Montes-Ortiz, Lucia ¹ ; Cohuo, Sergio ² and Elías-Gutiérrez, Manuel ³

¹✉ Tecnológico Nacional de México–I. T. Chetumal., Av. Insurgentes 330, Quintana Roo 77013, Mexico.
²Tecnológico Nacional de México–I. T. Chetumal., Av. Insurgentes 330, Quintana Roo 77013, Mexico.
³El Colegio de la Frontera Sur, Av. Centenario Km 5.5, Chetumal 77014, Quintana Roo, Mexico.

2024 - Volume: 64 Issue: 2 pages: 612-625

Original research

Keywords

Abstract

Introduction

Water mites represent the most abundant, diverse, and ecologically important group of arachnids in all freshwater environments (Smith et al. 2001; Di Sabatino et al. 2008), including interstitial waters, lotic, lentic, and temporary ponds (Di Sabatino et al. 2003). Water mites establish significant and multiple biotic interactions within their communities, primarily as parasites of, among others, several aquatic insects groups, such as plecopterans, dipterans, trichopterans, odonates, hemipterans, and also as predators of insect eggs and larvae, cladocerans, ostracods, and copepods (Proctor et al. 2015). Several studies have documented the effects of water mites on the survival or longevity of their hosts and the influence on the size and structure of their prey populations (Paterson 1970; Gliwicz and Biesiadka 1975; Proctor and Pritchard 2008). Even more significantly, the viability of water mites as bioindicators of the ecosystems they inhabit has been demonstrated, as a clear and predictable relationship exists between the quality of a water body and the composition of its mite fauna (Goldschmidt 2016). Global warming effects has been studied in semi-terrestrial mites (Zhang et al. 2021), but not with hydrachnids. However, some authors affirm that water mites could be a good model for long-term monitoring impacts of freshwater ecosystems including global climate change (Klimov et al. 2022). Therefore, the group constitutes an important tool for environmental diagnosis and biomonitoring.

Approximately 7,500 species have been described worldwide so far (Smit 2020); but the knowledge of faunal diversity is heterogeneous, with some regions being more explored than others and some not yet even explored (Goldschmidt 2002; Goldschmidt et al. 2022). Consequently, some time ago it was estimated that more than 10,000 species could reasonably occur in inland waters (Di Sabatino et al. 2003).

In Mexico, despite being a megadiverse country (Morrone 2019) only 271 species have been recorded since the formal study of the group in 1873 to the present. Therefore, it can be considered that historically, the taxonomic knowledge for this group increased recently, but still it is limited here (Cervantes-Martínez et al. 2023). Currently, the global trend of integrating molecular information (COI barcoding) for water mite studies seems promising in revealing new species, resolving taxonomic questions, and contributing to biodiversity baselines, and assessments (Blattner et al. 2019). This approach could allow their potential use in ecological research or biomonitoring while curatorial work is carried out (Pešić et al. 2021; Więcek et al. 2020).

For Mexican mite fauna, there are already multiple public COI barcode records and identifications down to the genus and family levels, revealing a potential high diversity of this group (Montes-Ortiz et al. 2020, 2022). However, most of these records are not identified to species-level, there is no checklist of water mite species and their distribution in Mexico and COI barcode sequences remain dispersed, challenging a comprehensive integration of data. This paper summarizes the species described in Mexico, their distribution areas, habitat types, and literature sources, and highlights the prospects for the future study of these organisms in the country.

Historical background

A brief summary of the historical background on water mites can be found at Cervantes-Martínez et al. (2023). Taxonomic studies in Mexico began formally with the description of two species of the genera Eylais and Limnesia collected in Guanajuato by the naturalist Alfred Dugés in 1873 and 1884, respectively. Later, in 1936 Ruth Marshall listed three new species (of which Neumania cenotea was synonymized) from the Yucatán state. During the decades of the 70's and 80's, the taxonomic knowledge of water mites experienced substantial growth, due to the contributions of Vidrine (1985a, 1985b, 1985c, 1986, 1996), and David Cook. The latter described in 1974 five species for Mexico, and added six years later 166 species more, on his book about neotropical water mites (Cook 1980). However, it is crucial to acknowledge that during this period, several species that were described, or recorded, were based on a small series of individuals, often lacking information on both sexes and assuming a cosmopolitan distribution of the species. Consequently, it is likely that different morphotypes and their variants described in the literature (e.g., Hydrodroma peregrina perigrina ''The small genital form'' and ''The large genital form'') or subspecies (e.g., Unionicola (Unionicola) gracilipalpis tenuis) pertained in fact to different species.

Simultaneous to new species descriptions, Dr. Ana Hoffmann promoted the studies of mites in Mexico by establishing two laboratories dedicated to acarology, one at the National Polytechnic Institute in 1965 and the other at the National Autonomous University of Mexico in 1977. A year later, one of her students, Dr. Cristina Cramer, made significant contributions in acarology, with the ''Taxonomy, Ecology, and Distribution of Water Mites of Mexico'' project. This led to the addition of 50 new species descriptions between 1977 and 2000 (Rivas and Hoffmann 2000). In the same decade, Dr. Otero-Colina (1987a, 1987b) described another 11 new species from the southeast of Mexico and redescribed some others previously documented by Cook (1980).

Following this period, research on water mites in Mexico experienced a significant decline, only to be reignited with the addition of four new species of arrenurids in 2013 by Dr. Marcia Ramírez (Ramírez-Sánchez and Rivas 2013) and five more under the paradigm of integrative taxonomy by two of us in 2022 (Montes Ortiz et al. 2022).

Material and methods

An exhaustive review of journals and/or books and online literature containing data on water mites in Mexico was carried out. The main reference sources for species described or reported before 1980 were the catalog from Viets (1987) in which the contributions of various authors, such as Müller (1776), Berlese (1888), Piersig (1904), Dugés (1873, 1884) and Marshall (1903, 1908, 1919, 1936) are included. The book ''Water mite genera and subgenera'' (Cook 1974) was also consulted, it includes five species descriptions. The book ''Studies on neotropical water mites'' (Cook 1980) contains 166 original descriptions of species from southern and some northern regions of Mexico. All other records after 1980 included in this list were confirmed through the original publications.

The localities from which species were described or reported, accompanied by the habitat type and the reference are included in the checklist.

For DNA sequences, a public dataset, DS-MEXWM was created at Barcode of Life Data System (https://boldsystems.org ) in which all the sequences from Mexican species available until now are included.

Results and discussion

In total, 271 species of water mites have been described or reported so far in Mexico since 1873. The species belong to 25 families and 61 genera. The families with the greatest species richness were Unionicolidae, Arrenuridae, Aturidae, Hygrobatidae, Limnesiidae, and Torrenticolidae (Figure 1), which together represent more than 75% of the species listed. The richest genera in the list were Arrenurus with 43 species described, followed by Koenikea with 24 species, Unionicola with 18, Atractides with 16 species, and Limnesia with 15.

This number of species is far from representing the true diversity of water mites in the country. Mexico, with an area of 1,964,375 km², boasts a heterogeneous mosaic of ecosystems and complex biogeographic patterns, notably the confluence of the Nearctic and Neotropical regions, making it one of the countries with the highest biodiversity; it occupies second place in the world in species of reptiles and is among the first five as far as species of plants, amphibians, and mammals are concerned (Morrone 2019). Former studies have estimated that there are over 1,000 species of water mites in the Nearctic and over 1,300 in the Neotropical regions (Di Sabatino et al. 2008). However, some authors (Goldschmidt 2002) estimated that the true figure could be much higher, reaching up to 5,500 species in the Neotropical regions alone. Similar cases are observable in other aquatic groups, such as cladocerans, for which the taxonomic list is not complete either and clear differentiation between the Nearctic and Neotropical regions as well as restricted distribution patterns (Elías-Gutiérrez et al. 2001) are observed.

Politically Mexico is divided into 32 states. Nevertheless, this list cataloged water mites in only 19 states (Figure 2). There is no information available for the remaining 13 states located in Neartic and transition zones, representing over half of the country's unexplored territory.

For the states with records or descriptions, the majority corresponded to a few localities within each region. For example, Mexico state (a region in the Mexican highlands, in which the hydrological systems are located at an average altitude > 1000 above sea level) has 39 listed species (Figure 2). Of those, 21 belong to one location, the Stream Peña Blanca in San Francisco Oxtotilpan town (Neotorrenticola davecooki, Neomamersa temazcala, Stygalbiella mexicana, Diamphidaxona anitae, Neoatractides mexicana, Torrenticola keesdavitsi, T. kurtvietsi, T. carlbaderi, T. esbelta, Aturus matlatzinca, A. catoantoni, A. guillecrameri, A. celiahemkesae, A. oxtotilpanensis, A. primitivus, Kongsbergia antoniocatoi, K. cosita, K. tigrina, K. eterna, K. maternal and Kawamuracarus iansmithi) (Cook 1980; Cramer 1987, 2000; Cramer and Smith 1991;Cramer 1992a, 1992b; Cramer and Letechipía 1996), 11 more were described from a stream in the Highway 134 at km 29 (Bandakia mexicana, B. hoffmannae, Chappuisides notialis, Neoacarus adocetus, Eylais mexicana, Thyopsella obscura, Tyrrelia ovalis, Neumania conroyi, Mideopsis mexicana and Limnesia abenda) (Cook 1980; Cramer and Smith 1991). Another five were described from Zempoala lagoons (Sperchon mexicanus, Lebertia azteca, Atractides tanutus, Atractides zempoalus, Arrenurus zempoala) (Cook 1980) and the last two (Hydrachna leovazquezae and Stygarrenurus armoensis) correspond to a pond in Highway Mexico-Tulancingo and a stream in Avándaro, near Valle de Bravo town (Cramer and Costero 1986; Cramer and Cook 1996). This implies that the water mite diversity in the state of Mexico, a region where three hydrological basins converge and with 11, 859 water reservoirs, mostly dams and lakes, but also wetlands (Gobierno del Estado de México 2024), is represented here by only five sampling locations.

Similarly, a study conducted in Queretaro state, a region with high climatic heterogeneity and complex physiogeography characterized by mountainous elevations between 2000 and 3000 above sea level, highlights the limited knowledge of water mite diversity across most of Mexico. In 2014, during a workshop at the Latin American Congress of Freshwater Macroinvertebrates some collections were made for educational purposes at a single sampling point in the ''La Barranca'' stream. This effort yielded 112 specimens from 7 families and 10 genera, including the first record of the genus Protzia in the country. This revealed that Queretaro, for which no prior information was available, now stands as one of the states with more significant water mite diversity in terms of genera (Goldschmidt et al. 2015).

In the Neotropical region of Mexico, the tendency is similar to the transition zone. In Quintana Roo state, which lies adjacent to the Caribbean Sea, seven species have been described out from five localities and the diversity in these environments reaches two families and two genera (Table 1). For this area, Montes-Ortiz and Elías-Gutiérrez (2020) found a richness of 77 putative species in 24 sites using COI barcodes, some of which have been already described (Montes-Ortiz et al. 2022) while the rest are still in the process of identification or description. In recent years, molecular analyses of water mites have demonstrated a consistent increase in relevance for species delimitation. It has been possible to pair sexes of a single species when sexual dimorphism challenges their association and match different stages of their life cycle (Alarcon-Elbal et al. 2020; Montes-Ortiz et al. 2022), reveal species assemblages, infer ecological similarities between water mite communities, and recognize seasonality (Montes-Ortiz et al. 2020; Vasquez et al. 2020; Klimov et al. 2022). Pešić et al. (2021) emphasized the significance of creating COI barcode libraries for water mites.

Currently, from Mexico there are 794 sequences of 898 specimens exclusively from the Yucatán Peninsula (southern Mexico). These sequences corresponded to 103 BINs (Barcode Index Numbers) or putative species, of which 82 are unique in the BOLD database. It means that these sequences are novel in the system and there are no genetically identical taxa in the database, therefore they could be the indicators of endemicity, or limited distribution (Table 2). The families with the most records of COI gene are the Unionicolidae, Arrenuridae, Limnesiidae, Krendowskiidae, Hygrobatidae, Mideopsidae, and Pionidae.

An example of the potential of barcode libraries can be seen in a cosmopolitan genus such as Neumania; with publicly available sequences from 12 countries, it was possible to generate an identification tree. This tree shows that clusters in Mexico, represented by the BINs AEA8101 and ACY6829, form a group separate from those in other regions of the world. Additionally, the differentiation of the two Neumania putative species is evident (Figure 3). The barcode analysis has been useful in determining species distributions (Więcek et al. 2020), and it illustrated that species considered cosmopolitan and recorded in regions far from their type locality are most likely new species. This may be the case of previous identification in Mexico, in which recorded species could originate from different ecoregions from the American continent.

The public barcode database (DS-YUCWM Water Mites Diversity from PY) presented in this work, represents the first reference framework of molecular data of water mites in Mexico. It will allow water mites to be integrated successfully and with certainty as putative species in ecological studies, facilitating a deeper understanding of biotic relationships (Vasquez et al. 2021), and their use in assessments of water quality. However, a Linnean name is still pending. It will be assigned after the fine taxonomical study. We must emphasize that BIN number is not a solution for the so-called taxonomic impediment, and taxonomic study of these putative species should be a priority in further studies.

Conclusion

This is the first checklist reporting all species described or recorded in Mexico, and a public project is made available, integrating all barcode sequences from the country. The analysis of Mexican states with records such as Mexico state, Queretaro and Quintana Roo highlighted that the sampling effort has been limited, underscoring the real water mite diversity here. Similarly, molecular data of the COI gene correspond only to species from Yucatán Peninsula, revealing the necessity of adequate studies in the rest of the country.

This manuscript represents the first reference framework of the current knowledge of morphological diversity and molecular data of water mites in Mexico and we propose this integrative approach to minimize the uncertainty of taxonomic determination toward increasing the reliability of ecological studies using water mites, or assessing their value as indicators of change.

Acknowledgements

This manuscript is part of the results after a Conahcyt postdoctoral fellowship of the first author and belongs to the project ''Ácaros acuáticos (Acari: Hydrachnidia) como (paleo) bioindicadores de cambio Ambiental y antropogénico en sistemas cársticos de la Península de Yucatán'' and is linked to the project Paradigms and Controversies of Science 2022: ''Influencia de la velocidad y magnitud del cambio climático Pleistoceno tardío-Holoceno temprano en las interacciones ecológicas de los ambientes acuáticos del norte del Neotrópico'', project number 319857.

Part of the DNA results were financed by the Global Environment Fund through the United Nations Development Programme (UNDP, Mexico), Comisión Nacional para el Conocimiento y Uso de la Biodiversidad (CONABIO) and Comisión Nacional de Áreas Naturales Protegidas (CONANP) as part of the investigation called: Programa de detección temprana piloto de especies acuáticas invasoras a través de los métodos de código de barras de la vida y análisis de ADN ambiental en la Reserva de la Biosfera Sian Ka'an within Project 00089333 ''Aumentar las capacidades de México para manejar especies exóticas invasoras a través de la implementación de la Estrategia Nacional de Especies Invasoras''.

We thank the editor and the reviewers for their valuable comments that helped to significantly improve this manuscript.

References

1. Alarcón-Elbal, P. M., García-Jiménez, R., Peláez, M. L., Horreo, J. L., Valdecasas, A. G. 2020. Molecular Correlation between Larval, Deutonymph and Adult Stages of the Water Mite Arrenurus (Micruracarus) novus. Life, 10 (7): 108. https://doi.org/10.3390/life10070108
2. Cervantes-Martínez, A., Durán Ramírez, C. A., Elías-Gutiérrez, M., García-Morales, A. E., Gutiérrez-Aguirre, M., Jaime, S., Macek, M., Maeda-Martínez, A. M., Martínez-Jerónimo, F., Mayén-Estrada, R. et al. 2023. Freshwater Diversity of Zooplankton from Mexico: Historical Review of Some of the Main Groups. Water, 15 (5): 858. https://doi.org/10.3390/w15050858
3. Cook, D. R. 1980. Studies on neotropical water mites. Memoirs of the American Entomological Institute, 31: 645.
4. Cramer, C. 1987. Nuevas especies mexicanas de hidracaridos, pertenecientes a los géneros Neomamersa, Kawamuracarus y Neotorrenticola (Acarida: Limnesiidae). Folia Entomológica Mexicana, 73: 185-203.
5. Cramer, C. 1988. Ácaros dulceacuícolas (Acarida: Prostigmata) del Arroyo Peña Blanca en San Francisco Oxtotilpan, México (Doctoral dissertation, Universidad Nacional Autónoma de México), 269.
6. Cramer, C. 1991. Estudios sobre hidracáridos mexicanos. Especies nuevas de la familia Aturidae. Parte 1. Folia Entomológica Mexicana, 82: 173-186.
7. Cramer, C. 1992a. Estudios sobre hidracáridos mexicanos, especies nuevas de la familia Aturidae. Parte II (Aturus: Stygalbiella). Folia Entomológica Mexicana, 84: 113-129.
8. Cramer, C. 1992b. Estudios sobre hidracáridos mexicanos, familia Torrenticolidae. Cinco especies nuevas de Neoatractides y Torrenticola y primer registro de Testudacarus para México. Anales del Instituto de Biología, Universidad Nacional Autónoma de México, Serie Zoología, 63: 13-27.
9. Cramer, C. 2000. New species of the water mite genus Kongsbergia (Acari: Hydrachnidia: Aturidae) from Mexico. International Journal of Acarology, 26: 271-278. https://doi.org/10.1080/01647950008684199
10. Cramer, C., Cook, D. R. 1992a. New species of Arrenurus (Acari: Arrenuridae) from Mexican lakes. Acarologia, 33 (4): 349-366.
11. Cramer, C., Cook, D. R. 1992b. New species of Arrenurus (Dadayella) (Acari: Arrenuridae) from Mexico, with a discussion of the latter's relationships. International Journal of Acarology, 18 (3): 221-229. https://doi.org/10.1080/01647959208683954
12. Cramer, C., Cook, D. R. 1996. A new species of Stygarrenurus (Acari, Hungarohydracarida) and a discussion of its systematic position. International Journal of Acarology, 22 (1): 29-32. https://doi.org/10.1080/01647959608684078
13. Cramer, C., Cook, D. R. 1998. Description of a new subgenus and species of Recifella Viets and a new species of Corticacarus (Acari: Hydracarina) from Mexico. International Journal of Acarology, 24 (2): 131-136. https://doi.org/10.1080/01647959808684141
14. Cramer, C., Cook, D. R. 2000. Water mites of the genera Neoatractides Lundblad and Pseudotorrenticola Walter (Acari: Hydrachnida: Torrenticolidae) from Mexico. International Journal of Acarology, 26 (1): 51-61. https://doi.org/10.1080/01647950008683635
15. Cramer, C. 1986. Primer registro de la familia Feltriidae (Acarida: Hydrachnellae) para México y Descripción de una especie nueva. Folia Entomológica Mexicana, 67: 37-43.
16. Di Sabatino, A., Boggero, A., Miccoli, F.P., Cicolani, B. 2002. Diversity, distribution, and ecology of water mites (Acari: Hydrachnidia and Halacaridae) in high Alpine lakes (Central Alps, Italy). Experimental and Applied Acarology, 34 (1-2): 199-210. https://doi.org/10.1023/B:APPA.0000045251.44202.58
17. Di Sabatino, A., Smit, H., Gerecke, R., Goldschmidt, T., Matsumoto, N., Cicolani, B. 2008. Global diversity of water mites (Acari, Hydrachnidia; Arachnida) in freshwater. Hydrobiologia, 595 (1): 303-315. https://doi.org/10.1007/s10750-007-9025-1
18. Elías-Gutiérrez, M., Smirnov, N., Suárez-Morales, E., Dimas-Flores, N. 2001. New and little known cladocerans (Crustacea: Anomopoda) from southeastern Mexico. Hydrobiologia, 442: 41-54. https://doi.org/10.1023/A:1017506804840
19. Gliwicz, Z.M., Biesiadka E. 1975. Pelagic water mites (Hydracarina) and their effect on the plancton community in a Neotropical man-made lake. Archiv für Hydrobiologie, 76: 65-88.
20. Gobierno del Estado de México. 2024. Inventario estatal de cuerpos de agua. [Internet]. [25 March 2023]. Available from: https://datos.edomex.gob.mx/ar/dataset/apoyar-al-campo/resource/33d2d2aa-f50e-42c9-9190-63712c198e78?inner_span=True
21. Goldschmidt, T. 2002. Environmental parameters determining water mite assemblages in Costa Rica. Experimental and Applied Acarology, 34: 171-197. https://doi.org/10.1023/B:APPA.0000045250.06565.72
22. Goldschmidt, T. 2016. Water mites (Acari, Hydrachnidia): powerful but widely neglected bioindicators-a review. Neotropical Biodiversity, 2 (1): 12-25. https://doi.org/10.1080/23766808.2016.1144359
23. Goldschmidt, T., Ramírez-Sánchez, M.M., Pedroza-Ramos, A., Rico-Sánchez, A.E., Ojeda-Carrasco, M., Pérez-Munguía, R.M., Pimiento-Ortega, M., Carlos-Delgado, A.L., Durán-Suárez, S. 2015. Primer registro de ácaros acuáticos (Acari\,: Hydrachnidia) del estado de Querétaro. Dugesiana, 22 (1): 21-27.
24. Klimov, P.B., Stolbov, V.A., Kazakov, D.V., Filimonova, M.O. 2022. A DNA barcoding and photo-documentation resource of water mites (Acariformes: Hydrachnidia) of Siberia: accurate species identification for global climate change monitoring programs. Systematic and Applied Acarology, 27 (12): 2493-2567. https://doi.org/10.11158/saa.27.12.8
25. Smith, I.M., Cook, D., Smith., B.P. 2001. Water Mites (Hydrachnida) and other Arachnids. In: Thorp, J.H., Covich, A.P. (eds.). Ecology and Classification of North American Freshwater Invertebrates, 2nd Edition. Dairy Science & Technology, CRC Taylor & Francis Group. pp. 551-659. https://doi.org/10.1016/B978-012690647-9/50017-X
26. Marín-Hernández, C.H., Cramer-Hemkes, C. 2009. Especie nueva de hidracárido del género Piona (Acari: Hydrachnidia: Pionidae) de los canales de Xochimilco, México. Revista Mexicana de Biodiversidad, 80 (1): 51-54. https://doi.org/10.22201/ib.20078706e.2009.001.583
27. Montes-Ortiz, L., Elías-Gutiérrez, M. 2020. Water Mite Diversity (Acariformes: Prostigmata: Parasitengonina: Hydrachnidiae) from Karst Ecosystems in Southern Mexico: A Barcoding Approach. Diversity, 12 (329): 1-16. https://doi.org/10.3390/d12090329
28. Montes-Ortiz, L., Elías-Gutiérrez, M., Ramírez-Sánchez, M.M. 2022. Checklist of Arrenurids (Acari: Hydrachnidia: Arrenuridae) of Mexico, with New Records from the Yucatan Peninsula, and the Description of Five New Species of the Subgenera Megaluracarus and Dadayella. Diversity, 14 (4): 276. https://doi.org/10.3390/d14040276
29. Morrone, J.J. 2019. Regionalización biogeográfica y evolución biótica de México: encrucijada de la biodiversidad del Nuevo Mundo. Revista Mexicana de Biodiversidad, 90, e902980. https://doi.org/10.22201/ib.20078706e.2019.90.2980
30. Otero-Colina, G. 1987a. Seis nuevas especies de ácaros acuáticos (Prostigmata: Parasitengona) de los estados de Chiapas, Oaxaca y Tabasco, México. Folia Entomológica Mexicana, 71: 67-85.
31. Otero-Colina, G. 1987b. Nuevas especies de ácaros acuáticos (Prostigmata: Parasitengona) del sureste de México. Folia Entomológica Mexicana, 72: 89-109.
32. Otero-Colina, G. 1988. Datos originales sobre los ácaros acuáticos (Prostigmata: Parasitengona) del sureste de México. Folia Entomológica Mexicana, 76: 195-223.
33. Paterson, C.G. 1970. Water mites (Hydracarina) as predators of chironomid larvae (Insecta: Diptera). Canadian Journal of Zoology, 48: 610-14. https://doi.org/10.1139/z70-112
34. Pešić, V., Zawal, A., Manović, A., Bańkowska, A., Jovanović, M. 2021. A DNA barcode library for the water mites of Montenegro. Biodiversity Data Journal, 9: e78311. https://doi.org/10.3897/BDJ.9.e78311
35. Proctor, H., Pritchard, G. 2008. Neglected Predators: Water Mites (Acari: Parasitengona: Hydrachnellae) in Freshwater Communities. Journal of the North American Benthological Society, 8 (1): 100-111. https://doi.org/10.2307/1467406
36. Proctor H.C., Smith I.M., Cook, D.R., Smith, B.P. 2015. Subphylum Chelicerata. Class Arachnida. In: Thorp, J., Rogers, D.C. (eds.). Ecology and General Biology: Thorp and Covich's Freshwater Invertebrades, Academic Press, pp. 599-660. https://doi.org/10.1016/B978-0-12-385026-3.00025-5
37. Ramírez-Sánchez, M.M., Rivas, G. 2013. New species of subgenus Megaluracarus (Acari: Hydrachnidiae: Arrenuridae: Arrenurus) from Mexico. Zootaxa, 3718 (4): 317-330. https://doi.org/10.11646/zootaxa.3718.4.2
38. Rivas-Lechuga, G., Cramer-Hemkes, C. 1998. Especie nueva de Hidrácarido Arrenurus (Truncaturus) (Acarida: Prostigmata) de Veracruz, México. Anales del Instituto de Biología, Universidad Nacional Autónoma de México, Serie Zoología, 69: 173-179.
39. Smit, H. 2020. Water mites of the world, with keys to the families, subfamilies, genera and subgenera (Acari: Hydrachnidia). Monografieën van de Nederlandse Entomologische Vereniging, 12: 1-774
40. Vasquez, A.A., Carmona-Galindo, V., Qazazi, M.S., Walker, X.N., Ram, J.L. 2020. Water mite assemblages reveal diverse genera, novel DNA barcodes and transitional periods of intermediate disturbance. Experimental and Applied Acarology, 80 (4): 491-507. h https://doi.org/10.1007/s10493-020-00476-4
41. Vasquez, A.A., Mohiddin, O., Li, Z., Bonnici, B.L., Gurdziel, K., Ram, J.L. 2021. Molecular diet studies of water mites reveal prey biodiversity. PLoS ONE, 16 (7): e0254598. https://doi.org/10.1371/journal.pone.0254598
42. Viets, K. 1987. Die Milben des Süßwassers (Hydrachnellae und Halacaridae [part.], Acari). 2: Katalog. Hamburg (Parey). Sonderbände des Naturwissenschaftlichen Vereins in Hamburg, 8. pp. 1012.
43. Vidrine, M. F. 1985a. Berezatax, new subgenus (Acari: Unionicolidae: Unionicolinae: Unionicola), in subtropical and tropical North America, with a re-evaluation of the higher taxonomic groups of related American unionicoline mites. International Journal of Acarology, 11 (2): 133-141. https://doi.org/10.1080/01647958508683403
44. Vidrine, M. F. 1985b. Nine new species in the subgenus Atacella (Acari: Unionicolidae: Unionicola) from Mexico and Brazil. International Journal of Acarology, 11 (4): 255-271. https://doi.org/10.1080/01647958508683426
45. Vidrine, M. F. 1985c. Six new species in the subgenus Polyatax (Acari: Unionicolidae: Unionicola) from North America, with a re-evaluation of related taxa. International Journal of Acarology, 11 (4): 273-287. https://doi.org/10.1080/01647958508683427
46. Vidrine, M. F. 1986. Ten new species in the subgenus Unionicolides (Acari: Unionicolidae: Unionicola) from North and South America, with a re-evaluation of related species. International Journal of Acarology, 12 (3): 123-140. https://doi.org/10.1080/01647958608683455
47. Vidrine, M. F. 1994. Revision of the subgenus Polyatax (Acari: Unionicolidae: Unionicolinae: Unionicola). Texas Journal of Science, 46 (2): 157-163.
48. Więcek, M., Szydło, W., Dabert, J., Proctor, H. 2020. Delimiting species of water mites of the genus Hydrodroma (Acari: Hydrachnidiae: Hydrodromidae) from North America and Europe: Integrative evidence of species status from COI sequences and morphology. Zoologischer Anzeiger, 284: 16-29. https://doi.org/10.1016/j.jcz.2019.11.004
49. Zhang, H., Sun, X., Liu, D., Wu, H. Chen, H. 2021. Air Warming and Drainage Influences Soil Microarthropod Communities. Frontiers in Ecology and Evolution, 9:731735. https://doi.org/10.3389/fevo.2021.731735