families, seven of which have been reported from Brazil,
namely Aturidae Thor, 1900, Frontipodopsidae Viets, 1931,
Hygrobatidae Koch, 1842, Pionidae Thor, 1900, Pontarach-
nidae Koenike, 1910, Unionicolidae Oudemans, 1909, and
Wettinidae Cook, 1956. This is the most diverse Anystina
superfamily in Brazil, with 242 valid species. Within it,
Unionicolidae is the best represented family, with 128 spe-
cies, followed by Hygrobatidae with 54 species.
## Lebertioidea Thor, 1900

This water mite superfamily comprises 11 families, of which four have been reported from Brazil: Anisitsiellidae Koenike, 1910 (three species), Limnesiidae Thor, 1900 (55 species), Sperchontidae Thor, 1900 (one species), and Oxidae Viets, 1926 (five species).

## Paratydeoidea Baker, 1949

Paratydeoidea is a small group of edaphic and arboreal mites of worldwide distribution (Khaustov 2017). This superfamily comprises the families Paratydeidae Baker, 1949 and Stigmocheylidae Kethley, 1990. Within this superfamily, only one species of Paratydeidae has been recorded from Brazil, represented only by *Scolotydaeus corticicola* Flechtmann, 1992 (Flechtmann 1992).

## Trombiculoidea Ewing, 1929

Trombiculoidea is organized in seven families, Auedyanidae Southcott, 1987, Johnstonianidae Thor, 1935, Leeuwenhoekiidae Womersley, 1944, Neotrombidiidae Feider, 1959, Trombellidae Leach, 1815, Trombiculidae Ewing, 1929, and Walchiidae Ewing, 1946. Only a few species of Johnstonianidae, Neotrombidiidae, and Trombellidae have been recorded in Brazil, all parasitizing arthropods. Additionally, about 80 species of chiggers of the families Leeuwenhoekiidae and Trombiculidae have been recorded parasitizing vertebrates. Species of these last two families are of great veterinarian and public health importance. Within the first, *Apolonia tigipioensis* Torres & Braga, 1938 causes what has been termed nodular trombiculiasis in humans and birds, in northeastern Brazil (Bassini-Silva et al. 2018). In the second, some species of *Eutrombicula* Ewing, 1938 have been reported causing human trombiculiasis in northern and southeastern Brazil (Bassini-Silva et al. 2019b, 2022).

## Trombidioidea Leach, 1815

This superfamily consists of four families worldwide, namely Achaemenothrombiidae Saboori, Wohltmann & Hakimitabar, 2010, Neothrombiidae Feider, 1959, Microtrombidiidae Thor, 1935, and Trombidiidae Leach, 1815 (Zhang et al. 2011). Although representatives of these families are very common and easily found in soils, the Brazilian Trombidioidea are mostly unknown and are represented by few species formally ascribed to the families Microtrombidiidae and Trombidiidae.

## Infraorder Eleutherengona Oudemans, 1909

Eleutherengona are divided traditionally into two

groups (not in TCBF): Raphignathina and Heterostigmata. The first comprises five superfamilies including predators (mainly Raphignathoidea Kramer, 1877 and Cheyletoidea Leach, 1815), phytophages (Tetranychoidea Donnadieu, 1875), and specialized animal parasites (Myobioidea Mégnin, 1877 and Pterygosomatoidea Oudemans, 1910). In contrast, Heterostigmatina comprises seven superfamilies, including some that are predominantly fungivorous (Dolichocyboidea Mahunka, 1970, Heterocheyloidea Trägårdh, 1950, Pygmephoroidea, Tarsocheyloidea Atyeo & Baker, 1964, and Trochometridioidea), arthropod parasites (Podapolipidae Ewing, 1922 and Pyemotoidea Oudemans, 1937), turtle parasites (Cloacaroidea Camin, Moss, Oliver & Singer, 1967) and of variable feeding habits (Tarsonemoidea Kramer, 1877). Tarsocheyloidea and Cloacaroidea have not been reported from Brazil.

## Cheyletoidea Leach, 1815

This superfamily comprises the families Cheyletidae, Demodicidae Nicolet, 1855, Harpirhynchidae Dubinin, 1957, Psorergatidae Dubinin, 1955, and Syringophilidae Lavoipierre, 1953 (Bochkov 2002, Walter et al. 2009). Cheyletidae contains species that are ectoparasites or endoparasites of vertebrates (mammals, birds or reptiles), or predators of arthropods, including other mites. Only Psorergatidae has not been reported from Brazil.

Most of the 19 cheyletids species reported from Brazil were collected from house dust (Rosa and Flechtmann 1979a, Galvão and Guitton 1986, Binotti et al. 2001), plants (Feres and Flechtmann 1995, Buosi et al. 2006) and parasitically associated with domestic rabbits and dogs (Londero et al. 1964, Alvim et al. 2005, Barros-Battesti et al. 2020); few species live on bird skin (Fain 1972, Rosa and Flechtmann 1979b, Fain and Bochkov 2002) or are phoretic on louse flies (Valim and Gazêta 2007).

A recent checklist of demodicids reported three *Demodex* species (*D. bovis* Stiles, 1892, *D. cati* Mégnin, 1877 and *D. injai* Desch & Hillier, 2003) from Brazil (Izdebska and Rolbiecki 2020), although in a more recent publication, Beron (2021) listed only two species (*D. bovis* and *D. phylloides* Csokor, 1879). In an effort to review the literature, Valim (2023) confirmed the existence of eleven species of *Demodex* in Brazil, all considered important agents of folliculitis (demodicosis) in domestic animals (Gazi et al. 2019).

Mites of the family Harpirhynchidae are highly specialized permanent monoxenic or narrowly oligoxenic parasites (Bochkov et al. 1999). This family includes permanent skin parasites inhabiting the feather follicles and epidermis
of birds (Harpirhynchinae) and the area under snake scales (Ophioptinae). In Brazil, these mites are represented by five species in each subfamily. Beron (2021) reported nine species from the country, overlooking the record of *Ophioptes tropicalis* Ewing, 1933 by Fain (1964).

Syringophilidae consists of mites mostly adapted as endoparasites of quill feathers, and the 27 reported species are an underestimation of the diversity of this family in Brazil, considering the highly diverse Brazilian bird fauna (Pacheco et al. 2021, Fain et al. 2000, Bochkov and Fain 2003b, Skoracki et al. 2016, Marciniak-Musial and Sikora 2022).

## Dolichocyboidea Mahunka, 1970

This heterostigmatid superfamily consists of two families Dolichocybidae Mahunka, 1970 and Crotalomorphidae Lindquist & Krantz, 2002. Only Dolichocybidae has been reported from Brazil. This is a small group of poorly studied early-derivative heterostigmatid mites. They are possibly fungivorous (Khaustov and Viacheslav 2018). Only the genera *Pavania* Lombardini, 1949 and *Formicomotes* Sevastianov, 1980 have been recorded from Brazil, associated with scarab beetles and termites, respectively (Mahunka 1970, Khaustov and Frolov 2017, 2018).

## Heterocheyloidea Trägårdh, 1950

This superfamily comprises the single family Heterocheylidae Trägårdh, 1950 and one genus, *Heterocheylus* Lombardini, 1926. All known species are subelytral parasites of passalid beetles. They occur with their hosts in decaying wood of tropical and warm temperate forests throughout the world (Lindquist and Kethley 1975). In Brazil, only *Heterocheylus fusiformis* Lombardini, 1938 have been recorded (Lombardini 1938).

## Myobioidea Mégnin, 1877

Myobioidea comprises only the family Myobiidae, which is distributed worldwide. It is a relatively large family, with more than 600 species (Bochkov and Fain 2003a, Bochkov 2009). Species of this family are characterized by the strongly modified first pair of legs, bearing clasping structures used for attachment to the hairs of the hosts when feeding on lymph and cell content. *Myobia musculi* (Schrank, 1781) is an ectoparasite of worldwide importance as agent of pruriginous dermatitis in laboratory rodents. Mites of this family are known as fur mites, having parasitic specificity defined at different taxonomic levels of the mammal host; they have monoxenous or oligoxenous parasitic habits (Dus-

bábek 2002). They are divided into five subfamilies, of which Archemyobiinae Fain, 1973 and Myobiinae Ewing, 1938 have been reported from Brazil (Jacinavicius and Bassini-Silva 2021), the first as ectoparasites of opossums and the second, of bats and rodents. The number of species so far reported in Brazil fewer than expected, as more than 500 species of susceptible mammals are known to occur in this country.

## Pterygosomatoidea Oudemans, 1910

This superfamily includes only Pterygosomatidae Oudemans, 1910. Almost all species of this family are external permanent parasites of lizards and are highly specific to their host (Fajfer 2012). Despite the high diversity of lizards in Brazil, only two species of pterygosomatids have been reported from this country; one native, *Geckobiella harrisi* Davidson, 1985, and one introduced, *G. hemidactyl* Lawrence, 1936, parasitic on introduced geckos. Some records need to be reviewed as they do not provide taxonomic information to confirm the identifications.

## Pyemotoidea Oudemans, 1937

According to Krantz and Walter (2009) and Zhang et al. (2011), it is presently considered to include four families. The largest family is Acarophenacidae Cross, 1965 (31 species) and Pyemotidae Oudemans, 1937 (24 species) and the smallest are Caraboacaridae Mahunka, 1970 (seven species) and Resinacaridae Mahunka, 1976 (two species). Where known, adult females of these mites are parasitoids of insects of different orders (Krantz and Walter 2009). Only species of Acarophenacidae and Pyemotidae have been reported from Brazil, by three and one species, respectively.

## Pygmephoroidea Cross, 1965

Krantz and Walter (2009) considered Pygmephoroidea to be composed of two families, Pygmephoridae and Siteroptidae Mahunka, 1970, they do not consider Neopygmephoridae Cross, 1965 to be a separate family. Zhang et al. (2011) considered Siteroptidae to be a synonym of the Pygmephoridae (based on Khaustov 2008) and included Neopygmephoridae as a valid family (based on Khaustov 2004). While Krantz and Walter (2009) considered Microdispidae Cross, 1965 and Scutacaridae Oudemans, 1916 to constitute the superfamily Scutacaroidea, we are following Zhang et al. (2011) and including these two families in Pygmephoroidea.

Most species of Pygmephoroidea appear to be fungivorous; some are known to have phoretic associations with insects (mainly), and occasionally also with Chilopoda, Dip-
lopoda or groups of Arachnida. A few microdispids and pyg-
mephorids have been cited as pests of cultivated mushrooms.
Most pygmephoroids are soil inhabitants, especially in places
where the substrate favors fungal development; some are
found in the nests of insects, birds and small mammals,
while others are common on plants. Physogastry (wherein
the abdomen becomes greatly enlarged and membranous)
is known to occur in pygmephorids, but is not common in
microdipids or scutacarids (although some small degree of
physogastry has been reported in microdispids).

Representatives of the four pygmephoroid families are
reported from Brazil: 11 of Microdispidae, 15 of Neopygme-
phoridae, 11 of Pygmephoridae, and 36 of Scutacaridae. Two
species of the first family *Pediculaster flechtmanni* (Wicht
Jr., 1970) and *Siteroptes reniformes* Krantz, 1957 have been
reported respectively on cultivation substrates or on plants
in Brazil, the first as a pest of mushrooms and the second
feeding on the fungus *Nigrospora* Zimmerman on orchids.
The pygmephoroids from Brazil were mostly studied by
Mahunka (1969, 1970, and 1977).

Raphignathoidea Kramer, 1877

This superfamily is diverse and widely distributed,
consisting of about 1,112 species organized in 12 families.
Barbutiidae Robaux, 1975, Caligonellidae Grandjean, 1944,
Camerobiidae Southcott, 1957, Cryptognathidae Oudemans,
1902, Dasythyreidae Walter & Gerson, 1998, Dytiscacaridae
Hajiqanbar & Lindquist, 2018, Eupalopsellidae Willmann,
1952, Homocaligidae Wood, 1970, Mecognathidae Gerson
& Walter, 1998, Raphignathidae Kramer, 1877, Stigmaei-
dae, and Xenocaligonellididae Gonzalez, 1978 (Zhang et
al. 2011). Stigmaeidae is the largest family, accounting for
about half of the genera and the majority of the species
(Fan et al. 2016, 2019, Beron, 2020b). In Brazil, only six fami-
lies have been reported: Caligonellidae (two genera, three
species), Cryptognathidae (one genus with two species),
Eupalopsellidae (one species), Camerobiidae (one species),
Stigmaeidae (seven genera, 33 species) and Raphignathidae
(only to genus level).

Mites of this superfamily are mostly cited as predators
of small arthropods in habitats such as soil, moss, lichen,
leaf litter, plants, tree bark, mushrooms, stored food and
bird nests. Some cryptognathids have also been reported
as microphytophages (Swift 1996, Swift and Goff 2001),
being speculated to feed on fungal spores, moss, algae and
even pierce plant cells and drain the contents (Luxton 1973,
1993). Some raphignathids have been reported from pigeon
nests and intestine of a wedded seal, plants, house dust and

even in the urine of humans (Atyeo et al. 1961, Atyeo 1963,
Chaudhri et al. 1979, Rosa and Flechtmann 1979a, Meyer and
Ueckermann 1989, Outhouse and Castro 1990).

Many members of Stigmaeidae are considered eco-
nomically important predators of plant feeding mites and
small insects (Santos and Laing 1985, Gerson et al. 2003, Fan
and Zhang 2005, Fan and Flechtmann 2015, Bizarro et al.
2020b, 2022). A world catalog of the mites of this family was
published by Fan et al. (2016). Over 35% of the stigmaeid
species are free-living predators on plants, while approxi-
mately 27% of the species are edaphic, feeding on moss or
lichen (Gerson 1972).

Tarsonemoidea Canestrini & Fanzago, 1877

Tarsonemoidea is composed of two families (Lindquist
1986): Podapolipidae and Tarsonemidae, both reported from
Brazil. Tarsonemids are mostly known from plants, while
mites of the two families are commonly found in association
with insects. The tarsonemids constitute one of the most im-
portant mite groups of agricultural pests (Gerson et al. 2003),
especially *Phytonemus pallidus* (Banks, 1899) and *Polyphagotar-
sonemus latus* (Banks, 1904) (Lindquist 1986). Podapolipids
have been found externally and internally on several insect
orders, including Blattaria, Orthoptera, Heteroptera, Hyme-
noptera and, mainly, Coleoptera (Walter et al. 2009).

The family Podapolipidae includes around 32 genera and more than 306 species (Seeman and Walter 2023, Seeman and Katlav 2024). In Brazil, nine genera are registered: *Bombacarus* Stammer, 1951, *Buprestapolipus* Husband, Ochoa & Naskrecki, 1995, *Chrysomelobia* Regenfuss, 1968, *Cydnipolipus* Husband & Husband, 2017, *Eutarsopolipus* Berlese, 1913, *Kurosapolipus* Husband & Li, 1993, *Podapolipus* Rovelli & Grassi, 1888, *Rhynchopolipus* Husband & Flechtmann, 1972, and *Tetrapolipus* Feldman-Muhsam & Mumcuoglu, 1981 (Husband and Flechtmann 1972, Feldman-Muhsam and Mumcuoglu 1981, Fain 1987, Husband 1987, Husband and Li 1993, Husband and Eidelberg 1996, Husband and Husband 2017).

Tarsonemids comprises around 45 genera and 624
species (Seeman and Walter 2023). Adults of most species
are only about 200 µm long. These are divided into three
subfamilies, namely Acarapinae Schaarschmidt, 1959,
Pseudotarsonemoidinae Lindquist, 1986 and Tarsoneminae
Canestrini & Fanzago, 1877. Twenty tarsonemid genera are
reported from Brazil, seven of which have been described
based on Brazilian specimens: *Amcortarsonemus* Fain,
1987, *Biscutulumemus* Lofego & Feres 2006, *Crossacarapis*
Ochoa & O'Connor, 1996, *Flechtmannus* Moraes, Lindquist
& Lofego, 2002, *Kaliszewskia* Lofego, Demite & Moraes, 2015, *Ochoanemus* Lofego, Pitton & Rezende, 2016, and *Pseudotarsonemus* Lindquist, 1986 (Lin and Zhang 2002, Lofego and Feres 2006, Lofego et al. 2015, 2016). A catalog of the world tarsonemid species was published by Lin and Zhang (2002).

## Tetranychoidea Donnadieu, 1875

This superfamily includes approximately 2,500 described species of plant-feeding mites in five families: Tetranychidae, Tenuipalpidae Berlese, 1913, Tuckerellidae Baker & Tuttle, 1953, Linotetranidae Baker & Pritchard, 1953, and Allochaetophoridae Reck, 1959 (Lindquist 1985, Walter et al. 2009). The majority of species of this superfamily belong to the families Tetranychidae and Tenuipalpidae, while Allochaetophoridae, Linotetranidae, and Tuckerellidae contain relatively few species. Tetranychidae and Tenuipalpidae include species that cause great economic losses to many crops worldwide (Jeppson et al. 1975, Bolland et al. 1998, Moraes and Flechtmann 2008). Allochaetophoridae has not been reported from Brazil (Meyer and Ueckermann 1997, Walter et al. 2009). Characteristics shared by the tetranychoids are the reduction of the fixed digit in parallel with the fusion of the cheliceral bases, forming a stylophore, which bears the movable cheliceral digits modified into whip-like structures able to pierce and withdraw plant cell contents for nourishment (Lindquist 1985, Meyer and Ueckermann 1997, Walter et al. 2009).

Tetranychidae includes around 1,300 described species in 71 genera, six tribes and two subfamilies, Bryobinae and Tetranychinae (Bolland et al. 1998, Migeon and Dorkeld 2023). Members of the Tetranychinae are the only tetranychoids able to produce web, hence the common name spider mites for the whole family (Lindquist 1985). The largest genus of the Tetranychidae are *Oligonychus* Berlese, 1876 (211 species), *Eotetranychus* Oudemans, 1931 (200), *Tetranychus* Dufor, 1832 (153) and *Bryobia* Koch, 1835 (147). A taxonomic data base is available on the internet for the world tetranychid mites (Migeon and Dorkeld 2023). In Brazil, 126 species of 22 genera are reported, with *Oligonychus*, *Tetranychus*, and *Eotetranychus* being the most diverse, with 31, 26, and 12 species, respectively. A similar database dealing specifically with the species reported from Brazil species is maintained by Flechtmann and Moraes (2017).

Tenuipalpidae includes about 1,100 described species distributed in 41 genera (Mesa et al. 2009, Castro et al. 2020, 2023, Beard et al. 2022). They have a worldwide distribution but are especially diverse in subtropical and tropical regions (Mesa et al. 2009, Walter et al. 2009). Mites of this family are

distinguishable from other tetranychoids by having simple palps, without the structure known as the “thumb-claw” complex, and with a variable number of segments. Many species have the body flattened dorsoventrally, and for this reason they are known as flat mites. The genera with the largest number of valid species are *Tenuipalpus* Donnadieu, 1875 (329), *Brevipalpus* Donnadieu, 1875 (292), *Aegyptobia* Sayed, 1950 (104) and *Cenopalpus* Pritchard and Baker, 1958 (70). These four genera alone represent about 80% of the described tenuipalpid species worldwide (Castro et al. 2020, 2023). In Brazil, 41 species of seven genera have been reported, with *Tenuipalpus* (26 species) and *Brevipalpus* (10 species) the most speciose genera. A database of mites of this family is maintained by Castro et al. (2023).

Tuckerellidae includes about 30 species in a single genus, *Tuckerella* Womersley, 1940 (Beard et al. 2013, Khadem and Asadi 2018). Most of these appear to have restricted distributions, contrasting in this regard with *Tuckerella ornata* (Tucker, 1926), *T. pavoniformis* (Ewing, 1922), *T. knorri* Baker & Tuttle, 1975 and *T. japonica* Ehara, 1975, of relatively wide distribution (Walter et al. 2009, Beard et al. 2013). Only three tuckerellid species have been reported from Brazil: *T. ornata*, *T. pavoniformis*, and *T. knorri* (Flechtmann 1979, Barbosa et al. 2003, Mineiro et al. 2005, Brito et al. 2023). Recently, *T. ornata* was reported causing damage on cocoa fruits (*Theobroma cacao* L.) in the Amazon Region, Pará State, Brazil (Brito et al. 2023).

Linotetranidae includes only 18 species in four genera, namely: *Afrolinotus* Meyer & Ueckermann, 1997, *Anoplopalpus* Meyer & Ueckermann, 1997, *Austrolinus* Beard & Walter, 2004 and *Linotetranus* Berlese, 1910 (Beard and Walter 2004, Walter et al. 2009, Tassi et al. 2020). *Linotetranus* is the largest genus, with 14 described species worldwide (Walter et al. 2009, Tassi et al. 2020). Linotetranidae differs from the other Tetranychoidea by the presence of ventral setae 2a and the absence of ocelli (Baker and Pritchard 1953, Beard and Walter 2004). They have been reported from moss, soil and rotting trees, but most collections indicated an association with crown and roots of grasses or sedges in dryland habitats (Beard and Walter 2004). Recently, Tassi et al. (2020) described a new species from Brazil, *Linotetranus faemensis* Tassi & Duarte, 2020 based on specimens collected in soil from a native pasture in Rio Grande do Sul, which represents the first record of the family for the country.

## Trochometridioidea Mahunka, 1970

Trochometridioidea comprises two fungivorous families *Trochometridiidae* Mahunka, 1970 and *Athyreacaridae*.
Lindquist, Kaliszewski & Rack, 1990. Of these, only *Trochometridiidae* has been reported from Brazil. This family is phoretically associated with Coleoptera, Dermaptera, Diptera and Hymenoptera (Loghmani et al. 2014). In Brazil, only a non-endemic species, *Trochometridium tribulatum* Cross, 1965, associated with the velvet ant (*Hymenoptera: Mutillidae*) has been recorded (Jacinavicius et al. 2018).

## Infraorder Eupodina Krantz, 1978

Eupodina is a rather morphologically and ecologically diverse mite group, constituted by five superfamilies: Bdelloidea Dugès, 1834, EriophyoideaALEPA, 1898, Eupodoidea Koch, 1842, Halacaroidea Murray, 1877, and Tydeoidea Kramer, 1877. Behaviorally they can be predators, parasites, fungivores and phytophagous. The superfamily Halacaroidea comprise mites of mostly marine and estuarine habitats, while mites of other superfamilies are terrestrial. All superfamilies are represented in Brazil.

## Bdelloidea Dugès, 1834

Members of this superfamily are organized in two families of strictly predatory mites, found mainly in the soil and on plants: Bdellidae, with 278 species (Hernandes et al. 2016) and Cunaxidae Thor, 1902, with more than 400 species (Skvarla et al. 2014) worldwide. These families are presently represented in Brazil by nine and 42 species, respectively.

## EriophyoideaALEPA, 1898

Eriophyoid mites usually have an intimate relationship with their host plants and, thus, high host specificity and great diversity (Lindquist et al. 1996, Oldfield 2002, 2005, Skoracka et al. 2010, 2012). They surpass all other groups of phytophagous mites in the extent of their morphological and biological specialization for obligate phytophagy and specificity to their hosts (Lindquist et al. 1996). Morphologically, eriophyoids are distinguishable from other mites by being vermiform, four-legged organisms (hence, the vernacular name, 'four-legged mites') and are among the smallest known arthropods, less than 0.25 millimeters long (Lindquist et al. 1996, Klimov et al. 2018).

Eriophyoidea constitutes an exclusively phytophagous lineage representing one of the largest chelicerate radiations, with fossils known from the Triassic (Schmidt et al. 2012). Although previously considered a sister group to a lineage including Tydeidae Kramer, 1877, Ereynetidae Oudemans, 1931, and Eupodidae, recent molecular phylogenetic evidence, supported by the rDNA and CO1 partitions, indicate

eriophyoids as the sister group of the deep soil-dwelling, vermiform family Nematalycidae in the suborder Endosiphigmata (Klimov et al. 2018, Bolton et al. 2017, 2023).

The host plants include angiosperms (flowering plants) and gymnosperms (e.g., conifers). Eriophyoid mites can be found on leaf buds, stems, bulbs and fruits. Most eriophyoid species (ca. 80–95%) are highly host-specific, colonizing only a single host plant species or a few hosts within a single genus (Oldfield 1996, Skoracka et al. 2010).

Some eriophyoid species can seriously damage their host plants, assuming pest status in crops of social or economic importance (Ueckermann 2010, Hoy 2011). In number of species, EriophyidaeALEPA, 1898 is the main family of phytophagous mites that can transmit plant viruses; de Lillo et al. (2021) listed 23 species confirmed as plant virus vectors and 37 different viruses.

The Eriophyoidea consist of three families: Diptilomiopidae Keifer, 1944, Eriophyidae, and Phytoptidae Murray, 1877. Worldwide, Eriophyidae is the most diverse family, comprising around 3,880 species of 330 genera, divided into 11 tribes (Colopodacini, Nothopodini, Cecidophyini, Colomerini, Aceriini, Eriophyimi, Acaricalini, Calacarini, Tegonotini, Phyllocoptini, and Anthocoptini) in turn distributed into six subfamilies (Nothopodinae, Cecidophyinae, Eriophyinae, Phyllocoptinae, Aberoptinae and Ashieldophyinae) (J.W. Amrine, pers. comm.). The Diptilomiopidae consists of approximately 513 valid species distributed into 78 genera; these are divided into two subfamilies: Diptilomiopinae and Rhycaphytoptinae (Amrine et al. 2003). The Phytoptidae consists of 193 valid species in 28 genera; these are divided into five subfamilies: Phytoptinae, Sierraphytoptinae, Nalepellinae, Novphytoptinae and Prothricinae; and five tribes: Sierraphytoptini, Mackielini, Trisetacini, Nalepellini and Pentasetacini.

In Brazil, 198 species of 77 genera, 11 tribes and five subfamilies of Eriophyidae, 25 species of nine genera and two subfamilies of Diptilomiopidae, and 18 species of eight genera, four tribes and four subfamilies of Phytoptidae have been reported (Navia et al. 2021).

## Eupodoidea Koch, 1842

This superfamily comprises nine families of soil mites, Cocceupodidae Jesionowska, 2010, Eupodidae, Dendrochaetidae Olivier, 2008, Eriorhynchidae Qin & Halliday, 1997, Pentapalpidae Olivier & Theron, 2000, Penthaleidae Oudemans, 1931, Penthalodidae Thor, 1933, Rhagidiidae Oudemans, 1922, and Strandtmanniidae Zacharda, 1979. Most of these are mycophagous, but some are predacious
(e.g., Rhagidiidae) or phytophagous (e.g., Penthaleidae and Penthalodidae) (Laniecki and Magowski 2023). Some species of Penthaleidae are serious pests. The Brazilian Eupodoidea are mostly unknown and represented by only one species of each of the following families: Eupodidae, Penthaleidae and Rhagidiidae. The penthaleid species is *Penthaleus major* (Dugès, 1834), considered a major pest of different crops in several countries and only recently reported for the first time in Brazil.

## Halacaroidea Murray, 1877

The Halacaroidea are a cosmopolitan meiobenthic group with the majority of marine and a few freshwater mite species that are organized in two families, Halacariidae Murray, 1877 and Pezidae Harvey, 1990. The Brazilian Halacaroidea are represented by 16 species of Halacaridae.

## Tydeoidea Kramer, 1877

André and Fain (2000) reorganized Tydeoidea to comprise four families: Ereynetidae, Iolinidae, Triophtydeidae André, 1979 (= Edbakerellidae) and Tydeidae. Many species are free-living or parasitic, but most species are poorly known in relation to their feeding habits, life cycle, and distribution (Kazmierski 1998, André and Fain 2000, Silva et al. 2016a). These mites are fungivores, phytophages, predators, scavengers, or even endoparasites (e.g., Ereynetidae) (Gerson et al. 2003, Sadeghi et al. 2012, Bernardon et al. 2015). The Triophtydeidae and Tydeidae, and to some extent Iolinidae and Ereynetidae, are most often reported from plants (Zacarias and Moraes 2002, Spongoski et al. 2005, Hernandes and Feres 2006, Silva et al. 2014, Tempfli et al. 2015), but are also found in caves, mosses, lichens, stingless bees, stored products and in the soil (Walter et al. 2009, Walter and Proctor 2013, Silva et al. 2016a, 2016b, Da-Costa et al. 2019, Bernardi et al. 2022). Members of the Tydeidae are the most widely studied. This is the largest family of the superfamily, comprising about 332 species (Krantz and Walter 2009, Silva et al. 2016a, Nuvoloni et al. 2020, Bizarro et al. 2020a), with 26 species in Brazil.

Iolinidae comprises about 135 species of three sub-families: Iolininae Pritchard, Pronematinae André, 1980 and Tydaeolinae André, 1908 and this group is the most likely group in Tydeoidea to form relationships with insects (Rosa and Flechtmann 1983, Rosa et al. 1985, André and Fain 2000, Zhang et al. 2011, Theron et al. 2012, Ahmad-Hosseini et al. 2017, Silva et al. 2017, Da-Costa et al. 2020). Their bioecology is still poorly known (Sadeghi et al. 2012, Silva et al. 2017). However, a species of this family has recently

been considered promising as a biological control agent of a pest eriophyid (Duarte et al. 2021, Pijnakker et al. 2021). In Brazil, only ten species have been recorded.

Ereynetidae comprises around 180 species of four sub-families. Speleognathinae Fain, 1957 is the most commonly reported subfamily in Brazil as bird intranasal parasites (Zhang et al. 2011, André and N'Dri 2013). Triophtydeidae is presently known only from a single report in Brazil.

## Infraorder Labidostommatina Krantz, 1978

Mites of this infraorder comprise a single superfamily (Labidostommatoidea Oudemans, 1904) containing a single family, Labidostommatidae Oudemans, 1904. This is one of the few well sclerotised prostigmatid mites; prodorsum with two pairs of trichobothria; usually with a pair of anterolateral ocelli and an unpaired anteromedian ocellus; coxal fields expanded and contiguous mesally. They are predators and have a worldwide distribution (Bertrand 1990). Six species of this group, one in the genus *Labidostomma* Kramer, 1879 and five *Sellnickiella* Feider & Vasiliu, 1969 species have been reported from Brazil.

## Suborder Sphaerolichida O'Connor, 1984

In contrast to the Prostigmata, Sphaerolichida, the only other Trombidiformes order, is a small suborder consisting of two superfamilies, Lordalychoidea Grandjean, 1939 and Sphaerolichoidea Berlese, 1913, each containing a single family, Lordalychidae Grandjean, 1939 and Sphaerolichidae Berlese, 1913. These mites are characterized by having chemicera usually chelate and denticulate; stigma and tracheal system absent; prodorsum with three or six pairs of setae, including two pairs of filamentous trichobothria. Representatives of both superfamilies were collected in Brazil by Antony (2004) and Bernardi et al. (2022). However, in both cases, the mites were not identified to the species level.

## DISCUSSION

Earth's biodiversity is among the most fundamental topics in science and requires collective action to document, understand, and conserve. Mora et al. (2011) estimated that approximately 8.7 million eukaryotic species exist worldwide, based on the 1.2 million species already described, suggesting that nearly 86 % of terrestrial species and 91 % of marine species remain to be described. The global taxonomic workforce is unevenly distributed: about one-third of taxonomists focus on vertebrates, which represent only ~1 % of all species; another third study plants (~10 % of species);
and the remaining taxonomists address invertebrates, which comprise roughly 90 % of species worldwide (May 2010).

The data presented in this document show the current state of knowledge of the mite fauna reported from Brazil in the published literature. We record 3,678 species belonging to six mite orders within the two superorders Acariformes and Parasitiformes. This makes mites the most diverse arachnid group reported from Brazil, with the following numbers of species per order: 2 (Holothyrida), 77 (Ixodida), 990 (Mesostigmata), 18 (Opilioacarida), 1,122 (Sarcoptiformes), and 1,469 (Trombidiformes). Despite our efforts, these numbers likely underestimate the actual published diversity, probably due especially to missing records in older literature. Considering the estimated numbers of species in different mite groups reported by Beaulieu et al. (2011), Schatz et al. (2011), Walter et al. (2011), Zhang (2011) and Zhang et al. (2011), the total number of species determined so far from Brazil represents only about 7 % of the estimated global total. The true diversity is expected to be much greater, as evidenced by recent surveys in which undescribed species from several families (e.g., Chyzeriidae, Lordalychidae, and Neothrombiidae) have been collected and remain to be described (Antony 2004, Bernardi et al. 2022).

A preliminary analysis of the data indicates that a few species of public health importance, listed by Linnaeus (1758) in Systema Naturae, such as the respiratory allergy mites – *Acarus siro* Linnaeus, 1758 (Acaridae); scabies – *Sarcoptes scabiei* (cited as *Acarus scabiei*) (Sarcoptidae); anthropophilic ectoparasites that cause trombiculiasis – *Eutrombicula batatas* (Linnaeus, 1758) (cited as *Acarus batatas*) (Trombiculidae), as well as one of agricultural pest – *Tetranychus urticae* (Linnaeus, 1758) (cited as *Acarus telarius*) (Tetranychidae), were among the first to be confirmed to be part of the Brazilian mite fauna. It can also be observed that, as one would expect, the bulk of the early efforts to determine the Brazilian oribatid mites' fauna were done by European researchers, especially A. Berlese, M. Sellnick, R. Schuster, K. Märkel, J. Balogh, P. Balogh, S. Mahunka, W. Niedala, C. Pérez-Iñigo, L. Beck, and S. Woas. H. Schubart was the first oribatid mite Brazilian specialist working in Brazil.

Erwin (1982) extrapolated data on the richness of Coleoptera and other insects from tropical forests based on host-specificity and spatial ratio. According to Mora et al. (2011), this methodology is adequate for other organisms. Applying these methods to the Brazilian mite fauna is highly informative; for instance, by estimating the number of unknown species in Brazil for mite groups with high host specificity. Myobioidea, a group of mammal parasites. Based

on the fact that only 16 species of these mites have been recorded from Brazil and knowing of the existence of 556 species of potentially susceptible hosts (sum of the Chiroptera, Didelphimorphia and Rodentia recorded in Brazil), it can be estimated that only about 2.9% of the Myobioidea species have been reported. This proportion could be even lower, considering that more than one parasitic species can be found on each host species. Similar calculations to estimate the proportion of known species of feather mites in Brazil, taking into account the reported number of species of each species to date (228 and 104 species, respectively) and the presently known number of susceptible host species in the same country (according to Pacheco et al. 2021, there are 1,971 bird species) suggest that only 11.6% of the Analgoidea and 5.3% of Pterolichoidea present in the country have been identified. Likewise, for plant mites with host-specificity, for example the Eriophyoidea, that attack angiosperms, gymnosperms and ferns, we can calculate the same ratio. These plant groups are represented in Brazil by 2,665 species (BFG 2015, Prado et al. 2015), while the number of Eriophyoidea recorded from this country is 265 species. Hence, the reported mites of this superfamily correspond to only 9.9% of the number that could be present in Brazil. Based on the average of these percentages and extrapolating this information to the other groups, considering the 3,678 species and a knowledge rate of 7.24%, we can roughly estimate the Brazilian acarofauna at about 50,000 species. These findings highlight the urgent need for further research in this area.

Brazil is the fifth largest country on Earth, yet mites remain widely unexplored across much of its territory. This initial survey highlights the urgent need for targeted exploration of habitats and hosts, as well as the incorporation of molecular data to validate and discover species. With the possible exception of Ixodoidea, whose diversity is relatively well known, information gaps persist across most mite taxa in the country. For many groups, specialists are unavailable to compile or update species lists necessary for the preparation and maintenance of the TCBF. In these cases, available records were added and validated by the coordinators—who are not specialists in those groups—until expert evaluations can be conducted.

Historically, this database is highly informative. It reveals the main contributors to mite diversity studies in Brazil, the geographic and taxonomic scope of research efforts, and temporal trends in faunal surveys. In practical terms, the results support updates to quarantine species lists, the development of teaching curricula, and initiatives for faunal conservation.
From a scientific perspective, the objective of the Taxonomic Catalog of the Brazilian Fauna extends well beyond the mere listing of records. It is designed as a comprehensive data source to advance knowledge of Brazil's mite fauna by documenting species distributions, assessing levels of endemism, identifying typical habitats, and recording associated hosts and prey. The effective use of such information will require substantial additional efforts and the active participation of researchers, students, and citizen scientists. Achieving these goals will require the continuous involvement of new personnel engaged in scientific work at Brazilian institutions or at collaborating institutions abroad. This initiative also has the potential to generate numerous undergraduate and graduate research projects and to foster coordinated, integrated research efforts. The authors welcome information on overlooked literature to ensure that this ongoing endeavor remains as comprehensive and productive as possible—an essential contribution in view of the urgent need for taxonomic revisions and the description of new species.

## ACKNOWLEDGEMENTS

The Smithsonian Natural History Museum and National Agricultural Library (NAL-USDA), SEL-USDA for support and assistance with specimens and references. Mention of trade names or commercial products in this publication is solely for purpose of providing specific information and does not imply recommendation or endorsement by the USDA; USDA is an equal opportunity provider and employer.

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