# Checklist of Water mites in Mexico. Historical background and DNA barcoding perspectives

Lucia Montes-Ortiz, Sergio Cohuo, Manuel Elías-Gutiérrez

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Lucia Montes-Ortiz, Sergio Cohuo, Manuel Elías-Gutiérrez. Checklist of Water mites in Mexico. Historical background and DNA barcoding perspectives. Acarologia, 2024, 64 (2), pp.612-625. 10.24349/ah6q-r1rk. hal-04582915

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# Checklist of Water mites in Mexico.
Historical background and DNA
barcoding perspectives

Lucia Montes-Ortiz IDª, Sergio Cohuo IDª, Manuel Elías-Gutiérrez IDᵇ

## Original research

### ABSTRACT

Water mites are the most ubiquitous, abundant, diverse, and ecologically important group of arachnids in all freshwater environments. In Mexico, despite being a megadiverse country, only 271 species have been recorded so far since 1873. This is probably due to the scarcity of taxonomists in the country or the inherent complex morphology of this group, which challenges its systematics and its integration into other research studies. Molecular identification has been highly useful for practical applications of these organisms, while taxonomic identification work continues. Until now, a species checklist and reference information about water mites diversity and distribution in Mexico is lacking. In this work, we compiled species described or recorded in Mexico and provided preliminary data for a COI barcode library of the water mites in the country. This checklist includes 271 species from 25 families and 61 genera. In addition, the COI barcode library is supplied in a public dataset from the BOLD database (boldsystems.org); it contains 794 sequences corresponding to 103 BINs or putative species, of which only five of them have been identified to the species. We predict that the discovery of hidden diversity of mites will continue and improve after the use of integrative taxonomy tools. Also, their value as indicators of water quality will be re-evaluated, helping in the conservation of freshwater environments and the evaluation of the global warming impact.

**Keywords** barcode; catalogue; Hydrachnidia; species; richness

### 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.* 2012).

Received 19 January 2024

Accepted 24 April 2024

Published 02 May 2024

Corresponding author

Lucia Montes-Ortiz ID:
lucia.mon@chetumal.tecnm.mx

Academic editor

Mąkol, Joanna

https://doi.org/10.24349/ah6q-r1rk

ISSN 0044-586X (print)

ISSN 2107-7207 (electronic)

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

Montes-Ortiz L. *et al.*

Licensed under
Creative Commons CC-BY 4.0

<footnote>ª Tecnológico Nacional de México–I. T. Chetumal., Av. Insurgentes 330, Quintana Roo 77013, Mexico.</footnote>

<footnote>ᵇ El Colegio de la Frontera Sur, Av. Centenario Km 5.5, Chetumal 77014, Quintana Roo, Mexico.</footnote>
# Checklist of Water mites in Mexico. Historical background and DNA barcoding perspectives

Lucia Montes-Ortiz IDª, Sergio Cohuo IDª, Manuel Elías-Gutiérrez IDᵇ

## Original research

### ABSTRACT

Water mites are the most ubiquitous, abundant, diverse, and ecologically important group of arachnids in all freshwater environments. In Mexico, despite being a megadiverse country, only 271 species have been recorded so far since 1873. This is probably due to the scarcity of taxonomists in the country or the inherent complex morphology of this group, which challenges its systematics and its integration into other research studies. Molecular identification has been highly useful for practical applications of these organisms, while taxonomic identification work continues. Until now, a species checklist and reference information about water mites diversity and distribution in Mexico is lacking. In this work, we compiled species described or recorded in Mexico and provided preliminary data for a COI barcode library of the water mites in the country. This checklist includes 271 species from 25 families and 61 genera. In addition, the COI barcode library is supplied in a public dataset from the BOLD database (boldsystems.org); it contains 794 sequences corresponding to 103 BINs or putative species, of which only five of them have been identified to the species. We predict that the discovery of hidden diversity of mites will continue and improve after the use of integrative taxonomy tools. Also, their value as indicators of water quality will be re-evaluated, helping in the conservation of freshwater environments and the evaluation of the global warming impact.

**Keywords** barcode; catalogue; Hydrachnidia; species; richness

### 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.* 2012).

Received 19 January 2024

Accepted 24 April 2024

Published 02 May 2024

Corresponding author

Lucia Montes-Ortiz ID: lucia.mon@chetumal.tecnm.mx

Academic editor

Mąkol, Joanna

https://doi.org/10.24349/ah6q-r1rk

ISSN 0044-586X (print)

ISSN 2107-7207 (electronic)

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

Montes-Ortiz L. *et al.*

Licensed under

Creative Commons CC-BY 4.0

<footnote>ªTecnológico Nacional de México–I. T. Chetumal., Av. Insurgentes 330, Quintana Roo 77013, Mexico.</footnote>

<footnote>ᵇEl Colegio de la Frontera Sur, Av. Centenario Km 5.5, Chetumal 77014, Quintana Roo, Mexico.</footnote>
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
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
<figure><img src="image_2.png" /><figcaption>**Figure 1** Species richness by family in Mexico.</figcaption></figure>

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
<figure><img src="image_3.png" /><figcaption>**Figure 1** Species richness by family in Mexico.</figcaption></figure>

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
<figure><img src="image_3.png" /><figcaption>**Figure 2** Number of species recorded by state in Mexico.</figcaption></figure>

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, Neoattractides 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 leovazqueae 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
<figure><img src="image_4.png" /><figcaption>**Figure 2** Number of species recorded by state in Mexico.</figcaption></figure>

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, Neoattractides 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 leovazqueae 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
<table><caption>Table 1 Checklist of water mite species in Mexico. * Without data collection.</caption><thead><tr><th>Species</th><th>State</th><th>Habitat</th><th>Reference</th></tr></thead><tbody><tr><td colspan="4"><strong>Arrenuridae</strong></td></tr><tr><td>Arrenurus (Arrenurus) dentipetiolatus</td><td>Oaxaca, Guanajuato</td><td>Pond</td><td>Marshall 1908</td></tr><tr><td>Arrenurus (Arrenurus) valenciatus</td><td>Campeche, Tabasco</td><td>Water-filled roadside</td><td>Marshall 1919</td></tr><tr><td>Arrenurus (Arrenurus) munovus</td><td>Chiapas</td><td>Stream</td><td>Cook 1980</td></tr><tr><td>Arrenurus (Arrenurus) wucabus</td><td>Oaxaca</td><td>Pond</td><td>Cook 1980</td></tr><tr><td>Arrenurus (Arrenurus) tamaulipensis</td><td>Tamaulipas</td><td>Lake</td><td>Cook 1980</td></tr><tr><td>Arrenurus (Arrenurus) xochimilcoensis</td><td>Mexico City</td><td>Standing waters</td><td>Cramer and Cook 1992</td></tr><tr><td>Arrenurus (Megaluracarus) manubriator</td><td>Tabasco</td><td>Pond</td><td>Marshall 1903</td></tr><tr><td>Arrenurus (Megaluracarus) birgei</td><td>Campeche</td><td>Lagoon</td><td>Marshall 1903</td></tr><tr><td>Arrenurus (Megaluracarus) marshallae</td><td>Campeche</td><td>Water-filled ditch</td><td>Piersig 1904</td></tr><tr><td>Arrenurus (Megaluracarus) gricalus</td><td>Veracruz</td><td>Pond</td><td>Cook 1980</td></tr><tr><td>Arrenurus (Megaluracarus) hartesus</td><td>Tabasco</td><td>Pond</td><td>Cook 1980</td></tr><tr><td>Arrenurus (Megaluracarus) neoexpansus</td><td>Tabasco</td><td>Pond</td><td>Cook 1980</td></tr><tr><td>Arrenurus (Megaluracarus) tabascoensis</td><td>Campeche</td><td>Water-filled ditch</td><td>Cook 1980</td></tr><tr><td>Arrenurus (Megaluracarus) trassamus</td><td>Tabasco</td><td>Pond</td><td>Cook 1980</td></tr><tr><td>Arrenurus (Megaluracarus) zitavus</td><td>Campeche</td><td>Water filled-ditch</td><td>Cook 1980</td></tr><tr><td>Arrenurus (Megaluracarus) campechensis</td><td>Campeche</td><td>Water filled-ditch</td><td>Cook 1980</td></tr><tr><td>Arrenurus (Megaluracarus) wolarbus</td><td>Veracruz, Colima</td><td>Pond</td><td>Cook 1980</td></tr><tr><td>Arrenurus (Megaluracarus) costeroae</td><td>Tamaulipas</td><td>Lake</td><td>Cramer and Cook 1992</td></tr><tr><td>Arrenurus (Megaluracarus) alloexpansus</td><td>Colima</td><td>Not specified</td><td>Cramer and Cook 1992</td></tr><tr><td>Arrenurus (Megaluracarus) apizanus</td><td>Tamaulipas</td><td>Lake</td><td>Cramer and Cook 1992</td></tr><tr><td>Arrenurus (Megaluracarus) catoi</td><td>Tamaulipas</td><td>Lake</td><td>Cramer and Cook 1992</td></tr><tr><td>Arrenurus (Megaluracarus) champayanus</td><td>Tamaulipas</td><td>Lake</td><td>Cramer and Cook 1992</td></tr><tr><td>Arrenurus (Megaluracarus) colitus</td><td>Tamaulipas</td><td>Lake</td><td>Cramer and Cook 1992</td></tr><tr><td>Arrenurus (Megaluracarus) anae</td><td>Tamaulipas</td><td>Lake</td><td>Cramer and Cook 1998</td></tr><tr><td>Arrenurus (Megaluracarus) anitahoffmannae</td><td>Tabasco</td><td>Lake, pond, canal</td><td>Ramirez-Sánchez and Rivas 2013</td></tr><tr><td>Arrenurus (Megaluracarus) olmeca</td><td>Tabasco</td><td>Lake, pond, canal</td><td>Ramirez-Sánchez and Rivas 2013</td></tr><tr><td>Arrenurus (Megaluracarus) maya</td><td>Yucatán, Quintana Roo</td><td>Cenote</td><td>Ramirez-Sánchez and Rivas 2013</td></tr><tr><td>Arrenurus (Megaluracarus) urbanus</td><td>Mexico city</td><td>Canal</td><td>Ramirez-Sánchez and Rivas 2013</td></tr><tr><td>Arrenurus (Megaluracarus) eduardoi</td><td>Quintana Roo</td><td>Pool (in a stream)</td><td>Montes-Ortiz et al. 2022</td></tr><tr><td>Arrenurus (Megaluracarus) federicoi</td><td>Quintana Roo</td><td>Pool (in a stream)</td><td>Montes-Ortiz et al. 2022</td></tr><tr><td>Arrenurus (Megaluracarus) ecosur</td><td>Quintana Roo</td><td>Cenote, lagoon, wetlands</td><td>Montes-Ortiz et al. 2022</td></tr><tr><td>Arrenurus (Megaluracarus) beatrizae</td><td>Quintana Roo, Tabasco</td><td>Wetland, lagoon</td><td>Montes-Ortiz et al. 2022</td></tr><tr><td>Arrenurus (Dadayella) zempoala</td><td>Mexico state</td><td>Small stream</td><td>Cook 1980</td></tr><tr><td>Arrenurus (Dadayella) adrianae</td><td>Colima, Michoacan</td><td>Wetland, lagoon</td><td>Cramer and Cook 1992</td></tr><tr><td>Arrenurus (Dadayella) veracruzensis</td><td>Veracruz</td><td>Pond</td><td>Cramer and Cook 1992</td></tr><tr><td>Arrenurus (Dadayella) aztecus</td><td>Veracruz</td><td>Wetland, lagoon</td><td>Cramer and Cook 1992</td></tr><tr><td>Arrenurus (Dadayella) colimensis</td><td>Colima</td><td>Wetland, lagoon</td><td>Cramer and Cook 1992</td></tr><tr><td>Arrenurus (Dadayella) cristinae</td><td>Quintana Roo</td><td>Wetland</td><td>Montes-Ortiz et al. 2022</td></tr><tr><td>Arrenurus (Truncatus) plevamus</td><td>Guerrero</td><td>Small stream</td><td>Cook 1980</td></tr><tr><td>Arrenurus (Truncatus) zukovus</td><td>Chiapas</td><td>Gravel-bottom stream</td><td>Cook 1980</td></tr><tr><td>Arrenurus (Truncatus) teoceloensis</td><td>Veracruz</td><td>Stream</td><td>Rivas and Cramer 1992</td></tr><tr><td>Arrenurus (?) Arrhenuropsis mexicanus</td><td>Tamaulipas, Colima</td><td>Lagoon</td><td>Cramer and Cook 1992</td></tr><tr><td>Arrenurus (?) nayaritensis</td><td>Nayarit</td><td>Small stream</td><td>Cook 1980</td></tr><tr><td colspan="4"><strong>Hydrachnidae</strong></td></tr><tr><td>Hydrachna miliaria</td><td>Oaxaca</td><td>Pond</td><td>Berlese 1888 in Cook 1980</td></tr><tr><td>Hydrachna leovazquezae</td><td>Mexico state</td><td>Temporal pond</td><td>Cramer and Costero 1986</td></tr><tr><td>Hydrachna guanajuatensis</td><td>Guanajuato</td><td>Pond</td><td>Cook 1980</td></tr><tr><td>Hydrachna mexicana</td><td>Veracruz</td><td>Pond</td><td>Cook 1980</td></tr><tr><td>Hydrachna rotunda</td><td>Oaxaca, Nayarit, Guanajuato.</td><td>Pond</td><td>Marshall 1930 in Cook 1980</td></tr><tr><td colspan="4"><strong>Limnocharidae</strong></td></tr><tr><td>Rhyncholimnochores (Rhyncholimnochores)</td><td>Chiapas</td><td>River</td><td>Cook 1980</td></tr><tr><td>Rhyncholimnochores (Paralimnochores)</td><td>Chiapas</td><td>River</td><td>Cook 1980</td></tr><tr><td colspan="4"><strong>Eylaidae</strong></td></tr><tr><td>Eylais guanajuatensis</td><td>Guanajuato</td><td>*</td><td>Dugés 1873</td></tr><tr><td>Eylais mexicana</td><td>Mexico state</td><td>Helocrene spring</td><td>Cook 1980</td></tr><tr><td colspan="4"><strong>Hydriphyantidae</strong></td></tr><tr><td>Hydryphantes (Hydryphantes) ramosus ramosus</td><td>Oaxaca</td><td>Pond</td><td>Daday 1905 in Cook 1980</td></tr><tr><td>Thyopsella obscura</td><td>Mexico state.</td><td>Helocrene spring</td><td>Cook 1980</td></tr><tr><td>Protzia</td><td>Querétaro</td><td>Stream</td><td>Goldschmidt et al. 2015</td></tr><tr><td colspan="4"><strong>Hydrodromidae</strong></td></tr><tr><td>Hydrodroma peregrina perigrina</td><td>Campeche, Guanajuato</td><td>Pond</td><td>Lundblad 1941 in Cook 1980</td></tr><tr><td>Hydrodroma (Clavipes) despisciens</td><td>Oaxaca, Chiapas, Veraruz, San Luis Potosí.</td><td>Stream, river.</td><td>Lundblad 1953 in Cook 1980</td></tr><tr><td colspan="4"><strong>Rhynchohydracaridae</strong></td></tr><tr><td>Clathrosperchon punctatus</td><td>Oaxaca, Chiapas, Veracruz.</td><td>Matted roots, bottom of stream</td><td>Cook 1980</td></tr><tr><td colspan="4"><strong>Sperchontidae</strong></td></tr><tr><td>Sperchon mexicanus</td><td>Mexico state</td><td>Stream</td><td>Cook 1980</td></tr><tr><td>Sperchon neotropicus</td><td>Oaxaca, Coahuila, Veracruz.</td><td>Algae on rocks, stream</td><td>Cook 1980</td></tr><tr><td>Sperchon (Hispidopershon) gledhilli</td><td>Oaxaca, Veracruz, Guerrero.</td><td>Stream</td><td>Viets 1977</td></tr><tr><td>Sperchon (Acadiosperchon) pontifex</td><td>Oaxaca</td><td>Submerged vegetation</td><td>Otero-Colina 1987a</td></tr><tr><td>Sperchon (Mixosperchon) oaxacensis</td><td>Oaxaca</td><td>Submerged vegetation</td><td>Otero-Colina 1987a</td></tr><tr><td colspan="4"><strong>Anisitsielidae</strong></td></tr><tr><td>Bandakia hoffmannae</td><td>Mexico state.</td><td>Spring</td><td>Cramer and Smith 1991</td></tr><tr><td>Bandakia mexicana</td><td>Mexico state.</td><td>Spring</td><td>Cramer and Smith 1991</td></tr><tr><td>Mamersella mesoamericana</td><td>Tabasco</td><td>Lagoon</td><td>Otero-Colina 1987a</td></tr><tr><td>Mamersellides costenius</td><td>Veracruz, Colima, Estados Unidos (Florida)</td><td>Ponds, Marshes, backwater pools of streams.</td><td>Cramer and Smith 1993</td></tr></tbody></table>
<table><caption>Table 1 Checklist of water mite species in Mexico. * Without data collection.</caption><thead><tr><th>Species</th><th>State</th><th>Habitat</th><th>Reference</th></tr></thead><tbody><tr><td colspan="4"><strong>Arrenuridae</strong></td></tr><tr><td>Arrenurus (Arrenurus) dentipetiolatus</td><td>Oaxaca, Guanajuato</td><td>Pond</td><td>Marshall 1908</td></tr><tr><td>Arrenurus (Arrenurus) valenciatus</td><td>Campeche, Tabasco</td><td>Water-filled roadside</td><td>Marshall 1919</td></tr><tr><td>Arrenurus (Arrenurus) munovus</td><td>Chiapas</td><td>Stream</td><td>Cook 1980</td></tr><tr><td>Arrenurus (Arrenurus) wucabus</td><td>Oaxaca</td><td>Pond</td><td>Cook 1980</td></tr><tr><td>Arrenurus (Arrenurus) tamaulipensis</td><td>Tamaulipas</td><td>Lake</td><td>Cook 1980</td></tr><tr><td>Arrenurus (Arrenurus) xochimilcoensis</td><td>Mexico City</td><td>Standing waters</td><td>Cramer and Cook 1992</td></tr><tr><td>Arrenurus (Megaluracarus) manubriator</td><td>Tabasco</td><td>Pond</td><td>Marshall 1903</td></tr><tr><td>Arrenurus (Megaluracarus) birgei</td><td>Campeche</td><td>Lagoon</td><td>Marshall 1903</td></tr><tr><td>Arrenurus (Megaluracarus) marshallae</td><td>Campeche</td><td>Water-filled ditch</td><td>Piersig 1904</td></tr><tr><td>Arrenurus (Megaluracarus) gricalus</td><td>Veracruz</td><td>Pond</td><td>Cook 1980</td></tr><tr><td>Arrenurus (Megaluracarus) hartesus</td><td>Tabasco</td><td>Pond</td><td>Cook 1980</td></tr><tr><td>Arrenurus (Megaluracarus) neoexpansus</td><td>Tabasco</td><td>Pond</td><td>Cook 1980</td></tr><tr><td>Arrenurus (Megaluracarus) tabascoensis</td><td>Campeche</td><td>Water-filled ditch</td><td>Cook 1980</td></tr><tr><td>Arrenurus (Megaluracarus) trassamus</td><td>Tabasco</td><td>Pond</td><td>Cook 1980</td></tr><tr><td>Arrenurus (Megaluracarus) zitavus</td><td>Campeche</td><td>Water filled-ditch</td><td>Cook 1980</td></tr><tr><td>Arrenurus (Megaluracarus) campechensis</td><td>Campeche</td><td>Water filled-ditch</td><td>Cook 1980</td></tr><tr><td>Arrenurus (Megaluracarus) wolarbus</td><td>Veracruz, Colima</td><td>Pond</td><td>Cook 1980</td></tr><tr><td>Arrenurus (Megaluracarus) costeroae</td><td>Tamaulipas</td><td>Lake</td><td>Cramer and Cook 1992</td></tr><tr><td>Arrenurus (Megaluracarus) alloexpansus</td><td>Colima</td><td>Not specified</td><td>Cramer and Cook 1992</td></tr><tr><td>Arrenurus (Megaluracarus) apizanus</td><td>Tamaulipas</td><td>Lake</td><td>Cramer and Cook 1992</td></tr><tr><td>Arrenurus (Megaluracarus) catoi</td><td>Tamaulipas</td><td>Lake</td><td>Cramer and Cook 1992</td></tr><tr><td>Arrenurus (Megaluracarus) champayanus</td><td>Tamaulipas</td><td>Lake</td><td>Cramer and Cook 1992</td></tr><tr><td>Arrenurus (Megaluracarus) colitus</td><td>Tamaulipas</td><td>Lake</td><td>Cramer and Cook 1992</td></tr><tr><td>Arrenurus (Megaluracarus) anae</td><td>Tamaulipas</td><td>Lake</td><td>Cramer and Cook 1998</td></tr><tr><td>Arrenurus (Megaluracarus) anitahoffmannae</td><td>Tabasco</td><td>Lake, pond, canal</td><td>Ramirez-Sánchez and Rivas 2013</td></tr><tr><td>Arrenurus (Megaluracarus) olmeca</td><td>Tabasco</td><td>Lake, pond, canal</td><td>Ramirez-Sánchez and Rivas 2013</td></tr><tr><td>Arrenurus (Megaluracarus) maya</td><td>Yucatán, Quintana Roo</td><td>Cenote</td><td>Ramirez-Sánchez and Rivas 2013</td></tr><tr><td>Arrenurus (Megaluracarus) urbanus</td><td>Mexico city</td><td>Canal</td><td>Ramirez-Sánchez and Rivas 2013</td></tr><tr><td>Arrenurus (Megaluracarus) eduardoi</td><td>Quintana Roo</td><td>Pool (in a stream)</td><td>Montes-Ortiz et al. 2022</td></tr><tr><td>Arrenurus (Megaluracarus) federicoi</td><td>Quintana Roo</td><td>Pool (in a stream)</td><td>Montes-Ortiz et al. 2022</td></tr><tr><td>Arrenurus (Megaluracarus) ecosur</td><td>Quintana Roo</td><td>Cenote, lagoon, wetlands</td><td>Montes-Ortiz et al. 2022</td></tr><tr><td>Arrenurus (Megaluracarus) beatrizae</td><td>Quintana Roo, Tabasco</td><td>Wetland, lagoon</td><td>Montes-Ortiz et al. 2022</td></tr><tr><td>Arrenurus (Dadayella) zempoala</td><td>Mexico state</td><td>Small stream</td><td>Cook 1980</td></tr><tr><td>Arrenurus (Dadayella) adrianae</td><td>Colima, Michoacan</td><td>Wetland, lagoon</td><td>Cramer and Cook 1992</td></tr><tr><td>Arrenurus (Dadayella) veracruzensis</td><td>Veracruz</td><td>Pond</td><td>Cramer and Cook 1992</td></tr><tr><td>Arrenurus (Dadayella) aztecus</td><td>Veracruz</td><td>Wetland, lagoon</td><td>Cramer and Cook 1992</td></tr><tr><td>Arrenurus (Dadayella) colimensis</td><td>Colima</td><td>Wetland, lagoon</td><td>Cramer and Cook 1992</td></tr><tr><td>Arrenurus (Dadayella) cristinae</td><td>Quintana Roo</td><td>Wetland</td><td>Montes-Ortiz et al. 2022</td></tr><tr><td>Arrenurus (Truncatus) plevamus</td><td>Guerrero</td><td>Small stream</td><td>Cook 1980</td></tr><tr><td>Arrenurus (Truncatus) zukovus</td><td>Chiapas</td><td>Gravel-bottom stream</td><td>Cook 1980</td></tr><tr><td>Arrenurus (Truncatus) teoceloensis</td><td>Veracruz</td><td>Stream</td><td>Rivas and Cramer 1992</td></tr><tr><td>Arrenurus (?) Arrhenuropsis mexicanus</td><td>Tamaulipas, Colima</td><td>Lagoon</td><td>Cramer and Cook 1992</td></tr><tr><td>Arrenurus (?) nayaritensis</td><td>Nayarit</td><td>Small stream</td><td>Cook 1980</td></tr><tr><td colspan="4"><strong>Hydrachnidae</strong></td></tr><tr><td>Hydrachna miliaria</td><td>Oaxaca</td><td>Pond</td><td>Berlese 1888 in Cook 1980</td></tr><tr><td>Hydrachna leovazquezae</td><td>Mexico state</td><td>Temporal pond</td><td>Cramer and Costero 1986</td></tr><tr><td>Hydrachna guanajuatensis</td><td>Guanajuato</td><td>Pond</td><td>Cook 1980</td></tr><tr><td>Hydrachna mexicana</td><td>Veracruz</td><td>Pond</td><td>Cook 1980</td></tr><tr><td>Hydrachna rotunda</td><td>Oaxaca, Nayarit, Guanajuato.</td><td>Pond</td><td>Marshall 1930 in Cook 1980</td></tr><tr><td colspan="4"><strong>Limnocharidae</strong></td></tr><tr><td>Rhyncholimnochores (Rhyncholimnochores)</td><td>Chiapas</td><td>River</td><td>Cook 1980</td></tr><tr><td>Rhyncholimnochores (Paralimnochores)</td><td>Chiapas</td><td>River</td><td>Cook 1980</td></tr><tr><td colspan="4"><strong>Eylaidae</strong></td></tr><tr><td>Eylais guanajuatensis</td><td>Guanajuato</td><td>*</td><td>Dugés 1873</td></tr><tr><td>Eylais mexicana</td><td>Mexico state</td><td>Helocrene spring</td><td>Cook 1980</td></tr><tr><td colspan="4"><strong>Hydriyphantidae</strong></td></tr><tr><td>Hydryphantes (Hydryphantes) ramosus ramosus</td><td>Oaxaca</td><td>Pond</td><td>Daday 1905 in Cook 1980</td></tr><tr><td>Thyopsella obscura</td><td>Mexico state.</td><td>Helocrene spring</td><td>Cook 1980</td></tr><tr><td>Protzia</td><td>Querétaro</td><td>Stream</td><td>Goldschmidt et al. 2015</td></tr><tr><td colspan="4"><strong>Hydrodromidae</strong></td></tr><tr><td>Hydrodroma peregrina perigrina</td><td>Campeche, Guanajuato</td><td>Pond</td><td>Lundblad 1941 in Cook 1980</td></tr><tr><td>Hydrodroma (Clavipes) despisciens</td><td>Oaxaca, Chiapas, Veraruz, San Luis Potosí.</td><td>Stream, river.</td><td>Lundblad 1953 in Cook 1980</td></tr><tr><td colspan="4"><strong>Rhynchohydracaridae</strong></td></tr><tr><td>Clathrosperchon punctatus</td><td>Oaxaca, Chiapas, Veracruz.</td><td>Matted roots, bottom of stream</td><td>Cook 1980</td></tr><tr><td colspan="4"><strong>Sperchontidae</strong></td></tr><tr><td>Sperchon mexicanus</td><td>Mexico state</td><td>Stream</td><td>Cook 1980</td></tr><tr><td>Sperchon neotropicus</td><td>Oaxaca, Coahuila, Veracruz.</td><td>Algae on rocks, stream</td><td>Cook 1980</td></tr><tr><td>Sperchon (Hispidopershon) gledhilli</td><td>Oaxaca, Veracruz, Guerrero.</td><td>Stream</td><td>Viets 1977</td></tr><tr><td>Sperchon (Acadiosperchon) pontifex</td><td>Oaxaca</td><td>Submerged vegetation</td><td>Otero-Colina 1987a</td></tr><tr><td>Sperchon (Mixosperchon) oaxacensis</td><td>Oaxaca</td><td>Submerged vegetation</td><td>Otero-Colina 1987a</td></tr><tr><td colspan="4"><strong>Anisitsielidae</strong></td></tr><tr><td>Bandakia hoffmannae</td><td>Mexico state.</td><td>Spring</td><td>Cramer and Smith 1991</td></tr><tr><td>Bandakia mexicana</td><td>Mexico state.</td><td>Spring</td><td>Cramer and Smith 1991</td></tr><tr><td>Mamersella mesoamericana</td><td>Tabasco</td><td>Lagoon</td><td>Otero-Colina 1987a</td></tr><tr><td>Mamersellides costenius</td><td>Veracruz, Colima, Estados Unidos (Florida)</td><td>Ponds, Marshes, backwater pools of streams.</td><td>Cramer and Smith 1993</td></tr></tbody></table>
<table><caption>Table 1 Continued.</caption><thead><tr><th>Species</th><th>State</th><th>Habitat</th><th>Reference</th></tr></thead><tbody><tr><td><strong>Lebertiidae</strong></td><td></td><td></td><td></td></tr><tr><td>Lebertia (Pseudolebertia) azteca</td><td>Mexico state</td><td>Stream</td><td>Cook 1980</td></tr><tr><td><strong>Oxidae</strong></td><td></td><td></td><td></td></tr><tr><td>Oxus dugesi</td><td>Veracruz, Costa Rica</td><td>Stream</td><td>Cook 1980</td></tr><tr><td>Oxus stolli</td><td>Guerrero</td><td>Tributary river</td><td>Cook 1980</td></tr><tr><td><strong>Torrenticolidae</strong></td><td></td><td></td><td></td></tr><tr><td>Torrenticola (Monoattractides) lembada</td><td>Chiapas, Costa Rica</td><td>Stream, river.</td><td>Cook 1980</td></tr><tr><td>Torrenticola (Monoattractides) gorda</td><td>Nayarit, Guerrero, Oaxaca</td><td>River</td><td>Cook 1980</td></tr><tr><td>Torrenticola (Monoattractides) veracruzensis</td><td>Veracruz, Chiapas</td><td>Stream</td><td>Cook 1980</td></tr><tr><td>Torrenticola (Torrenticola) rala</td><td>Guerrero</td><td>River</td><td>Cook 1980</td></tr><tr><td>Torrenticola (Torrenticola) keesdavidsi</td><td>Mexico state.</td><td>Stream</td><td>Cramer 1992</td></tr><tr><td>Torrenticola (Torrenticola) carlbaderi</td><td>Mexico state.</td><td>Stream</td><td>Cramer 1992</td></tr><tr><td>Torrenticola (Torrenticola) esbelta</td><td>Mexico state.</td><td>Stream</td><td>Cramer 1992</td></tr><tr><td>Torrenticola (Torrenticola) kurtvietsi</td><td>Mexico state</td><td>Stream</td><td>Cramer 1992</td></tr><tr><td>Neotorrenticola (Neotorrenticola) davecooki</td><td>Mexico state</td><td>Stream</td><td>Cramer 1987</td></tr><tr><td>Pseudotorrenticola (Pseudotorrenticola)</td><td>Veracruz</td><td>Stream</td><td>Cramer and Cook 2000</td></tr><tr><td>Pseudotorrenticola (Pseudotorrenticola) boettgeri</td><td>Veracruz</td><td>Stream</td><td>Viets 1977 in Cramer and Cook 2000</td></tr><tr><td><strong>Limnesiidae</strong></td><td></td><td></td><td></td></tr><tr><td>Neomamersa (Neomamersa) mexicana</td><td>Oaxaca, Guerrero</td><td>Bottom deposits of stream</td><td>Cook 1980</td></tr><tr><td>Neomamersa (Neomamersa) triacebulata</td><td>Chiapas</td><td>Gravels deposits of stream</td><td>Cook 1980</td></tr><tr><td>Neomamersa (Neomamersa) temazcala</td><td>Mexico state</td><td>Stream</td><td>Cramer 1987</td></tr><tr><td>Meramecia (Meramecia)mexicana</td><td>Chiapas, Oaxaca, Puebla</td><td>Gravel deposits of stream</td><td>Cook 1980</td></tr><tr><td>Kawamuracarus (Kawamuracarus) expansipes</td><td>Oaxaca, Chiapas</td><td>Gravel deposits of stream</td><td>Cook 1980</td></tr><tr><td>Kawamuracarus (Kawamuracarus) novus</td><td>Puebla</td><td>Gravel deposits of stream</td><td>Cook 1980</td></tr><tr><td>Kawamuracarus (Kawamuracarus) iansmithi</td><td>Mexico state</td><td>Stream</td><td>Cramer 1987</td></tr><tr><td>Tyrrelia viuda</td><td>Puebla</td><td>Stream</td><td>Cook 1980</td></tr><tr><td>Tyrrelia ovalis</td><td>Puebla, Mexico state</td><td>Stream</td><td>Marshall 1940 in Cook 1980</td></tr><tr><td>Centrolimnesia (Centrolimnesia)motasi</td><td>Chiapas</td><td>Pond</td><td>Cook 1980</td></tr><tr><td>Centrolimnesia (Centrolimnesia) bondi</td><td>Veracruz, Campeche</td><td>Swam, Pond</td><td>Lundblad 1935 in Cook 1980</td></tr><tr><td>Limnesia alzatei</td><td>Guanajuato</td><td>*</td><td>Dugés 1884</td></tr><tr><td>Limnesia laeta</td><td>Oaxaca, Tabasco</td><td>Pond</td><td>Stoll 1887 in Cook 1980</td></tr><tr><td>Limnesia puteorum</td><td>Guanajuato</td><td>*</td><td>Stoll 1887 in Wolcott 1903</td></tr><tr><td>Limnesia (Limnesia) abenda</td><td>Mexico state</td><td>Helocrene spring</td><td>Cook 1980</td></tr><tr><td>Limnesia (Limnesia) banola</td><td>Nayarit, Veracruz</td><td>Spring</td><td>Cook 1980</td></tr><tr><td>Limnesia (Limnesia) canvada</td><td>Nayarit</td><td>River</td><td>Cook 1980</td></tr><tr><td>Limnesia (Limnesia) dimorpha</td><td>Nayarit</td><td>Spring</td><td>Cook 1980</td></tr><tr><td>Limnesia (Limnesia) dornera</td><td>Veracruz</td><td>Stream</td><td>Cook 1980</td></tr><tr><td>Limnesia (Limnesia) mesoamericana</td><td>Guerrero, Oaxaca, Chiapas, Morelos, Nayarit</td><td>River, stream</td><td>Cook 1980</td></tr><tr><td>Limnesia (Limnesia) neodentipalpis</td><td>Veracruz</td><td>Pond</td><td>Cook 1980</td></tr><tr><td>Limnesia (Limnesia) setacoxalis</td><td>Veracruz, Chiapas, Guerrero</td><td>Stream</td><td>Cook 1980</td></tr><tr><td>Limnesia (Limnesia) trehona</td><td>Guerrero</td><td>Spring</td><td>Cook 1980</td></tr><tr><td>Limnesia (Limnesia) zomplina</td><td>Nayarit, Veracruz</td><td>Pond</td><td>Cook 1980</td></tr><tr><td>Limnesia (Limnesiella) mexicana</td><td>Tabasco</td><td>Pond</td><td>Cook 1980</td></tr><tr><td>Limnesia parafalsificata</td><td>Tabasco, Chiapas</td><td>Pond</td><td>Otero-Colina 1987a</td></tr><tr><td>Psammolimnesia (Psammolimnesia) mexicana</td><td>Chiapas</td><td></td><td>Cook 1974</td></tr><tr><td><strong>Omartacaridae</strong></td><td></td><td></td><td></td></tr><tr><td>Omartacarus elongatus</td><td>Puebla</td><td>Interstitial waters</td><td>Cook 1963 in Cook 1980</td></tr><tr><td>Omartacarus brevipalpis</td><td>Chiapas</td><td>Interstitial waters</td><td>Cook 1974</td></tr><tr><td>Omartacarus motasi</td><td>Chiapas</td><td>Sand and gravel of stream</td><td>Cook 1980</td></tr><tr><td><strong>Hygrobatidae</strong></td><td></td><td></td><td></td></tr><tr><td>Corticacarus novum</td><td>Veracruz</td><td>Stream</td><td>Cramer and Cook 1998</td></tr><tr><td>Corticacarus (Paracorticacarus) mexicanus</td><td>Guerrero, Oaxaca, Chiapas</td><td>Stream</td><td>Cook 1980</td></tr><tr><td>Corticacarus (Polycorticacarellus) similis</td><td>Guerrero</td><td>Stream</td><td>Cook 1980</td></tr><tr><td>Corticacarus (Polycorticacarellus) mebosus</td><td>Chiapas, Veracruz</td><td>River</td><td>Cook 1980</td></tr><tr><td>Neoattractides curvispinatus</td><td>Tabasco</td><td>Gravel in a stream</td><td>Otero-Colina 1987a</td></tr><tr><td>Neoattractides mexicana</td><td>Mexico state</td><td>Stream</td><td>Cramer 1992</td></tr><tr><td>Neoattractides nayaritensis</td><td>Nayarit</td><td>Stream</td><td>Cramer and Cook 2000</td></tr><tr><td>Neoattractides abueloi</td><td>Veracruz</td><td>Stream</td><td>Cramer and Cook 2000</td></tr><tr><td>Neoattractides celiae</td><td>Veracruz</td><td>Stream</td><td>Cramer and Cook 2000</td></tr><tr><td>Hyrobates (Hyrobates) boettgeri</td><td>Oaxaca, Chiapas, Guerrero, Morelos, Nayarit</td><td>Stream</td><td>Viets 1975 in Cook 1980</td></tr><tr><td>Hyrobates (Hyrobates) plebejus</td><td>Oaxaca, Chiapas, Veracruz, Morelos, Nayarit.</td><td>River</td><td>Lundblad 1930 in Cook 1930</td></tr><tr><td>Hyrobates (Hyrobates) amplipalpis</td><td>Oaxaca, Veracruz</td><td>Stream</td><td>Viets 1936 in Cook 1980</td></tr><tr><td>Hyrobates (Hyrobates) ampliatus</td><td>Veracruz, San Luis Potosí, Morelos</td><td>Stream</td><td>Viets 1936 in Cook 1980</td></tr><tr><td>Hyrobates (Hyrobates) procursus</td><td>Guerrero</td><td>Stream</td><td>Viets 1936 in Cook 1980</td></tr><tr><td>Hyrobates (Hyrobates)amatolus</td><td>Oaxaca</td><td>Tributary river</td><td>Cook 1980</td></tr><tr><td>Hyrobates (Hyrobates)blatolus</td><td>Nayarit</td><td>River</td><td>Cook 1980</td></tr><tr><td>Hyrobates (Hyrobates) mexicanus</td><td>Guerrero</td><td>Stream</td><td>Lundblad 1942 in Cook 1980</td></tr><tr><td>Atractides (Atractides) crassitarsis</td><td>Oaxaca</td><td>River</td><td>Viets 1978 in Cook 1980</td></tr><tr><td>Atractides (Atractides) imitatus</td><td>Oaxaca</td><td>Tributary river, stream</td><td>Viets 1978 in Cook 1980</td></tr><tr><td>Atractides (Atractides) guatemaltecus</td><td>Oaxaca, Chiapas, Guerrero.</td><td></td><td></td></tr><tr><td>Atractides (Atractides) blazonus</td><td>Oaxaca</td><td>Tributary river</td><td>Cook 1980</td></tr><tr><td>Atractides (Atractides) mexicanus</td><td>Oaxaca</td><td>Tributary river</td><td>Cook 1980</td></tr><tr><td>Atractides (Atractides) neongitarsus</td><td>Coahuila</td><td>River</td><td>Cook 1980</td></tr><tr><td>Atractides (Atractides) prasadi</td><td>Chiapas</td><td>Stream</td><td>Cook 1980</td></tr><tr><td>Atractides (Atractides) tanutus</td><td>Mexico state</td><td>Stream</td><td>Cook 1980</td></tr><tr><td>Atractides (Atractides) tembolus</td><td>Chiapas</td><td>Stream</td><td>Cook 1980</td></tr><tr><td>Atractides (Atractides) tolas</td><td>Chiapas</td><td>Stream</td><td>Cook 1980</td></tr></tbody></table>
<table><caption>Table 1 Continued.</caption><thead><tr><th>Species</th><th>State</th><th>Habitat</th><th>Reference</th></tr></thead><tbody><tr><td><strong>Lebertiidae</strong></td><td></td><td></td><td></td></tr><tr><td>Lebertia (Pseudolebertia) azteca</td><td>Mexico state</td><td>Stream</td><td>Cook 1980</td></tr><tr><td><strong>Oxidae</strong></td><td></td><td></td><td></td></tr><tr><td>Oxus dugesi</td><td>Veracruz, Costa Rica</td><td>Stream</td><td>Cook 1980</td></tr><tr><td>Oxus stolli</td><td>Guerrero</td><td>Tributary river</td><td>Cook 1980</td></tr><tr><td><strong>Torrenticolidae</strong></td><td></td><td></td><td></td></tr><tr><td>Torrenticola (Monoattractides) lembada</td><td>Chiapas, Costa Rica</td><td>Stream, river.</td><td>Cook 1980</td></tr><tr><td>Torrenticola (Monoattractides) gorda</td><td>Nayarit, Guerrero, Oaxaca</td><td>River</td><td>Cook 1980</td></tr><tr><td>Torrenticola (Monoattractides) veracruzensis</td><td>Veracruz, Chiapas</td><td>Stream</td><td>Cook 1980</td></tr><tr><td>Torrenticola (Torrenticola) rala</td><td>Guerrero</td><td>River</td><td>Cook 1980</td></tr><tr><td>Torrenticola (Torrenticola) keesdavidsi</td><td>Mexico state.</td><td>Stream</td><td>Cramer 1992</td></tr><tr><td>Torrenticola (Torrenticola) carlbaderi</td><td>Mexico state.</td><td>Stream</td><td>Cramer 1992</td></tr><tr><td>Torrenticola (Torrenticola) esbelta</td><td>Mexico state.</td><td>Stream</td><td>Cramer 1992</td></tr><tr><td>Torrenticola (Torrenticola) kurtvietsi</td><td>Mexico state</td><td>Stream</td><td>Cramer 1992</td></tr><tr><td>Neotorrenticola (Neotorrenticola) davecooki</td><td>Mexico state</td><td>Stream</td><td>Cramer 1987</td></tr><tr><td>Pseudotorrenticola (Pseudotorrenticola)</td><td>Veracruz</td><td>Stream</td><td>Cramer and Cook 2000</td></tr><tr><td>Pseudotorrenticola (Pseudotorrenticola) boettgeri</td><td>Veracruz</td><td>Stream</td><td>Viets 1977 in Cramer and Cook 2000</td></tr><tr><td><strong>Limnesiidae</strong></td><td></td><td></td><td></td></tr><tr><td>Neomamersa (Neomamersa) mexicana</td><td>Oaxaca, Guerrero</td><td>Bottom deposits of stream</td><td>Cook 1980</td></tr><tr><td>Neomamersa (Neomamersa) triacebulata</td><td>Chiapas</td><td>Gravels deposits of stream</td><td>Cook 1980</td></tr><tr><td>Neomamersa (Neomamersa) temazcala</td><td>Mexico state</td><td>Stream</td><td>Cramer 1987</td></tr><tr><td>Meramecia (Meramecia)mexicana</td><td>Chiapas, Oaxaca, Puebla</td><td>Gravel deposits of stream</td><td>Cook 1980</td></tr><tr><td>Kawamuracarus (Kawamuracarus) expansipes</td><td>Oaxaca, Chiapas</td><td>Gravel deposits of stream</td><td>Cook 1980</td></tr><tr><td>Kawamuracarus (Kawamuracarus) novus</td><td>Puebla</td><td>Gravel deposits of stream</td><td>Cook 1980</td></tr><tr><td>Kawamuracarus (Kawamuracarus) iansmithi</td><td>Mexico state</td><td>Stream</td><td>Cramer 1987</td></tr><tr><td>Tyrrelia viuda</td><td>Puebla</td><td>Stream</td><td>Cook 1980</td></tr><tr><td>Tyrrelia ovalis</td><td>Puebla, Mexico state</td><td>Stream</td><td>Marshall 1940 in Cook 1980</td></tr><tr><td>Centrolimnesia (Centrolimnesia)motasi</td><td>Chiapas</td><td>Pond</td><td>Cook 1980</td></tr><tr><td>Centrolimnesia (Centrolimnesia) bondi</td><td>Veracruz, Campeche</td><td>Swam, Pond</td><td>Lundblad 1935 in Cook 1980</td></tr><tr><td>Limnesia alzatei</td><td>Guanajuato</td><td>*</td><td>Dugés 1884</td></tr><tr><td>Limnesia laeta</td><td>Oaxaca, Tabasco</td><td>Pond</td><td>Stoll 1887 in Cook 1980</td></tr><tr><td>Limnesia puteorum</td><td>Guanajuato</td><td>*</td><td>Stoll 1887 in Wolcott 1903</td></tr><tr><td>Limnesia (Limnesia) abenda</td><td>Mexico state</td><td>Helocrene spring</td><td>Cook 1980</td></tr><tr><td>Limnesia (Limnesia) banola</td><td>Nayarit, Veracruz</td><td>Spring</td><td>Cook 1980</td></tr><tr><td>Limnesia (Limnesia) canvada</td><td>Nayarit</td><td>River</td><td>Cook 1980</td></tr><tr><td>Limnesia (Limnesia) dimorpha</td><td>Nayarit</td><td>Spring</td><td>Cook 1980</td></tr><tr><td>Limnesia (Limnesia) dornera</td><td>Veracruz</td><td>Stream</td><td>Cook 1980</td></tr><tr><td>Limnesia (Limnesia) mesoamericana</td><td>Guerrero, Oaxaca, Chiapas, Morelos, Nayarit</td><td>River, stream</td><td>Cook 1980</td></tr><tr><td>Limnesia (Limnesia) neodentipalpis</td><td>Veracruz</td><td>Pond</td><td>Cook 1980</td></tr><tr><td>Limnesia (Limnesia) setacoxalis</td><td>Veracruz, Chiapas, Guerrero</td><td>Stream</td><td>Cook 1980</td></tr><tr><td>Limnesia (Limnesia) trehona</td><td>Guerrero</td><td>Spring</td><td>Cook 1980</td></tr><tr><td>Limnesia (Limnesia) zomplina</td><td>Nayarit, Veracruz</td><td>Pond</td><td>Cook 1980</td></tr><tr><td>Limnesia (Limnesiella) mexicana</td><td>Tabasco</td><td>Pond</td><td>Cook 1980</td></tr><tr><td>Limnesia parafalsificata</td><td>Tabasco, Chiapas</td><td>Pond</td><td>Otero-Colina 1987a</td></tr><tr><td>Psammolimnesia (Psammolimnesia) mexicana</td><td>Chiapas</td><td></td><td>Cook 1974</td></tr><tr><td><strong>Omartacaridae</strong></td><td></td><td></td><td></td></tr><tr><td>Omartacarus elongatus</td><td>Puebla</td><td>Interstitial waters</td><td>Cook 1963 in Cook 1980</td></tr><tr><td>Omartacarus brevipalpis</td><td>Chiapas</td><td>Interstitial waters</td><td>Cook 1974</td></tr><tr><td>Omartacarus motasi</td><td>Chiapas</td><td>Sand and gravel of stream</td><td>Cook 1980</td></tr><tr><td><strong>Hygrobatidae</strong></td><td></td><td></td><td></td></tr><tr><td>Corticacarus novum</td><td>Veracruz</td><td>Stream</td><td>Cramer and Cook 1998</td></tr><tr><td>Corticacarus (Paracorticacarus) mexicanus</td><td>Guerrero, Oaxaca, Chiapas</td><td>Stream</td><td>Cook 1980</td></tr><tr><td>Corticacarus (Polycorticacarellus) similis</td><td>Guerrero</td><td>Stream</td><td>Cook 1980</td></tr><tr><td>Corticacarus (Polycorticacarellus) mebosus</td><td>Chiapas, Veracruz</td><td>River</td><td>Cook 1980</td></tr><tr><td>Neoattractides curvispinatus</td><td>Tabasco</td><td>Gravel in a stream</td><td>Otero-Colina 1987a</td></tr><tr><td>Neoattractides mexicana</td><td>Mexico state</td><td>Stream</td><td>Cramer 1992</td></tr><tr><td>Neoattractides nayaritensis</td><td>Nayarit</td><td>Stream</td><td>Cramer and Cook 2000</td></tr><tr><td>Neoattractides abueloi</td><td>Veracruz</td><td>Stream</td><td>Cramer and Cook 2000</td></tr><tr><td>Neoattractides celiae</td><td>Veracruz</td><td>Stream</td><td>Cramer and Cook 2000</td></tr><tr><td>Hyrobates (Hyrobates) boettgeri</td><td>Oaxaca, Chiapas, Guerrero, Morelos, Nayarit</td><td>Stream</td><td>Viets 1975 in Cook 1980</td></tr><tr><td>Hyrobates (Hyrobates) plebejus</td><td>Oaxaca, Chiapas, Veracruz, Morelos, Nayarit.</td><td>River</td><td>Lundblad 1930 in Cook 1930</td></tr><tr><td>Hyrobates (Hyrobates) amplipalpis</td><td>Oaxaca, Veracruz</td><td>Stream</td><td>Viets 1936 in Cook 1980</td></tr><tr><td>Hyrobates (Hyrobates) ampliatus</td><td>Veracruz, San Luis Potosí, Morelos</td><td>Stream</td><td>Viets 1936 in Cook 1980</td></tr><tr><td>Hyrobates (Hyrobates) procursus</td><td>Guerrero</td><td>Stream</td><td>Viets 1936 in Cook 1980</td></tr><tr><td>Hyrobates (Hyrobates)amatolus</td><td>Oaxaca</td><td>Tributary river</td><td>Cook 1980</td></tr><tr><td>Hyrobates (Hyrobates)blatolus</td><td>Nayarit</td><td>River</td><td>Cook 1980</td></tr><tr><td>Hyrobates (Hyrobates) mexicanus</td><td>Guerrero</td><td>Stream</td><td>Lundblad 1942 in Cook 1980</td></tr><tr><td>Atractides (Atractides) crassitarsis</td><td>Oaxaca</td><td>River</td><td>Viets 1978 in Cook 1980</td></tr><tr><td>Atractides (Atractides) imitatus</td><td>Oaxaca</td><td>Tributary river, stream</td><td>Viets 1978 in Cook 1980</td></tr><tr><td>Atractides (Atractides) guatemaltecus</td><td>Oaxaca, Chiapas, Guerrero.</td><td></td><td></td></tr><tr><td>Atractides (Atractides) blazonus</td><td>Oaxaca</td><td>Tributary river</td><td>Cook 1980</td></tr><tr><td>Atractides (Atractides) mexicanus</td><td>Oaxaca</td><td>Tributary river</td><td>Cook 1980</td></tr><tr><td>Atractides (Atractides) neongitarsus</td><td>Coahuila</td><td>River</td><td>Cook 1980</td></tr><tr><td>Atractides (Atractides) prasadi</td><td>Chiapas</td><td>Stream</td><td>Cook 1980</td></tr><tr><td>Atractides (Atractides) tanutus</td><td>Mexico state</td><td>Stream</td><td>Cook 1980</td></tr><tr><td>Atractides (Atractides) tembolus</td><td>Chiapas</td><td>Stream</td><td>Cook 1980</td></tr><tr><td>Atractides (Atractides) tolas</td><td>Chiapas</td><td>Stream</td><td>Cook 1980</td></tr></tbody></table>
<table><caption>Table 1 Continued.</caption><thead><tr><th>Species</th><th>State</th><th>Habitat</th><th>Reference</th></tr></thead><tbody><tr><td>Atractides (Atractides) toldomus</td><td>Coahuila</td><td>River</td><td>Cook 1980</td></tr><tr><td>Atractides (Atractides) travanus</td><td>Chiapas, Oaxaca, Guerrero</td><td>Stream</td><td>Cook 1980</td></tr><tr><td>Atractides (Atractides) wevanus</td><td>Nayarit</td><td>River bottom deposits</td><td>Cook 1980</td></tr><tr><td>Atractides (Atractides) zempoalus</td><td>Mexico state</td><td>Stream</td><td>Cook 1980</td></tr><tr><td>Atractides (Atractides) bassolsae</td><td>Oaxaca</td><td>Stream</td><td>Otero-Colina 1987a</td></tr><tr><td>Atractides (Octomegapus) peltatus</td><td>Chiapas</td><td>River</td><td>Viets 1977 in Cook 1980</td></tr><tr><td>Atractidella obtusidens</td><td>Oaxaca, Chiapas, Veracruz</td><td>River, stream</td><td>Lundblad 1953 in Cook 1980</td></tr><tr><td>Atractidella mesoamericana</td><td>Chiapas</td><td>Stream</td><td>Cook 1980</td></tr><tr><td>Paraschizobates scutatus</td><td>Nayarit, Chiapas, Oaxaca</td><td>River, stream</td><td>Cook 1980</td></tr><tr><td>Diamphidaxona mexicana</td><td>Chiapas</td><td>Gravel deposits in stream</td><td>Cook 1980</td></tr><tr><td>Diamphidaxona anitae</td><td>Mexico state</td><td>Stream</td><td>Cramer and Letechipía 1996</td></tr><tr><td colspan="4"><strong>Unionicolidae</strong></td></tr><tr><td>Unionicola gracilipalpis tenuis</td><td>Oaxaca, Campeche</td><td>Pond, water filled roadside</td><td>Lundblad 1935 in Cook 1980</td></tr><tr><td>Unionicola (Unionicola) mexicana</td><td>Tabasco</td><td>Pond</td><td>Cook 1980</td></tr><tr><td>Unionicola (Pentatax) conjuncta</td><td>Chiapas</td><td>Stream</td><td>Viets 1954 in Cook 1980</td></tr><tr><td>Unionicola (Pentatax) furculopsis</td><td>Oaxaca</td><td>Pond</td><td>Cook 1980</td></tr><tr><td>Unionicola (Pentatax) aculeata</td><td>Tamaulipas, northern and central Mexico</td><td>Freshwater mussel Anodonta sp., Cyrtonaias tampicoensis, Disconaias sp., Friersonia sp.</td><td>Koenike 1914 in Vidrine 1996</td></tr><tr><td>Unionicola (Atacella) neoperforata</td><td>Tamaulipas, San Luis Potosí.</td><td>Freshwater mussel A. trapezialis glaucus</td><td>Vidrine 1985b</td></tr><tr><td>Unionicola (Atacella) petita</td><td>Veracruz</td><td>Freshwater mussel A. trapezialis glaucus</td><td>Vidrine 1985b</td></tr><tr><td>Unionicola (Atacella) entrerrianensis</td><td>Tamaulipas, San Luis Potosí, Veracruz.</td><td>Freshwater mussel A. trapezialis glaucus</td><td>Rosso de Ferradás 1976</td></tr><tr><td>Unionicola (Atacella) fissipes</td><td>Western Mexico</td><td>Freshwater mussel Anodontites</td><td>Koenike, 1891</td></tr><tr><td>Unionicola (Berezatax) berezai</td><td>Veracruz, San Luis Potosí</td><td>Freshwater mussel Nephronaias sp.</td><td>Vidrine 1985a</td></tr><tr><td>Unionicola (Berezatax) acylindrotarsa</td><td>Tamaulipas, San Luis Potosí</td><td>Freshwater mussel Popenaias, Nephronaias</td><td>Vidrine 1985a</td></tr><tr><td>Unionicola (Causeyatax) hensleyi</td><td>San Luis Potosí, Tamaulipas</td><td>Freshwater mussel Friersonia iridella</td><td>Vidrine 1985c</td></tr><tr><td>Unionicola (Clarkatax) serrata</td><td>San Luis Potosí, Tamaulipas, Veracruz (Panuco river system)</td><td>Friersonia iridella, F. moctezumensis</td><td>Wolcott 1898 in Vidrine 1996</td></tr><tr><td>Unionicola (Hexatax) laurentiana</td><td>San Luis Potosí, Tamaulipas, Veracruz (Panuco river system)</td><td>Freshwater mussel Disconaias fimbriata, Nephronaias sp., Popenaias sp.</td><td>Crowell and Davids 1979 in</td></tr><tr><td>Unionicola (Unionicolides) burchi</td><td>Guanajuato</td><td>Freshwater mussel Anodonta sp.</td><td>Vidrine 1996</td></tr><tr><td>Unionicola (Unionicolides) calnani</td><td>Tamaulipas, Veracruz</td><td>Freshwater mussel Cyrtonaias tampicoensis, Disconaias discus,</td><td>Vidrine 1986</td></tr><tr><td>Unionicola (Unionicolides) hoesei</td><td>San Luis Potosí, Tamaulipas, Veracruz (Panuco river system)</td><td>Freshwater mussel</td><td>Vidrine 1986</td></tr><tr><td>Unionicola (Neoatax) abnormipes</td><td>Tamaulipas</td><td>Freshwater mussel Cyrtonaias tampicoensis</td><td>Vidrine 1986</td></tr><tr><td>Neumania (Neumania) conroyi</td><td>Mexico state</td><td>Reocrene spring</td><td>Cook 1980</td></tr><tr><td>Neumania (Neumania) diversipalpa</td><td>Chiapas</td><td>River</td><td>Cook 1980</td></tr><tr><td>Neumania (Neumania) alticola</td><td>Chiapas, Campeche, Oaxaca, Tabasco, Veracruz.</td><td>Pond, stream.</td><td>Stoll 1886 in Cook 1980</td></tr><tr><td>Neumania (Tetraneumania) nondanda</td><td>Chiapas, Tabasco, Campeche</td><td>River</td><td>Cook 1980</td></tr><tr><td>Neumania (Tetraneumania) cesateca</td><td>Tabasco, Chiapas</td><td>River, Pond</td><td>Otero-Colina 1987b</td></tr><tr><td>Recifella (Recifella) bella</td><td>Veracruz, Chiapas</td><td>River, stream</td><td>Cook 1980</td></tr><tr><td>Recifella (Eorecifella?) azteca</td><td>Oaxaca</td><td>Pond</td><td>Cook 1980</td></tr><tr><td>Recifella (Eorecifella?) cemoba</td><td>Chiapas</td><td>Stream</td><td>Cook 1980</td></tr><tr><td>Recifella (Eorecifella) laversi</td><td>Chiapas</td><td>River</td><td>Cook 1980</td></tr><tr><td>Recifella (Eorecifella) alpa</td><td>Veracruz</td><td>Stream</td><td>Cook 1980</td></tr><tr><td>Recifella (Eorecifella) indistincta</td><td>Yucatán</td><td>Cenote (Sinkhole)</td><td>Marshall 1936 in Cook 1980</td></tr><tr><td>Recifella (Recifellopsis) mexicana</td><td>Chiapas, Veracruz</td><td>Stream</td><td>Cook 1980</td></tr><tr><td>Recifella (Vietsella) veracruzana</td><td>Veracruz</td><td>Pond</td><td>Cramer and Cook 1998</td></tr><tr><td>Koenikea (Koenikea) canerma</td><td>Tabasco</td><td>Pond</td><td>Cook 1980</td></tr><tr><td>Koenikea (Koenikea) clazona</td><td>Veracruz</td><td>Stream</td><td>Cook 1980</td></tr><tr><td>Koenikea (Koenikea) gracilipes</td><td>Nayarit, Chiapas, Guerrero</td><td>Stream</td><td>Cook 1980</td></tr><tr><td>Koenikea (Koenikea) tobarana</td><td>Nayarit</td><td>River, stream</td><td>Cook 1980</td></tr><tr><td>Koenikea (Koenikea) trocala</td><td>Guerrero</td><td>Spring</td><td>Cook 1980</td></tr><tr><td>Koenikea (Koenikea) yenona</td><td>Nayarit</td><td>Tributary river</td><td>Cook 1980</td></tr><tr><td>Koenikea (Koenikea) zedima</td><td>Chiapas, Tabasco</td><td>River</td><td>Cook 1980</td></tr><tr><td>Koenikea (Koenikea) zolada</td><td>Veracruz</td><td>River</td><td>Cook 1980</td></tr><tr><td>Koenikea mesoamericana</td><td>Chiapas</td><td>Stream</td><td>Cook 1980</td></tr><tr><td>Koenikea (?Diplokoenikea) bispina</td><td>Veracruz</td><td>River</td><td>Cook 1980</td></tr><tr><td>Koenikea (?Diplokoenikea) bleptá</td><td>Guerrero</td><td>Stream</td><td>Cook 1980</td></tr><tr><td>Koenikea (?Diplokoenikea) beltista</td><td>Chiapas, Veracruz, Puebla.</td><td>River</td><td>Cook 1980</td></tr><tr><td>Koenikea (Diplokoenikea) neopteinifera</td><td>Campeche, Tabasco</td><td>Waterfilled ditch, pond</td><td>Cook 1980</td></tr><tr><td>Koenikea (Diplokoenikea) clavigera</td><td>Chiapas, Oaxaca</td><td>Pond</td><td>Lundblad 1943 in Cook 1980</td></tr><tr><td>Koenikea (Notomideopsis) paragrossa</td><td>Oaxaca</td><td>Pond</td><td>Cook 1980</td></tr><tr><td>Koenikea (Notomideopsis) sexmaculata</td><td>Oaxaca, Veracruz</td><td>Pond</td><td>Cook 1980</td></tr><tr><td>Koenikea (Notomideopsis) toloma</td><td>Veracruz</td><td>Stream</td><td>Cook 1980</td></tr><tr><td>Koenikea (Notomideopsis) veracruzensis</td><td>Veracruz</td><td>Stream</td><td>Cook 1980</td></tr><tr><td>Koenikea (Notomideopsis) nayaritensis</td><td>Nayarit</td><td>Spring</td><td>Cook 1980</td></tr><tr><td>Koenikea (Notomideopsis) taninulensis</td><td>San Luis Potosí</td><td>Sulphurous stream</td><td>Cramer 1983</td></tr><tr><td>Koenikea (Sespekenikea) chiapasana</td><td>Chiapas</td><td>Stream</td><td>Cook 1980</td></tr><tr><td>Koenikea (Sespekenikea) piotiformis</td><td>Nayarit, Guerrero</td><td>River</td><td>Cook 1980</td></tr><tr><td>Koenikea (Tanaognathus) kurtvietsi</td><td>Guerrero</td><td>River</td><td>Cook 1980</td></tr><tr><td>Koenikea (Parakoenikea) curvipalpis</td><td>Oaxaca</td><td>River</td><td>Lundblad 1936 in Cook 1980</td></tr><tr><td colspan="4"><strong>Pionidae</strong></td></tr><tr><td>Piona bromada</td><td>Tabasco, Campeche, Veracruz, Oaxaca.</td><td>Pond, waterfilled ditch</td><td>Cook 1980</td></tr><tr><td>Piona mexicana</td><td>Veracruz</td><td>Stream</td><td>Cook 1980</td></tr><tr><td>Piona neoacutidens</td><td>Veracruz</td><td>Stream</td><td>Cook 1980</td></tr><tr><td>Piona (Piona) triangularis</td><td>*</td><td>*</td><td>Wolcott 1902 in Marín-Hernández and Cramer-Hemkes 2009</td></tr></tbody></table>
<table><caption>Table 1 Continued.</caption><thead><tr><th>Species</th><th>State</th><th>Habitat</th><th>Reference</th></tr></thead><tbody><tr><td>Atractides (Atractides) toldomus</td><td>Coahuila</td><td>River</td><td>Cook 1980</td></tr><tr><td>Atractides (Atractides) travanus</td><td>Chiapas, Oaxaca, Guerrero</td><td>Stream</td><td>Cook 1980</td></tr><tr><td>Atractides (Atractides) wevanus</td><td>Nayarit</td><td>River bottom deposits</td><td>Cook 1980</td></tr><tr><td>Atractides (Atractides) zempoalus</td><td>Mexico state</td><td>Stream</td><td>Cook 1980</td></tr><tr><td>Atractides (Atractides) bassolsae</td><td>Oaxaca</td><td>Stream</td><td>Otero-Colina 1987a</td></tr><tr><td>Atractides (Octomegapus) peltatus</td><td>Chiapas</td><td>River</td><td>Viets 1977 in Cook 1980</td></tr><tr><td>Atractidella obtusidens</td><td>Oaxaca, Chiapas, Veracruz</td><td>River, stream</td><td>Lundblad 1953 in Cook 1980</td></tr><tr><td>Atractidella mesoamericana</td><td>Chiapas</td><td>Stream</td><td>Cook 1980</td></tr><tr><td>Paraschizobates scutatus</td><td>Nayarit, Chiapas, Oaxaca</td><td>River, stream</td><td>Cook 1980</td></tr><tr><td>Diamphidaxona mexicana</td><td>Chiapas</td><td>Gravel deposits in stream</td><td>Cook 1980</td></tr><tr><td>Diamphidaxona anitae</td><td>Mexico state</td><td>Stream</td><td>Cramer and Letechipía 1996</td></tr><tr><td><strong>Unionicolidae</strong></td><td></td><td></td><td></td></tr><tr><td>Unionicola gracilipalpis tenuis</td><td>Oaxaca, Campeche</td><td>Pond, water filled roadside</td><td>Lundblad 1935 in Cook 1980</td></tr><tr><td>Unionicola (Unionicola) mexicana</td><td>Tabasco</td><td>Pond</td><td>Cook 1980</td></tr><tr><td>Unionicola (Pentatax) conjuncta</td><td>Chiapas</td><td>Stream</td><td>Viets 1954 in Cook 1980</td></tr><tr><td>Unionicola (Pentatax) furculopsis</td><td>Oaxaca</td><td>Pond</td><td>Cook 1980</td></tr><tr><td>Unionicola (Pentatax) aculeata</td><td>Tamaulipas, northern and central Mexico</td><td>Freshwater mussel Anodonta sp., Cyrtonaias tampicoensis, Disconaias sp., Friersonia sp.</td><td>Koenike 1914 in Vidrine 1996</td></tr><tr><td>Unionicola (Atacella) neoperforata</td><td>Tamaulipas, San Luis Potosí.</td><td>Freshwater mussel A. trapezialis glaucus</td><td>Vidrine 1985b</td></tr><tr><td>Unionicola (Atacella) petita</td><td>Veracruz</td><td>Freshwater mussel A. trapezialis glaucus</td><td>Vidrine 1985b</td></tr><tr><td>Unionicola (Atacella) entrerrianensis</td><td>Tamaulipas, San Luis Potosí, Veracruz.</td><td>Freshwater mussel A. trapezialis glaucus</td><td>Rosso de Ferradás 1976</td></tr><tr><td>Unionicola (Atacella) fissipes</td><td>Western Mexico</td><td>Freshwater mussel Anodontites</td><td>Koenike, 1891</td></tr><tr><td>Unionicola (Berezatax) berezai</td><td>Veracruz, San Luis Potosí</td><td>Freshwater mussel Nephronaias sp.</td><td>Vidrine 1985a</td></tr><tr><td>Unionicola (Berezatax) acylindrotarsa</td><td>Tamaulipas, San Luis Potosí</td><td>Freshwater mussel Popenaias, Nephronaias</td><td>Vidrine 1985a</td></tr><tr><td>Unionicola (Causeyatax) hensleyi</td><td>San Luis Potosí, Tamaulipas</td><td>Freshwater mussel Friersonia iridella</td><td>Vidrine 1985c</td></tr><tr><td>Unionicola (Clarkatax) serrata</td><td>San Luis Potosí, Tamaulipas, Veracruz (Panuco river system)</td><td>Friersonia iridella, F. moctezumensis</td><td>Wolcott 1898 in Vidrine 1996</td></tr><tr><td>Unionicola (Hexatax) laurentiana</td><td>San Luis Potosí, Tamaulipas, Veracruz (Panuco river system)</td><td>Freshwater mussel Disconaias fimbriata, Nephronaias sp., Popenaias sp.</td><td>Crowell and Davids 1979 in</td></tr><tr><td>Unionicola (Unionicolides) burchi</td><td>Guanajuato</td><td>Freshwater mussel Anodonta sp.</td><td>Vidrine 1996</td></tr><tr><td>Unionicola (Unionicolides) calnani</td><td>Tamaulipas, Veracruz</td><td>Freshwater mussel Cyrtonaias tampicoensis, Disconaias discus,</td><td>Vidrine 1986</td></tr><tr><td>Unionicola (Unionicolides) hoesei</td><td>San Luis Potosí, Tamaulipas, Veracruz (Panuco river system)</td><td>Freshwater mussel</td><td>Vidrine 1986</td></tr><tr><td>Unionicola (Neoatax) abnormipes</td><td>Tamaulipas</td><td>Freshwater mussel Cyrtonaias tampicoensis</td><td>Vidrine 1986</td></tr><tr><td>Neumania (Neumania) conroyi</td><td>Mexico state</td><td>Reocrene spring</td><td>Cook 1980</td></tr><tr><td>Neumania (Neumania) diversipalpa</td><td>Chiapas</td><td>River</td><td>Cook 1980</td></tr><tr><td>Neumania (Neumania) alticola</td><td>Chiapas, Campeche, Oaxaca, Tabasco, Veracruz.</td><td>Pond, stream.</td><td>Stoll 1886 in Cook 1980</td></tr><tr><td>Neumania (Tetraneumania) nondanda</td><td>Chiapas, Tabasco, Campeche</td><td>River</td><td>Cook 1980</td></tr><tr><td>Neumania (Tetraneumania) cesateca</td><td>Tabasco, Chiapas</td><td>River, Pond</td><td>Otero-Colina 1987b</td></tr><tr><td>Recifella (Recifella) bella</td><td>Veracruz, Chiapas</td><td>River, stream</td><td>Cook 1980</td></tr><tr><td>Recifella (Eorecifella?) azteca</td><td>Oaxaca</td><td>Pond</td><td>Cook 1980</td></tr><tr><td>Recifella (Eorecifella?) cemoba</td><td>Chiapas</td><td>Stream</td><td>Cook 1980</td></tr><tr><td>Recifella (Eorecifella) laversi</td><td>Chiapas</td><td>River</td><td>Cook 1980</td></tr><tr><td>Recifella (Eorecifella) alpa</td><td>Veracruz</td><td>Stream</td><td>Cook 1980</td></tr><tr><td>Recifella (Eorecifella) indistincta</td><td>Yucatán</td><td>Cenote (Sinkhole)</td><td>Marshall 1936 in Cook 1980</td></tr><tr><td>Recifella (Recifellopsis) mexicana</td><td>Chiapas, Veracruz</td><td>Stream</td><td>Cook 1980</td></tr><tr><td>Recifella (Vietsella) veracruzana</td><td>Veracruz</td><td>Pond</td><td>Cramer and Cook 1998</td></tr><tr><td>Koenikea (Koenikea) canerma</td><td>Tabasco</td><td>Pond</td><td>Cook 1980</td></tr><tr><td>Koenikea (Koenikea) clazona</td><td>Veracruz</td><td>Stream</td><td>Cook 1980</td></tr><tr><td>Koenikea (Koenikea) gracilipes</td><td>Nayarit, Chiapas, Guerrero</td><td>Stream</td><td>Cook 1980</td></tr><tr><td>Koenikea (Koenikea) tobarana</td><td>Nayarit</td><td>River, stream</td><td>Cook 1980</td></tr><tr><td>Koenikea (Koenikea) trocala</td><td>Guerrero</td><td>Spring</td><td>Cook 1980</td></tr><tr><td>Koenikea (Koenikea) yenona</td><td>Nayarit</td><td>Tributary river</td><td>Cook 1980</td></tr><tr><td>Koenikea (Koenikea) zedima</td><td>Chiapas, Tabasco</td><td>River</td><td>Cook 1980</td></tr><tr><td>Koenikea (Koenikea) zolada</td><td>Veracruz</td><td>River</td><td>Cook 1980</td></tr><tr><td>Koenikea mesoamericana</td><td>Chiapas</td><td>Stream</td><td>Cook 1980</td></tr><tr><td>Koenikea (?Diplokoenikea) bispina</td><td>Veracruz</td><td>River</td><td>Cook 1980</td></tr><tr><td>Koenikea (?Diplokoenikea) blepta</td><td>Guerrero</td><td>Stream</td><td>Cook 1980</td></tr><tr><td>Koenikea (?Diplokoenikea) beltista</td><td>Chiapas, Veracruz, Puebla.</td><td>River</td><td>Cook 1980</td></tr><tr><td>Koenikea (Diplokoenikea) neopteinifera</td><td>Campeche, Tabasco</td><td>Waterfilled ditch, pond</td><td>Cook 1980</td></tr><tr><td>Koenikea (Diplokoenikea) clavigera</td><td>Chiapas, Oaxaca</td><td>Pond</td><td>Lundblad 1943 in Cook 1980</td></tr><tr><td>Koenikea (Notomideopsis) paragrossa</td><td>Oaxaca</td><td>Pond</td><td>Cook 1980</td></tr><tr><td>Koenikea (Notomideopsis) sexmaculata</td><td>Oaxaca, Veracruz</td><td>Pond</td><td>Cook 1980</td></tr><tr><td>Koenikea (Notomideopsis) toloma</td><td>Veracruz</td><td>Stream</td><td>Cook 1980</td></tr><tr><td>Koenikea (Notomideopsis) veracruzensis</td><td>Veracruz</td><td>Stream</td><td>Cook 1980</td></tr><tr><td>Koenikea (Notomideopsis) nayaritensis</td><td>Nayarit</td><td>Spring</td><td>Cook 1980</td></tr><tr><td>Koenikea (Notomideopsis) taninulensis</td><td>San Luis Potosí</td><td>Sulphurous stream</td><td>Cramer 1983</td></tr><tr><td>Koenikea (Sespekenikea) chiapasana</td><td>Chiapas</td><td>Stream</td><td>Cook 1980</td></tr><tr><td>Koenikea (Sespekenikea) piotiformis</td><td>Nayarit, Guerrero</td><td>River</td><td>Cook 1980</td></tr><tr><td>Koenikea (Tanaognathus) kurtvietsi</td><td>Guerrero</td><td>River</td><td>Cook 1980</td></tr><tr><td>Koenikea (Parakoenikea) curvipalpis</td><td>Oaxaca</td><td>River</td><td>Lundblad 1936 in Cook 1980</td></tr><tr><td><strong>Pionidae</strong></td><td></td><td></td><td></td></tr><tr><td>Piona bromada</td><td>Tabasco, Campeche, Veracruz, Oaxaca.</td><td>Pond, waterfilled ditch</td><td>Cook 1980</td></tr><tr><td>Piona mexicana</td><td>Veracruz</td><td>Stream</td><td>Cook 1980</td></tr><tr><td>Piona neoacutidens</td><td>Veracruz</td><td>Stream</td><td>Cook 1980</td></tr><tr><td>Piona (Piona) triangularis</td><td>*</td><td>*</td><td>Wolcott 1902 in Marín-Hernández and Cramer-Hemkes 2009</td></tr></tbody></table>
<table><caption>Table 1 Continued.</caption><thead><tr><th>Species</th><th>State</th><th>Habitat</th><th>Reference</th></tr></thead><tbody><tr><td>Piona (Tetrapiona) rotunda sudamericana</td><td>*</td><td>*</td><td>Viets 1910 in Marín-Hernández and Cramer-Hemkes 2009</td></tr><tr><td>Piona alzatei</td><td>*</td><td>*</td><td>Dugés 1884</td></tr><tr><td>Piona pearsei</td><td>*</td><td>*</td><td>Marshall 1936</td></tr><tr><td>Piona amimitli</td><td>Mexico city</td><td>Stream</td><td>Marín-Hernández 2009</td></tr><tr><td colspan="4"><strong>Aturidae</strong></td></tr><tr><td>Aturus matlatzinca</td><td>Mexico state</td><td>Stream</td><td>Cramer 1991</td></tr><tr><td>Aturus catoantoni</td><td>Mexico state</td><td>Stream</td><td>Cramer 1991</td></tr><tr><td>Aturus guillecrameri</td><td>Mexico state</td><td>Stream</td><td>Cramer 1992</td></tr><tr><td>Aturus celiahemkesae</td><td>Mexico state</td><td>Stream</td><td>Cramer 1992</td></tr><tr><td>Aturus oxtoilpanensis</td><td>Mexico state</td><td>Stream</td><td>Cramer 1992</td></tr><tr><td>Aturus primitivus</td><td>Mexico state</td><td>Stream</td><td>Cramer 1991</td></tr><tr><td>Neoaturus kurtvietsi</td><td>Oaxaca, Chiapas, Veracruz, Guerrero</td><td>River, stream</td><td>Cook 1980</td></tr><tr><td>Neoaturus projectus</td><td>Veracruz</td><td>River</td><td>Lundblad 1936 in Cook 1980</td></tr><tr><td>Adelazonopsella pallida</td><td>Veracruz</td><td>Interstitial water stream</td><td>Cook 1974</td></tr><tr><td>Albia (Anchistalbia) iantha</td><td>Nayarit</td><td>Spring</td><td>Cook 1980</td></tr><tr><td>Albia (Anchistalbia) ianthopsis</td><td>Quintana Roo</td><td>Lagoon</td><td>Otero-Colina 1987b</td></tr><tr><td>Axonopsella (?) mesoamericana</td><td>Guerrero, Morelos, Nayarit</td><td>River</td><td>Cook 1980</td></tr><tr><td>Axonopsella (Neoazonopsella) nayaritensis</td><td>Nayarit</td><td>Cataracts</td><td>Cook 1980</td></tr><tr><td>Axonopsella (Neoazonopsella) vicina</td><td>Veracruz</td><td>Stream</td><td>Cook 1980</td></tr><tr><td>Axonopsis (Brachypodopsis) columbica</td><td>Chiapas, Oaxaca, Puebla. Nayarit</td><td>River, stream</td><td>Lundblad 1953 in Cook 1980</td></tr><tr><td>Axonopsis (Brachypodopsis) cemoba</td><td>Veracruz</td><td>Stream</td><td>Cook 1980</td></tr><tr><td>Axonopsis (Brachypodopsis) cerucha</td><td>Veracruz, San Luis Potosí</td><td>River, Stream</td><td>Cook 1980</td></tr><tr><td>Axonopsis (Brachypodopsis) mesoamericana</td><td>Veracruz, Chiapas</td><td>Stream</td><td>Cook 1980</td></tr><tr><td>Axonopsis (Brachypodopsis) trocala</td><td>Chiapas</td><td>Chiapas</td><td>Cook 1980</td></tr><tr><td>Polyazonopsella mexicana</td><td>Oaxaca</td><td>Stream</td><td>Otero-Colina 1987b</td></tr><tr><td>Frontipodopsis mesoamericana</td><td>Chiapas</td><td>Bottom deposits of stream</td><td>Cook 1980</td></tr><tr><td>Kongsbergia antoniocatoi</td><td>Mexico state</td><td>Gravel deposits of stream</td><td>Cramer 2000</td></tr><tr><td>Kongsbergia cosita</td><td>Mexico state</td><td>Gravel deposits of stream</td><td>Cramer 2000</td></tr><tr><td>Kongsbergia tigrina</td><td>Mexico state</td><td>Stream</td><td>Cramer 2000</td></tr><tr><td>Kongsbergia eterna</td><td>Mexico state</td><td>Stream</td><td>Cramer 2000</td></tr><tr><td>Kongsbergia (Kongsbergia) mexicana</td><td>Oaxaca</td><td>River</td><td>Cook 1980</td></tr><tr><td>Kongsbergia glopipalpis</td><td>Chiapas and Veracruz</td><td>Stream</td><td>Lundblad 1953 in Cook 1980</td></tr><tr><td>Kongsbergia materna</td><td>Chiapas, Mexico state, Oaxaca, Veracruz.</td><td>Tributary river</td><td>Thor 1899 in Cramer 2000</td></tr><tr><td>Submiraxona (Submiraxona) bella</td><td>Nayarit</td><td>Stream</td><td>Cook 1980</td></tr><tr><td>Submiraxona (Submiraxona) lundbladi</td><td>Guerrero, Chiapas</td><td>River</td><td>Cook 1980</td></tr><tr><td>Submiraxona (Submiraxona) stolli</td><td>Guerrero</td><td>River</td><td>Cook 1980</td></tr><tr><td>Miraxona (Miraxona) expansipes</td><td>Guerrero</td><td>River</td><td>Cook 1980</td></tr><tr><td>Miraxona (Miraxona) mexicana</td><td>Guerrero</td><td>River</td><td>Cook 1980</td></tr><tr><td>Mirazonides (Mirazonidella) similis</td><td>Nayarit, Veracruz</td><td>River</td><td>Cook 1980</td></tr><tr><td>Miraxona (Mirazonides) geronimoi</td><td>Nayarit</td><td>Stream</td><td>Cramer and Cook 1991</td></tr><tr><td>Stygalbiella mexicana</td><td>Mexico state</td><td>Spring</td><td>Cramer 1992</td></tr><tr><td>Neoaxona (?Lamellaxona) mexicana</td><td>Nayarit</td><td>Stream</td><td>Cook 1980</td></tr><tr><td>Woolastookia gretae</td><td>Nayarit</td><td>River, stream</td><td>Viets 1978 in Cook 1980</td></tr><tr><td>Forelia mesoamericana</td><td>Chiapas</td><td>Lagoon</td><td>Otero-Colina 1987b</td></tr><tr><td colspan="4"><strong>Feltriidae</strong></td></tr><tr><td>Feltria (Tropifeltria) neotropica</td><td>Chiapas</td><td>Waterfalls</td><td>Otero-Colina 1987b</td></tr><tr><td>Feltria (Feltriella) anahoffmannae</td><td>Mexico state</td><td>Mountain spring</td><td>Cramer 1986</td></tr><tr><td colspan="4"><strong>Mideopsidae</strong></td></tr><tr><td>Mideopsis (Mideopsis) cartesa</td><td>Nayarit, Veracruz</td><td>Stream</td><td>Cook 1980</td></tr><tr><td>Mideopsis (Mideopsides) beta</td><td>Chiapas</td><td>Gravel deposits of stream</td><td>Cook 1980</td></tr><tr><td>Mideopsis (Mideopsis) magna</td><td>Veracruz, Morelos</td><td>River</td><td>Cook 1980</td></tr><tr><td>Mideopsis (Neoxystonotus) nobilis</td><td>Tabasco</td><td>Pond</td><td>Cook 1980</td></tr><tr><td>Mideopsis orbicularis</td><td>Yucatán</td><td>Cenote</td><td>Müller 1776</td></tr><tr><td>Mideopsis (Neoxystonotus) mexicana</td><td>Mexico state</td><td>Helocrene spring</td><td>Cook 1980</td></tr><tr><td>Mideopsis (Neoxystonotus) gennada</td><td>Chiapas</td><td>River</td><td>Cook 1980</td></tr><tr><td>Mideopsis (Neoxystonotus) canzolla</td><td>Guerrero</td><td>Tributary river</td><td>Cook 1980</td></tr><tr><td colspan="4"><strong>Krendowskiidae</strong></td></tr><tr><td>Krendowskia (Krendowskiella) azteca</td><td>Guerrero</td><td>Tributary river</td><td>Cook 1980</td></tr><tr><td>Krendowskia (Krendowskiella) moyara</td><td>Chiapas</td><td>River</td><td>Cook 1980</td></tr><tr><td>Krendowskia (Krendowskiella) trodroma</td><td>Veracruz, Nayarit</td><td>Stream</td><td>Cook 1980</td></tr><tr><td>Krendowskia (Krendowskiella) vicina</td><td>Tabasco</td><td>Pond</td><td>Cook 1980</td></tr><tr><td>Geayia (Geayia) amacuzaca</td><td>Morelos</td><td>River</td><td>Cook 1980</td></tr><tr><td>Geayia (Geayia) mitchelli</td><td>Tabasco, Chiapas</td><td>Pond</td><td>Cook 1980</td></tr><tr><td colspan="4"><strong>Arenohydracaridae</strong></td></tr><tr><td>Arenohydracarus eremitus</td><td>*</td><td>*</td><td>Cook 1974</td></tr><tr><td>Arenohydracarus minimus</td><td>Oaxaca, Guerrero</td><td>Interstitial waters</td><td>Cook 1974</td></tr><tr><td colspan="4"><strong>Chappuisididae</strong></td></tr><tr><td>Chappuisides notialis</td><td>Mexico state</td><td>Spring</td><td>Cramer and Smith 1991</td></tr><tr><td colspan="4"><strong>Hungarohydracaridae</strong></td></tr><tr><td>Stygarrenurus armoniensis</td><td>Mexico state</td><td>Interstitial deposits of spring.</td><td>Cramer and Cook 1996</td></tr><tr><td colspan="4"><strong>Neoacaridae</strong></td></tr><tr><td>Neoacarus adocetus</td><td>Mexico state</td><td>Spring</td><td>Cramer and Smith 1991</td></tr></tbody></table>
<table><caption>Table 1 Continued.</caption><thead><tr><th>Species</th><th>State</th><th>Habitat</th><th>Reference</th></tr></thead><tbody><tr><td>Piona (Tetrapiona) rotunda sudamericana</td><td>*</td><td>*</td><td>Viets 1910 in Marín-Hernández and Cramer-Hemkes 2009</td></tr><tr><td>Piona alzatei</td><td>*</td><td>*</td><td>Dugés 1884</td></tr><tr><td>Piona pearsei</td><td>*</td><td>*</td><td>Marshall 1936</td></tr><tr><td>Piona amimitli</td><td>Mexico city</td><td>Stream</td><td>Marín-Hernández 2009</td></tr><tr><td><strong>Aturidae</strong></td><td></td><td></td><td></td></tr><tr><td>Aturus matlatzinca</td><td>Mexico state</td><td>Stream</td><td>Cramer 1991</td></tr><tr><td>Aturus catoantoni</td><td>Mexico state</td><td>Stream</td><td>Cramer 1991</td></tr><tr><td>Aturus guillecrameri</td><td>Mexico state</td><td>Stream</td><td>Cramer 1992</td></tr><tr><td>Aturus celiahemkesae</td><td>Mexico state</td><td>Stream</td><td>Cramer 1992</td></tr><tr><td>Aturus oxtoilpanensis</td><td>Mexico state</td><td>Stream</td><td>Cramer 1992</td></tr><tr><td>Aturus primitivus</td><td>Mexico state</td><td>Stream</td><td>Cramer 1991</td></tr><tr><td>Neoaturus kurtvietsi</td><td>Oaxaca, Chiapas, Veracruz, Guerrero</td><td>River, stream</td><td>Cook 1980</td></tr><tr><td>Neoaturus projectus</td><td>Veracruz</td><td>River</td><td>Lundblad 1936 in Cook 1980</td></tr><tr><td>Adelazonopsella pallida</td><td>Veracruz</td><td>Interstitial water stream</td><td>Cook 1974</td></tr><tr><td>Albia (Anchistalbia) ianthia</td><td>Nayarit</td><td>Spring</td><td>Cook 1980</td></tr><tr><td>Albia (Anchistalbia) ianthopsis</td><td>Quintana Roo</td><td>Lagoon</td><td>Otero-Colina 1987b</td></tr><tr><td>Axonopsella (?) mesoamericana</td><td>Guerrero, Morelos, Nayarit</td><td>River</td><td>Cook 1980</td></tr><tr><td>Axonopsella (Neoazonopsella) nayaritensis</td><td>Nayarit</td><td>Cataracts</td><td>Cook 1980</td></tr><tr><td>Axonopsella (Neoazonopsella) vicina</td><td>Veracruz</td><td>Stream</td><td>Cook 1980</td></tr><tr><td>Axonopsis (Brachypodopsis) columbica</td><td>Chiapas, Oaxaca, Puebla. Nayarit</td><td>River, stream</td><td>Lundblad 1953 in Cook 1980</td></tr><tr><td>Axonopsis (Brachypodopsis) cemoba</td><td>Veracruz</td><td>Stream</td><td>Cook 1980</td></tr><tr><td>Axonopsis (Brachypodopsis) cerucha</td><td>Veracruz, San Luis Potosí</td><td>River, Stream</td><td>Cook 1980</td></tr><tr><td>Axonopsis (Brachypodopsis) mesoamericana</td><td>Veracruz, Chiapas</td><td>Stream</td><td>Cook 1980</td></tr><tr><td>Axonopsis (Brachypodopsis) trocala</td><td>Chiapas</td><td>Chiapas</td><td>Cook 1980</td></tr><tr><td>Polyazonopsella mexicana</td><td>Oaxaca</td><td>Stream</td><td>Otero-Colina 1987b</td></tr><tr><td>Frontipodopsis mesoamericana</td><td>Chiapas</td><td>Bottom deposits of stream</td><td>Cook 1980</td></tr><tr><td>Kongsbergia antoniocatoi</td><td>Mexico state</td><td>Gravel deposits of stream</td><td>Cramer 2000</td></tr><tr><td>Kongsbergia cosita</td><td>Mexico state</td><td>Gravel deposits of stream</td><td>Cramer 2000</td></tr><tr><td>Kongsbergia tigrina</td><td>Mexico state</td><td>Stream</td><td>Cramer 2000</td></tr><tr><td>Kongsbergia eterna</td><td>Mexico state</td><td>Stream</td><td>Cramer 2000</td></tr><tr><td>Kongsbergia (Kongsbergia) mexicana</td><td>Oaxaca</td><td>River</td><td>Cook 1980</td></tr><tr><td>Kongsbergia glopipalpis</td><td>Chiapas and Veracruz</td><td>Stream</td><td>Lundblad 1953 in Cook 1980</td></tr><tr><td>Kongsbergia materna</td><td>Chiapas, Mexico state, Oaxaca, Veracruz.</td><td>Tributary river</td><td>Thor 1899 in Cramer 2000</td></tr><tr><td>Submiraxona (Submiraxona) bella</td><td>Nayarit</td><td>Stream</td><td>Cook 1980</td></tr><tr><td>Submiraxona (Submiraxona) lundbladi</td><td>Guerrero, Chiapas</td><td>River</td><td>Cook 1980</td></tr><tr><td>Submiraxona (Submiraxona) stolli</td><td>Guerrero</td><td>River</td><td>Cook 1980</td></tr><tr><td>Miraxona (Miraxona) expansipes</td><td>Guerrero</td><td>River</td><td>Cook 1980</td></tr><tr><td>Miraxona (Miraxona) mexicana</td><td>Guerrero</td><td>River</td><td>Cook 1980</td></tr><tr><td>Mirazonides (Mirazonidella) similis</td><td>Nayarit, Veracruz</td><td>River</td><td>Cook 1980</td></tr><tr><td>Miraxona (Mirazonides) geronimoi</td><td>Nayarit</td><td>Stream</td><td>Cramer and Cook 1991</td></tr><tr><td>Stygalbiella mexicana</td><td>Mexico state</td><td>Spring</td><td>Cramer 1992</td></tr><tr><td>Neoaxona (?Lamellaxona) mexicana</td><td>Nayarit</td><td>Stream</td><td>Cook 1980</td></tr><tr><td>Woolastookia gretae</td><td>Nayarit</td><td>River, stream</td><td>Viets 1978 in Cook 1980</td></tr><tr><td>Forelia mesoamericana</td><td>Chiapas</td><td>Lagoon</td><td>Otero-Colina 1987b</td></tr><tr><td><strong>Feltriidae</strong></td><td></td><td></td><td></td></tr><tr><td>Feltria (Tropifeltria) neotropica</td><td>Chiapas</td><td>Waterfalls</td><td>Otero-Colina 1987b</td></tr><tr><td>Feltria (Feltriella) anahoffmannae</td><td>Mexico state</td><td>Mountain spring</td><td>Cramer 1986</td></tr><tr><td><strong>Mideopsidae</strong></td><td></td><td></td><td></td></tr><tr><td>Mideopsis (Mideopsis) cartesa</td><td>Nayarit, Veracruz</td><td>Stream</td><td>Cook 1980</td></tr><tr><td>Mideopsis (Mideopsides) beta</td><td>Chiapas</td><td>Gravel deposits of stream</td><td>Cook 1980</td></tr><tr><td>Mideopsis (Mideopsis) magna</td><td>Veracruz, Morelos</td><td>River</td><td>Cook 1980</td></tr><tr><td>Mideopsis (Neoxystonotus) nobilis</td><td>Tabasco</td><td>Pond</td><td>Cook 1980</td></tr><tr><td>Mideopsis orbicularis</td><td>Yucatán</td><td>Cenote</td><td>Müller 1776</td></tr><tr><td>Mideopsis (Neoxystonotus) mexicana</td><td>Mexico state</td><td>Helocrene spring</td><td>Cook 1980</td></tr><tr><td>Mideopsis (Neoxystonotus) gennada</td><td>Chiapas</td><td>River</td><td>Cook 1980</td></tr><tr><td>Mideopsis (Neoxystonotus) canzolla</td><td>Guerrero</td><td>Tributary river</td><td>Cook 1980</td></tr><tr><td><strong>Krendowskiidae</strong></td><td></td><td></td><td></td></tr><tr><td>Krendowskia (Krendowskiella) azteca</td><td>Guerrero</td><td>Tributary river</td><td>Cook 1980</td></tr><tr><td>Krendowskia (Krendowskiella) moyara</td><td>Chiapas</td><td>River</td><td>Cook 1980</td></tr><tr><td>Krendowskia (Krendowskiella) trodroma</td><td>Veracruz, Nayarit</td><td>Stream</td><td>Cook 1980</td></tr><tr><td>Krendowskia (Krendowskiella) vicina</td><td>Tabasco</td><td>Pond</td><td>Cook 1980</td></tr><tr><td>Geayia (Geayia) amacuzaca</td><td>Morelos</td><td>River</td><td>Cook 1980</td></tr><tr><td>Geayia (Geayia) mitchelli</td><td>Tabasco, Chiapas</td><td>Pond</td><td>Cook 1980</td></tr><tr><td><strong>Arenohydracaridae</strong></td><td></td><td></td><td></td></tr><tr><td>Arenohydracarus eremitus</td><td>*</td><td>*</td><td>Cook 1974</td></tr><tr><td>Arenohydracarus minimus</td><td>Oaxaca, Guerrero</td><td>Interstitial waters</td><td>Cook 1974</td></tr><tr><td><strong>Chappuisididae</strong></td><td></td><td></td><td></td></tr><tr><td>Chappuisides notialis</td><td>Mexico state</td><td>Spring</td><td>Cramer and Smith 1991</td></tr><tr><td><strong>Hungarohydracaridae</strong></td><td></td><td></td><td></td></tr><tr><td>Stygarrenurus armoniensis</td><td>Mexico state</td><td>Interstitial deposits of spring.</td><td>Cramer and Cook 1996</td></tr><tr><td><strong>Neoacaridae</strong></td><td></td><td></td><td></td></tr><tr><td>Neoacarus adocetus</td><td>Mexico state</td><td>Spring</td><td>Cramer and Smith 1991</td></tr></tbody></table>
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

<table><caption>**Table 2** Unidentified species with a DNA barcode in BOLD database. BINs indicate putative species. * Unique BINs this database.</caption><thead><tr><th>Taxa</th><th>BIN</th><th>Taxa</th><th>BIN</th><th>Taxa</th><th>BIN</th></tr></thead><tbody><tr><td><strong>Arrenuridae</strong></td><td></td><td><strong>Limnesiidae</strong></td><td></td><td><strong>Pionidae</strong></td><td></td></tr><tr><td>Arrenuridae</td><td>AEA4828*</td><td>Limnesiidae</td><td>AEA4382*</td><td>Pionidae</td><td>AEA4809*</td></tr><tr><td>Arrenurus cristinae</td><td>AEA7842*</td><td>Centrolimnesia</td><td>AEA3914*</td><td>Piona</td><td>AEA5358*</td></tr><tr><td>Arrenurus eduardoi</td><td>AEA7844*</td><td>Limnesia</td><td>AEA6471*</td><td>Piona</td><td>AEES501*</td></tr><tr><td>Arrenurus federicoi</td><td>AEB7095*</td><td>Limnesia</td><td>ACX7759</td><td>Piona</td><td>AEN6046*</td></tr><tr><td>Arrenurus ecosur</td><td>ACX8463</td><td>Limnesia</td><td>ACY7380</td><td>Piona</td><td>AEO7290*</td></tr><tr><td>Arrenurus marshallae</td><td>ACL2521</td><td>Limnesia</td><td>AEA5595</td><td>Piona</td><td>AER1599*</td></tr><tr><td>Arrenurus</td><td>ACX8462*</td><td><strong>Hygrobatidae</strong></td><td></td><td>Piona</td><td>AER1600*</td></tr><tr><td>Arrenurus</td><td>ACX8788*</td><td>Hygrobatidae</td><td>AEA5236*</td><td>Piona</td><td>AER1601*</td></tr><tr><td>Arrenurus</td><td>ACX8789*</td><td>Hygrobatidae</td><td>AEA4089*</td><td><strong>Mideopsidae</strong></td><td></td></tr><tr><td>Arrenurus</td><td>ACY6809*</td><td>Atractides</td><td>ACX7786*</td><td>Mideopsidae</td><td>AEB4633*</td></tr><tr><td>Arrenurus</td><td>ADI3752*</td><td>Hygrobates</td><td>AEA3689*</td><td>Mideopsis</td><td>ACY7169*</td></tr><tr><td>Arrenurus</td><td>ADI4458*</td><td>Hygrobates</td><td>AEA3690*</td><td>Mideopsis</td><td>AEA6512*</td></tr><tr><td>Arrenurus</td><td>AEA3972*</td><td>Hygrobates</td><td>AEA3924*</td><td>Mideopsis</td><td>ACX8679</td></tr><tr><td>Arrenurus</td><td>AEA7182*</td><td>Hygrobates</td><td>ACX7887</td><td><strong>Krendowskiidae</strong></td><td></td></tr><tr><td>Arrenurus</td><td>AEA7843*</td><td>Hygrobates</td><td>ADO7098</td><td>Krendowskia</td><td>ACX8435*</td></tr><tr><td>Arrenurus</td><td>AEA8234*</td><td><strong>Unionicolidae</strong></td><td></td><td>Geayia</td><td>ACT6195</td></tr><tr><td>Arrenurus</td><td>AEF1989*</td><td>Unionicolidae</td><td>AEA6658*</td><td><strong>Hydrachnidia</strong></td><td></td></tr><tr><td>Arrenurus</td><td>AEF8444*</td><td>Unionicolidae</td><td>ACY7381</td><td></td><td>AEA3823*</td></tr><tr><td>Arrenurus</td><td>ACL2418</td><td>Unionicolidae</td><td>AEB1594*</td><td></td><td>AEA4343*</td></tr><tr><td>Arrenurus</td><td>ACX8464</td><td>Unionicolidae</td><td>AEA7951*</td><td></td><td>AEF3494*</td></tr><tr><td><strong>Limnocharidae</strong></td><td></td><td>Unionicolidae</td><td>AEA6062*</td><td></td><td>AEF8255*</td></tr><tr><td>Limnocharidae</td><td>AEA4515*</td><td>Unionicolidae</td><td>AEA4829*</td><td></td><td>AEF0324*</td></tr><tr><td>Limnocharides</td><td>ACY6840*</td><td>Unionicolidae</td><td>AEA3726*</td><td></td><td>AEI2293*</td></tr><tr><td>Limnocharides</td><td>ADI4862*</td><td>Unionicolidae</td><td>AEA4514*</td><td></td><td>AEI2954*</td></tr><tr><td>Eylaidae</td><td></td><td>Unionicola</td><td>ACX8035*</td><td></td><td>AEI4296*</td></tr><tr><td>Eylaidae</td><td>AEA4696*</td><td>Unionicola</td><td>ADM7936*</td><td></td><td>AEI4327*</td></tr><tr><td>Eylaidae</td><td>AEA5669*</td><td>Unionicola</td><td>AEB4634*</td><td></td><td>AEI4328*</td></tr><tr><td>Eylais</td><td>ADD9174*</td><td>Unionicola</td><td>AEE0841*</td><td></td><td>AEI4329*</td></tr><tr><td><strong>Hydryptantidae</strong></td><td></td><td>Unionicola</td><td>ACX9008</td><td></td><td>AEI8266*</td></tr><tr><td>Hydryptantes</td><td>AEA5005*</td><td>Unionicola</td><td>AEF2345</td><td></td><td>AEI8707*</td></tr><tr><td><strong>Hydrodromidae</strong></td><td></td><td>Unionicola</td><td>ACX8034</td><td></td><td>AEI9124*</td></tr><tr><td>Hydrodroma</td><td>ADF3732</td><td>Neumania</td><td>AEA8101*</td><td></td><td>AEN6047*</td></tr><tr><td><strong>Anisitsiellidae</strong></td><td></td><td>Neumania</td><td>ACY6829</td><td></td><td>AEN8226*</td></tr><tr><td>Mamersellides</td><td>AEA6955*</td><td>Koenikea</td><td>ACY7384*</td><td></td><td>AEN8227*</td></tr><tr><td>Mamersellides</td><td>AEA6956*</td><td>Koenikea</td><td>ACB9299</td><td></td><td>AE05260*</td></tr><tr><td>Torrenticolidae</td><td></td><td>Koenikea</td><td>ADI2928</td><td></td><td>AER2566*</td></tr><tr><td>Torrenticolidae</td><td>AEA4395*</td><td>Koenikea</td><td>ADI3114</td><td></td><td>AEB1898</td></tr><tr><td>Torrenticola</td><td>AEA7372*</td><td></td><td></td><td></td><td></td></tr></tbody></table>
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

<table><caption>**Table 2** Unidentified species with a DNA barcode in BOLD database. BINs indicate putative species. * Unique BINs this database.</caption><thead><tr><th>Taxa</th><th>BIN</th><th>Taxa</th><th>BIN</th><th>Taxa</th><th>BIN</th></tr></thead><tbody><tr><td><strong>Arrenuridae</strong></td><td></td><td><strong>Limnesiidae</strong></td><td></td><td><strong>Pionidae</strong></td><td></td></tr><tr><td>Arrenuridae</td><td>AEA4828*</td><td>Limnesiidae</td><td>AEA4382*</td><td>Pionidae</td><td>AEA4809*</td></tr><tr><td>Arrenurus cristinae</td><td>AEA7842*</td><td>Centrolimnesia</td><td>AEA3914*</td><td>Piona</td><td>AEA5358*</td></tr><tr><td>Arrenurus eduardoi</td><td>AEA7844*</td><td>Limnesia</td><td>AEA6471*</td><td>Piona</td><td>AEES501*</td></tr><tr><td>Arrenurus federicoi</td><td>AEB7095*</td><td>Limnesia</td><td>ACX7759</td><td>Piona</td><td>AEN6046*</td></tr><tr><td>Arrenurus ecosur</td><td>ACX8463</td><td>Limnesia</td><td>ACY7380</td><td>Piona</td><td>AEO7290*</td></tr><tr><td>Arrenurus marshallae</td><td>ACL2521</td><td>Limnesia</td><td>AEA5595</td><td>Piona</td><td>AER1599*</td></tr><tr><td>Arrenurus</td><td>ACX8462*</td><td><strong>Hygrobatidae</strong></td><td></td><td>Piona</td><td>AER1600*</td></tr><tr><td>Arrenurus</td><td>ACX8788*</td><td>Hygrobatidae</td><td>AEA5236*</td><td>Piona</td><td>AER1601*</td></tr><tr><td>Arrenurus</td><td>ACX8789*</td><td>Hygrobatidae</td><td>AEA4089*</td><td><strong>Mideopsidae</strong></td><td></td></tr><tr><td>Arrenurus</td><td>ACY6809*</td><td>Atractides</td><td>ACX7786*</td><td>Mideopsidae</td><td>AEB4633*</td></tr><tr><td>Arrenurus</td><td>ADI3752*</td><td>Hygrobates</td><td>AEA3689*</td><td>Mideopsis</td><td>ACY7169*</td></tr><tr><td>Arrenurus</td><td>ADI4458*</td><td>Hygrobates</td><td>AEA3690*</td><td>Mideopsis</td><td>AEA6512*</td></tr><tr><td>Arrenurus</td><td>AEA3972*</td><td>Hygrobates</td><td>AEA3924*</td><td>Mideopsis</td><td>ACX8679</td></tr><tr><td>Arrenurus</td><td>AEA7182*</td><td>Hygrobates</td><td>ACX7887</td><td><strong>Krendowskiidae</strong></td><td></td></tr><tr><td>Arrenurus</td><td>AEA7843*</td><td>Hygrobates</td><td>ADO7098</td><td>Krendowskia</td><td>ACX8435*</td></tr><tr><td>Arrenurus</td><td>AEA8234*</td><td><strong>Unionicolidae</strong></td><td></td><td>Geayia</td><td>ACT6195</td></tr><tr><td>Arrenurus</td><td>AEF1989*</td><td>Unionicolidae</td><td>AEA6658*</td><td><strong>Hydrachnidia</strong></td><td></td></tr><tr><td>Arrenurus</td><td>AEF8444*</td><td>Unionicolidae</td><td>ACY7381</td><td></td><td>AEA3823*</td></tr><tr><td>Arrenurus</td><td>ACL2418</td><td>Unionicolidae</td><td>AEB1594*</td><td></td><td>AEA4343*</td></tr><tr><td>Arrenurus</td><td>ACX8464</td><td>Unionicolidae</td><td>AEA7951*</td><td></td><td>AEF3494*</td></tr><tr><td><strong>Limnocharidae</strong></td><td></td><td>Unionicolidae</td><td>AEA6062*</td><td></td><td>AEF8255*</td></tr><tr><td>Limnocharidae</td><td>AEA4515*</td><td>Unionicolidae</td><td>AEA4829*</td><td></td><td>AEF0324*</td></tr><tr><td>Limnocharides</td><td>ACY6840*</td><td>Unionicolidae</td><td>AEA3726*</td><td></td><td>AEI2293*</td></tr><tr><td>Limnocharides</td><td>ADI4862*</td><td>Unionicolidae</td><td>AEA4514*</td><td></td><td>AEI2954*</td></tr><tr><td>Eylaidae</td><td></td><td>Unionicola</td><td>ACX8035*</td><td></td><td>AEI4296*</td></tr><tr><td>Eylaidae</td><td>AEA4696*</td><td>Unionicola</td><td>ADM7936*</td><td></td><td>AEI4327*</td></tr><tr><td>Eylaidae</td><td>AEA5669*</td><td>Unionicola</td><td>AEB4634*</td><td></td><td>AEI4328*</td></tr><tr><td>Eylais</td><td>ADD9174*</td><td>Unionicola</td><td>AEE0841*</td><td></td><td>AEI4329*</td></tr><tr><td><strong>Hydryptantidae</strong></td><td></td><td>Unionicola</td><td>ACX9008</td><td></td><td>AEI8266*</td></tr><tr><td>Hydryptantes</td><td>AEA5005*</td><td>Unionicola</td><td>AEF2345</td><td></td><td>AEI8707*</td></tr><tr><td><strong>Hydrodromidae</strong></td><td></td><td>Unionicola</td><td>ACX8034</td><td></td><td>AEI9124*</td></tr><tr><td>Hydrodroma</td><td>ADF3732</td><td>Neumania</td><td>AEA8101*</td><td></td><td>AEN6047*</td></tr><tr><td><strong>Anisitsiellidae</strong></td><td></td><td>Neumania</td><td>ACY6829</td><td></td><td>AEN8226*</td></tr><tr><td>Mamersellides</td><td>AEA6955*</td><td>Koenikea</td><td>ACY7384*</td><td></td><td>AEN8227*</td></tr><tr><td>Mamersellides</td><td>AEA6956*</td><td>Koenikea</td><td>ACB9299</td><td></td><td>AE05260*</td></tr><tr><td>Torrenticolidae</td><td></td><td>Koenikea</td><td>ADI2928</td><td></td><td>AER2566*</td></tr><tr><td>Torrenticolidae</td><td>AEA4395*</td><td>Koenikea</td><td>ADI3114</td><td></td><td>AEB1898</td></tr><tr><td>Torrenticola</td><td>AEA7372*</td><td></td><td></td><td></td><td></td></tr></tbody></table>
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

<figure><img src="image_4.png" /><figcaption>**Figure 3** Neighbor joining tree based on all worldwide *Neumania* COI sequences. The name is followed by the barcode index number (BIN). * Without data collection.</figcaption></figure>
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

<figure><img src="image_5.png" /><figcaption>**Figure 3** Neighbor joining tree based on all worldwide *Neumania* COI sequences. The name is followed by the barcode index number (BIN). * Without data collection.</figcaption></figure>
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
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 signifi-
cantly improve this manuscript.

ORCID

Lucia Montes-Ortiz https://orcid.org/0000-0001-5959-7188

Sergio Cohuo https://orcid.org/0000-0002-7826-5303

Manuel Elías-Gutiérrez https://orcid.org/0000-0003-2562-4584

References

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 (Micru-racarus) novus*. Life, 10 (7): 108. https://doi.org/10.3390/life10070108

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

Cook, D. R. 1980. Studies on neotropical water mites. Memoirs of the American Entomological Institute, 31: 645.

Cramer, C. 1987. Nuevas especies mexicanas de hidracaridos, pertenecientes a los géneros *Neomamersa*, *Kawamuracarus* y *Neotorrenticola* (Acarida: Limnesiidae). Folia Entomológica Mexicana, 73: 185-203.

Cramer, C. 1988. Ácaros dulceacuícolas (Acarida: Prostigmata) del Arroyo Peña Blanca en San Francisco Oxtotilpan, México (Doctoral dissertation, Universidad Nacional Autónoma de México), 269.

Cramer, C. 1991. Estudios sobre hidracáridos mexicanos. Especies nuevas de la familia Aturidae. Parte 1. Folia Entomológica Mexicana, 82: 173-186.

Cramer, C. 1992a. Estudios sobre hidracáridos mexicanos, especies nuevas de la familia Aturidae. Parte II (*Aturus: Stygalbiella*). Folia Entomológica Mexicana, 84: 113-129.

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.

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

Cramer, C., Cook, D. R. 1992a. New species of *Arrenurus* (Acari: Arrenuridae) from Mexican lakes. Acarologia, 33 (4): 349-366.

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

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

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

Cramer, C., Cook, D. R. 2000. Water mites of the genera *Neoatractides* Lundblad and *Pseudotorrenticola* Walter (Acari: Hydrachnidia: Torrenticolidae) from Mexico. International Journal of Acarology, 26 (1): 51-61. https://doi.org/10.1080/01647950008683635

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.

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

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

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

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.

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
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 signifi-
cantly improve this manuscript.

ORCID

Lucia Montes-Ortiz https://orcid.org/0000-0001-5959-7188

Sergio Cohuo https://orcid.org/0000-0002-7826-5303

Manuel Elías-Gutiérrez https://orcid.org/0000-0003-2562-4584

References

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 (Micru-racarus) novus*. Life, 10 (7): 108. https://doi.org/10.3390/life10070108

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

Cook, D. R. 1980. Studies on neotropical water mites. Memoirs of the American Entomological Institute, 31: 645.

Cramer, C. 1987. Nuevas especies mexicanas de hidracaridos, pertenecientes a los géneros *Neomamersa*, *Kawamuracarus* y *Neotorrenticola* (Acarida: Limnesiidae). Folia Entomológica Mexicana, 73: 185-203.

Cramer, C. 1988. Ácaros dulceacuícolas (Acarida: Prostigmata) del Arroyo Peña Blanca en San Francisco Oxtotilpan, México (Doctoral dissertation, Universidad Nacional Autónoma de México), 269.

Cramer, C. 1991. Estudios sobre hidracáridos mexicanos. Especies nuevas de la familia Aturidae. Parte 1. Folia Entomológica Mexicana, 82: 173-186.

Cramer, C. 1992a. Estudios sobre hidracáridos mexicanos, especies nuevas de la familia Aturidae. Parte II (*Aturus: Stygalbiella*). Folia Entomológica Mexicana, 84: 113-129.

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.

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

Cramer, C., Cook, D. R. 1992a. New species of *Arrenurus* (Acari: Arrenuridae) from Mexican lakes. Acarologia, 33 (4): 349-366.

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

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

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

Cramer, C., Cook, D. R. 2000. Water mites of the genera *Neoatractides* Lundblad and *Pseudotorrenticola* Walter (Acari: Hydrachnidia: Torrenticolidae) from Mexico. International Journal of Acarology, 26 (1): 51-61. https://doi.org/10.1080/01647950008683635

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.

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

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

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

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.

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
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

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

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.

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

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

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

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

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

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

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.

Otero-Colina, G. 1987b. Nuevas especies de ácaros acuáticos (Prostigmata: Parasitengona) del sureste de México. Folia Entomológica Mexicana, 72: 89-109.

Otero-Colina, G. 1988. Datos originales sobre los ácaros acuáticos (Prostigmata: Parasitengona) del sureste de México. Folia Entomológica Mexicana, 76: 195-223.

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

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

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

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 Invertebrates, Academic Press, pp. 599-660. https://doi.org/10.1016/B978-0-12-385026-3.00025-5

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

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.

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

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. https://doi.org/10.1007/s10493-020-00476-4

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

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.

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

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

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
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

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

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.

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

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

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

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

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

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

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.

Otero-Colina, G. 1987b. Nuevas especies de ácaros acuáticos (Prostigmata: Parasitengona) del sureste de México. Folia Entomológica Mexicana, 72: 89-109.

Otero-Colina, G. 1988. Datos originales sobre los ácaros acuáticos (Prostigmata: Parasitengona) del sureste de México. Folia Entomológica Mexicana, 76: 195-223.

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

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

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

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 Invertebrides, Academic Press, pp. 599-660. https://doi.org/10.1016/B978-0-12-385026-3.00025-5

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

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.

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

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. https://doi.org/10.1007/s10493-020-00476-4

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

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.

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

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

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
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

Vidrine, M. F. 1994. Revision of the subgenus *Polyatax* (Acari: Unionicolidae: Unionicolinae: *Unioni-cola*). Texas Journal of Science, 46 (2): 157-163.

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

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
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

Vidrine, M. F. 1994. Revision of the subgenus *Polyatax* (Acari: Unionicolidae: Unionicolinae: *Unioni-cola*). Texas Journal of Science, 46 (2): 157-163.

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

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