# New species of a fur mite of the genus Alabidocarpus (Acari: Chirodiscidae) associated with Desmodus rotundus (Chiroptera: Phyllostomidae) from Mexico, including a key to the American species

Juan B Morales-Malacara, Naomi Vázquez-Xicoténcatl, Carlos García-Estrada, Helisama Colín-Martínez, Cenia E Almazán-Marín, Hugo A Valdes-Saucedo, Hugo A Ruiz-Piña, Alan Cuxim-Koyoc, Enrique Reyes-Novelo

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Juan B Morales-Malacara, Naomi Vázquez-Xicoténcatl, Carlos García-Estrada, Helisama Colín-Martínez, Cenia E Almazán-Marín, et al. New species of a fur mite of the genus Alabidocarpus (Acari: Chirodiscidae) associated with Desmodus rotundus (Chiroptera: Phyllostomidae) from Mexico, including a key to the American species. Acarologia, 2025, 65 (3), pp.736-755. 10.24349/235z-kw78. hal-05199142

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# New species of a fur mite of the genus *Alabidocarpus* (Acari: Chirodiscidae) associated with *Desmodus rotundus* (Chiroptera: Phyllostomidae) from Mexico, including a key to the American species

Juan B. Morales-Malacara IDª, Naomi Vázquez-Xicoténcatl IDª, Carlos

García-Estrada IDᵇ, Helisama Colín-Martínez IDᵇ, Cenia E.

Almazán-Marín IDª, Hugo A. Valdes-Saucedo IDª, Hugo A. Ruiz-Piña IDᶜ,

Alan Cuxim-Koyoc IDᶜ, Enrique Reyes-Novelo IDᶜ

## Original research

### ABSTRACT

Derived from an extensive survey on the common vampire bat *Desmodus rotundus* (Geoffroy, 1810) in some localities along the Central region, and the Pacific and Atlantic slopes of Mexico, Chirodiscid fur mites of the genus *Alabidocarpus* were found exclusively in the country's southern region in the state of Oaxaca, where they turned out to be a new species. Here, we describe the adults and juveniles of this new Chirodiscid species. The present study increases the number of species of *Alabidocarpus* associated with American bats to seven; therefore, we include an updated key to known species in the American continent.

**Keywords** Chirodiscidae; *Alabidocarpus*; bat ectoparasite; *Desmodus*

**Zoobank** http://zoobank.org/CCBBD646-AF2E-481F-9E81-B42AD4A3EE43

### Introduction

The fur mites of the family Chirodiscidae Trouessart, 1892, currently has 228 valid species, divided into four subfamilies associated with mammals: Chirodiscinae Trouessart, 1892 (1 species) on Marsupialia, Schizocoptinae Fain, 1970 (1 species) on Afrosoricida, Lemuroeciinae Fain, 1968 (1 species) on Primates, and Labidocarpinae Gunther, 1942 which has the greatest diversity. The latter comprises the tribe Schizocarpini Fain, 1971, which includes two species associated with Eulipotyphla, five species with Carnivora, 49 species with Rodentia, and the tribe Labidocarpini Fain, 1971 contains one species associated with Primates, and 168 species represented by 20 genera found on Chiroptera with a wide distribution in both Old and New World (Bochkov 2010).

In particular, the subfamily Labidocarpinae Gunther, 1942, consists of permanent ectosymbionts that have specific adaptations to exploit their food resources at the base of each hair

Received 29 January 2025

Accepted 17 July 2025

Published 22 July 2025

Corresponding author

Juan B. Morales-Malacara ID: morales.malacara@gmail.com

Academic editor

Faraji, Farid

https://doi.org/10.24349/235z-kw78

ISSN 0044-586X (print)

ISSN 2107-7207 (electronic)

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

Morales-Malacara J. B. et al.

Licensed under

Creative Commons CC-BY 4.0

<footnote>ªLaboratorio de Espeleobiología y Acarología, Unidad Multidisciplinaria de Docencia e Investigación, Facultad de Ciencias, Universidad Nacional Autónoma de México, campus Juriquilla, Boulevard Juriquilla 3001, C.P. 76230, Querétaro, México.</footnote>

<footnote>ᵇUniversidad del Mar. Instituto de Ecología. Km 1.5 de la carretera a Sola de Vega. C.P. 71980. Puerto Escondido, San Pedro Mixtepec, Juquila, Oaxaca, México.</footnote>

<footnote>ᶜCentro de Investigaciones Regionales "Dr. Hideyo Noguchi" Universidad Autónoma de Yucatán. Av. Itzaes por 59 No. 490 Centro, C.P. 97000, Mérida, Yucatán, México.</footnote>
follicle. These mites exhibit an elongated, subcylindrical or laterally compressed idiosoma with reduced leg segments or extreme fusion, particularly in legs I and II. A broad fan-like tarsus represents the free segment. Throughout their life cycle, chirodiscids exhibit viviparity in gravid females. These females give birth to larvae with two developmental lines: the female and the male. These larvae then molt into nymphs and eventually to adults (Fain 1971).

*Alabidocarpus* Ewing, 1929 currently includes 37 species found on bats from the Old World belonging to the families Rhinolophidae, Hipposideridae, Nycteridae, Megadermatidae, Molossidae, and Pteropodidae. Additionally, species from the families Emballonuridae and Vespertilionidae are found in both the Old and New Worlds, while those from the family Phyllostomidae are endemic to the New World (Bochkov 2010).

Twelve species of *Alabidocarpus* have been recorded in the American continent. However, it has been shown that six species of these records contain errors, misidentifications, undetermined species, or generic reassignment of some species:

1) *Alabidocarpus calcaratus* Lawrence, 1952, was described from specimens obtained from *Myotis tricolor* (Temminck, 1832) in South Africa (Lawrence 1952). Later, *A. longipilus* Pinichpongse, 1963 was described from specimens obtained from *M. yumanensis satu-ratus* Miller, 1897 from California, USA. Pinichpongse (1963) himself noted some mor-phological differences between the two species. However, later McDaniel (1970), after obtaining specimens of *M. nigricans* (Schninz, 1821) from Nicaragua, conducted a morpho-logical analysis of both species and concluded that one of the main similarities between them, the pair of spur-like setae on coxa IV, was enough to synonymize *A. longipilus* with *A. calcaratus*. Later, Fain (1973) argued that McDaniel's proposed synonymy might be plausible but suggested reviewing the type host specimens before making a final decision. Therefore, Fain (1973) questioned the synonymy, marking it with a question mark, which was seconded by Guerrero (1992). Then, the latter author concluded that *A. longipilus* should be revalidated, and this was also recognized by Bochkov (2010). We agree with these arguments, considering *A. calcaratus* a valid species, restricted to Africa, Eurasia, and Europe (Bochkov 2010), and *A. longipilus* distributed across the American continent (Bochkov 2010).

2) *Alabidocarpus eptesicus* Fain, 1970 was described from Congo, Africa (Fain, 1970a) on *Eptesicus tenuipinnis* (Peters, 1872) (currently *Pseudoromicia tenuipinnis* (Peters, 1872), *sensu* Monadjem *et al.* 2020), with some other records from bats in Europe (Bochkov 2010). However, records from the American continent have been cited as *A. nr. eptesi-cus*, indicating undetermined or undescribed species, from the vespertilionid bats *Eptesi-cus fuscus* (Beauvois, 1796) in Colorado, USA (Pearce and O'Shea 2007, Whitaker *et al.* 2007), and *Myotis occultus* Hollister, 1909 from Colorado and New Mexico, USA (Whitaker *et al.* 2007, Valdez *et al.* 2009). These specimens should be recognized as *Alabidocarpus* sp. associated with these bats distributed in America.

3) *Alabidocarpus americanus* (McDaniel and Lawrence, 1964), from Cuba (Torre and Cuervo 2019), was reclassified based on a generic change, but there is no taxonomic justification for the reassignment of this species to the genus *Alabidocarpus*; besides, this species was first cited from Cuba by Cuervo *et al.* (1995) as *Olabidocarpus americanus* McDaniel and Lawrence, 1964 following the original description made by McDaniel and Lawrence (1964) in the genus *Olabidocarpus*.

The other three species in the Neotropical region in America, initially described within the genus *Alabidocarpus*, were reassigned to the genus *Pseudoalabidocarpus* McDaniel, 1972 due to morphological differences defined by Guerrero (1992) and corroborated by Bochkov (2010). Therefore, original records no longer belong to the genus *Alabidocarpus*:

4) *Alabidocarpus macrotus* Cruz, 1974 associated with *Macrotus waterhousii minor* Gundlach, 1864 from Habana (Cruz 1974) and reported from Cuba by Cuervo *et al.* (1995), was correctly reconsidered as *Pseudoalabidocarpus macrotus* (Cruz, 1974) by Torre and Cuervo (2019).
5) *Alabidocarpus noctilio* Fain, 1970, associated with *Noctilio labialis* (Kerr, 1792) from Suriname (Fain 1970b; 1973), is now *Pseudoalabidocarpus noctilio* (Fain, 1970) (Bochkov 2010).

6) *Alabidocarpus phyllostomi* Fain, 1972 associated with *Phyllostomus hastatus* (Pallas, 1767) from Suriname (Fain 1972; 1973) is now considered as *Pseudoalabidocarpus phyllostomi* (Fain, 1972) (Bochkov 2010).

Therefore, this would lead to six species for the genus *Alabidocarpus* distributed in the American continent (Table 1): *A. furmani* Pinichpongse, 1963; *A. guyanensis* Fain, 1972; *A. jonesi* McDaniel, 1971; *A. longipilus* Pinichpongse, 1963; *A. nicaraguae* McDaniel, 1971; and *A. saccopteryx* Fain, 1970 (Pinichpongse 1963; Fain 1970b, 1972; McDaniel 1970; McDaniel and Coffman 1970; Guerrero 1992; Whitaker and Morales-Malacara 2005, Bochkov 2010).

After surveying common vampire bats *Desmodus rotundus* (Geoffroy, 1810) in different localities across Central, Pacific and Atlantic slopes of Mexico, we focused on searching for Chirodiscid fur mites, and we found mites of the genus *Alabidocarpus* only in southern Mexico (Pacific slope), in the state of Oaxaca, which represents a new species.

This study aims to describe all active stages of a new species of *Alabidocarpus* associated with the common vampire bat *Desmodus rotundus*; which is the 38th species described in the genus, the 7th species in the American continent, and the third species found in Mexico. Furthermore, a rediagnosis of the genus is presented, accompanied by an updated key for American species.

<table><caption>**Table 1** Host and location data for known species of *Alabidocarpus* from America continent.</caption><thead><tr><th>Species</th><th>Host</th><th>Location</th><th>Reference</th></tr></thead><tbody><tr><td rowspan="11">A. furmani</td><td>Anoura geoffroyi</td><td>Trinidad: Mount Tamana<br/>Nicaragua: N. Segovia<br/>México: Guerrero</td><td>Pinichpongse 1963; Fain 1973; Webb and Loomis 1977; Guerrero 1992<br/>McDaniel 1970; Webb and Loomis 1977</td></tr><tr><td>Glossophaga soricina</td><td>Nicaragua: Chontales</td><td>Morales-Malacara 1982; Hoffmann et al. 1986; Whitaker and Morales-Malacara 2005</td></tr><tr><td>Anoura caudifer</td><td>Venezuela: T.F. Amazonas</td><td>McDaniel 1970; Webb and Loomis 1977</td></tr><tr><td>Glossophaga longirostris</td><td>Venezuela: Puerto Paez</td><td>McDaniel 1972; Webb and Loomis 1977; Guerrero 1992; Smith 2012</td></tr><tr><td>Carollia perspicillata</td><td>Venezuela: T.F. Amazonas<br/>Surinam: Moeroekreek</td><td>McDaniel 1972; Webb and Loomis 1977; Guerrero 1992<br/>Fain 1973</td></tr><tr><td>Carollia brevicauda</td><td>French Guyana: Cayenne</td><td>Fain 1973</td></tr><tr><td>Platyrhinus helleri</td><td>Venezuela: Carabobo</td><td>McDaniel 1972; Webb and Loomis 1977; Guerrero 1992</td></tr><tr><td>Pteronotus parnellii</td><td>Venezuela: T.F. Amazonas</td><td>McDaniel 1972<sup>a</sup>; Webb and Loomis 1977<sup>a</sup>; Guerrero 1992<sup>a</sup></td></tr><tr><td>Artibeus cinereus</td><td>Venezuela: Yaracuy</td><td>McDaniel 1972; Guerrero 1992</td></tr><tr><td>Platyrhinus helleri</td><td>French Guyana: Rochambeau</td><td>Fain 1972, 1973; Webb and Loomis 1977; Guerrero 1992</td></tr><tr><td>A. guyanensis</td><td>Artibeus cinereus</td><td>Nicaragua: Chontales</td><td>McDaniel 1970<sup>a</sup>, 1972<sup>a</sup>; Fain 1973; Webb and Loomis 1977<sup>a</sup>; Guerrero 1992<sup>a</sup></td></tr><tr><td>A. jonesi</td><td>Platyrhinus helleri</td><td>Venezuela: T.F. Amazonas</td><td>McDaniel 1972<sup>a</sup>; Webb and Loomis 1977<sup>a</sup>; Guerrero 1992<sup>a</sup></td></tr><tr><td rowspan="10">A. longipilus</td><td>Myotis yumanensis</td><td>USA: California</td><td>Pinichpongse 1963; McDaniel and Coffman 1970<sup>b</sup>; Fain 1973<sup>b</sup>; Guerrero 1992; Braun et al. 2015</td></tr><tr><td>Myotis nigricans</td><td>Nicaragua: N. Segobia, Chontales</td><td>McDaniel 1970<sup>b</sup>; McDaniel and Coffman 1970<sup>b</sup>; Fain 1973<sup>b</sup>; Guerrero 1992</td></tr><tr><td>Myotis lucifugus</td><td>USA: Michigan</td><td>Dood and Kurta 1982<sup>b</sup></td></tr><tr><td>Myotis thysanodes</td><td>México: Nuevo León</td><td>Bassols Batalla et al. 1996<sup>b</sup>; Whitaker and Morales-Malacara 2005<sup>b</sup></td></tr><tr><td>Myotis volans</td><td>USA: Oregon</td><td>Fain 1982<sup>b</sup>; Whitaker et al. 2007<sup>b</sup></td></tr><tr><td>Myotis californicus</td><td>USA: Colorado</td><td>Fain 1982<sup>b</sup>; Whitaker et al. 2007<sup>b</sup></td></tr><tr><td>Myotis evotis</td><td>USA: Oregon</td><td>Whitaker et al. 1983<sup>b</sup>; Parson et al. 1991<sup>b</sup></td></tr><tr><td>Myotis occultus</td><td>USA: Colorado, New Mexico</td><td>Whitaker et al. 2007<sup>b</sup>; Valdez et al. 2009<sup>b</sup></td></tr><tr><td>A. nicaraguae</td><td>Uroderma bilobatum</td><td>Nicaragua: Boaco</td><td>McDaniel 1970, 1972; Fain 1973; Webb and Loomis 1977; Guerrero 1992</td></tr><tr><td>A. nicaraguae</td><td>Uroderma magnirostrum</td><td>Venezuela: T.F. Amazonas</td><td>McDaniel 1972; Webb and Loomis 1977; Guerrero 1992</td></tr><tr><td>A. saccopteryx</td><td>Saccopteryx bilineata</td><td>Suriname: Lelydorp<br/>Suriname: Sumatraweg</td><td>Fain 1970b; Fain 1973; Guerrero 1992<br/>Fain 1973; Guerrero 1992</td></tr><tr><td rowspan="2">Alabidocarpus sp.</td><td>Eptesicus fuscus</td><td>USA: Colorado</td><td>Pearce and O'Shea 2007<sup>c</sup>; Whitaker et al. 2007<sup>c</sup></td></tr><tr><td>Myotis occultus</td><td>USA: Colorado, New Mexico</td><td>Whitaker et al. 2007<sup>c</sup>; Valdez et al. 2009<sup>c</sup></td></tr></tbody></table>

<footnote>Notes<sup>a</sup> Host cited as *Vampyrops helleri*; <sup>b</sup> Parasite misidentified and cited as *Alabidocarpus calcaratus*; <sup>c</sup> Parasite misidentified and cited as *Alabidocarpus* near *eptesicus* or *Alabidocarpus eptesicus* [Alabidocarpus *eptesicus* it is a valid species, but its distribution is restricted to Africa and Europe (Bochkov 2010)].</footnote>
<figure><img src="image_3.png" /><figcaption>**Figure 1** Geographical distribution of sampling localities of *Desmodus rotundus*. Specimens from Central region (filled circle), Pacific slope (filled star) and Atlantic slope (filled triangle). Filled gray: *Desmodus rotundus* range.</figcaption></figure>

## Materials and methods

The mite collection was made through active surveying of the common vampire bats *Desmodus rotundus* from the Pacific and Atlantic slopes of Mexico, corresponding to the states of Yucatán and Oaxaca, respectively, and the central region of the country in the state of Querétaro (Table 2), during different expeditions from 2005 to 2015 (Figure 1; Table 2). Vampire bat collections were made in natural refuges such as caves (Querétaro), and cattle paddocks or feeding areas (Yucatán and Oaxaca), using nylon mist nets 6m or 12m long by 2.6 m tall.

Vampire bats were identified using the field guides of Medellín et al. (2007). Bats were euthanized following protocols under scientific collector license FAUT-0070 and Permission SGPA/DGVS/02748/13, both granted to JBMM by the Secretaría del Medio Ambiente y Recursos Naturales, México (SEMARNAT). The bats were placed individually in polyethylene bags, each with a paper label containing the location, date, collector's name, and catalog number. The captured bats were then kept refrigerated (-29°C) for later examination at the Laboratorio de Espeleobiología y Acarología, Facultad de Ciencias, UNAM, Campus Juriquilla, Querétaro.

All the bat carcases were deposited in the Colección Nacional de Mamíferos, Instituto de Biología, Universidad Nacional Autónoma de México [CNMA], and Laboratorio de Zoonosis, Centro de Investigaciones Regionales, Dr. Hideyo Noguchi, Universidad Autónoma de Yucatán (UADY).

The collected vampire bats were examined in the laboratory under a Zeiss, Stemi 2000 dissecting microscope (Göttingen, Niedersachsen, Germany). Chirodiscid fur mites were sought in the dorsal and ventral regions of bats. Mites were removed with entomological forceps, mounted in Hoyer's medium (Morales-Malacara and Juste 2002) and examined under a phase contrast Zeiss Axioskop 2 plus microscope (ZA2) (Göttingen, Niedersachsen, Germany) for genus determination using specialized keys from Fain (1973), Guerrero (1992) and Bochkov (2010). Parasite prevalence (P) was calculated for each host population using the
Quantitative Parasitology 3.0 program (Rózsa *et al.* 2000).

All collected *Alabidocarpus* mites were deposited in the collection of J.B.M-M (MM), housed at Laboratorio de Espeleobiología y Acarología, de la Unidad Multidisciplinaria de Docencia e Investigación, Facultad de Ciencias, Universidad Nacional Autónoma de México, campus Juriquilla, Querétaro [MM]. Some specimens were selected for the type series.

Measurements of all *Alabidocarpus* mites selected as type specimens, as well as some voucher specimens, were taken with a Zeiss Axio Imager.A2 differential interference contrast (DIC) microscope (AIA2), with an AxioCam MRc and AxioVision 4.8.2 software (Göttingen, Niedersachsen, Germany). All measurements of body structures and leg segment setae were made with 40x and 100x objectives using the corresponding scale. These measurements were taken directly on each structure or from a series of sequential microphotographs, adding up all the measures and allowing for accurate and precise measurements of each structure's dimensions.

Additionally, to compare morphological features among species, update the taxonomic key for the species from the American continent, and provide more accurate morphological features for the new species of *Alabidocarpus*. A review of type specimens of *Alabidocarpus* from the A. Fain Collection at the Royal Belgian Institute of Natural Sciences, Brussels, Belgium [RBINS] was carried out using a Leitz DMRB Leica differential interference contrast (DIC) microscope, and Surveyor software (Wetzlar, Germany), as well as a phase contrast Zeiss Axioplan2 microscope, equipped with a Lumenera Infinity X CMOS camera, monitored through Deltapix Infinity X-32 software.

<table><caption>**Table 2** Collecting sites of the common vampire bat *Desmodus rotundus*, with their municipality, State, and respective geographic coordinates, and number of bats collected, infested host and the finding of the fur mite *Alabidocarpus ricardoguerreroi* n. sp. with its prevalence and number of parasites found.</caption><thead><tr><th>Locality/Region</th><th>Municipality</th><th>State</th><th>Latitude</th><th>Longitude</th><th>Dates</th><th>NB</th><th>IH</th><th>P</th><th>NP</th></tr></thead><tbody><tr><td>Riscos Cave/Center Mexico</td><td>Pinal de Amoles</td><td>Querétaro</td><td>21.19389</td><td>-99.51389</td><td>19 Nov. 2005</td><td>2</td><td>0</td><td>0</td><td>0</td></tr><tr><td>Riscos Cave/Center Mexico</td><td>Pinal de Amoles</td><td>Querétaro</td><td>21.19389</td><td>-99.51389</td><td>01 Oct. 2006</td><td>12</td><td>0</td><td>0</td><td>0</td></tr><tr><td>Riscos Cave/Center Mexico</td><td>Pinal de Amoles</td><td>Querétaro</td><td>21.19389</td><td>-99.51389</td><td>10 Mar. 2007</td><td>4</td><td>0</td><td>0</td><td>0</td></tr><tr><td>Riscos Cave/Center Mexico</td><td>Pinal de Amoles</td><td>Querétaro</td><td>21.19389</td><td>-99.51389</td><td>30 Jun. 2007</td><td>2</td><td>0</td><td>0</td><td>0</td></tr><tr><td>Kax-Ek/Atlantic Slope</td><td>Tixkokob</td><td>Yucatán</td><td>20.91949</td><td>-89.38915</td><td>11 Dec. 2012</td><td>2</td><td>0</td><td>0</td><td>0</td></tr><tr><td>Kax-Ek/Atlantic Slope</td><td>Tixkokob</td><td>Yucatán</td><td>20.91949</td><td>-89.38915</td><td>12 Dec. 2012</td><td>3</td><td>0</td><td>0</td><td>0</td></tr><tr><td>San Luis/Atlantic Slope</td><td>Tunkas</td><td>Yucatán</td><td>21.05894</td><td>-88.72734</td><td>01 Apr. 2013</td><td>11</td><td>0</td><td>0</td><td>0</td></tr><tr><td>Santa Isabel/Atlantic Slope</td><td>Buctzotz</td><td>Yucatán</td><td>21.22977</td><td>-88.82947</td><td>01 Apr. 2013</td><td>1</td><td>0</td><td>0</td><td>0</td></tr><tr><td>Esperanza/Atlantic Slope</td><td>Mérida</td><td>Yucatán</td><td>20.95969</td><td>-89.33742</td><td>01 Apr. 2013</td><td>4</td><td>0</td><td>0</td><td>0</td></tr><tr><td>San Antonio Aku/Atlantic Slope</td><td>Mérida</td><td>Yucatán</td><td>21.05677</td><td>-89.38489</td><td>01 Apr. 2013</td><td>6</td><td>0</td><td>0</td><td>0</td></tr><tr><td>San Martín/Atlantic Slope</td><td>Buctzotz</td><td>Yucatán</td><td>21.23956</td><td>-88.81494</td><td>03 Jun. 2013</td><td>3</td><td>0</td><td>0</td><td>0</td></tr><tr><td>Yaxché/Atlantic Slope</td><td>Sinanche</td><td>Yucatán</td><td>21.29362</td><td>-89.19428</td><td>23 Aug. 2013</td><td>2</td><td>0</td><td>0</td><td>0</td></tr><tr><td>Sinaí/Atlantic Slope</td><td>Sinanche</td><td>Yucatán</td><td>21.29365</td><td>-89.19426</td><td>23 Aug. 2013</td><td>5</td><td>0</td><td>0</td><td>0</td></tr><tr><td>Chen Delgado/Atlantic Slope</td><td>Buctzotz</td><td>Yucatán</td><td>21.23534</td><td>-89.2071</td><td>23 Aug. 2013</td><td>3</td><td>0</td><td>0</td><td>0</td></tr><tr><td>San Miguel Xcanan I/Atlantic Slope</td><td>Chacsinkin</td><td>Yucatán</td><td>20.25681</td><td>-89.06775</td><td>01 Oct. 2013</td><td>3</td><td>0</td><td>0</td><td>0</td></tr><tr><td>San Miguel Xcanan II/Atlantic Slope</td><td>Chacsinkin</td><td>Yucatán</td><td>20.25942</td><td>-89.071</td><td>01 Oct. 2013</td><td>2</td><td>0</td><td>0</td><td>0</td></tr><tr><td>Panaba I/Atlantic Slope</td><td>Chacsinkin</td><td>Yucatán</td><td>20.28606</td><td>-89.1145</td><td>01 Oct. 2013</td><td>2</td><td>0</td><td>0</td><td>0</td></tr><tr><td>Itzinte II/Atlantic Slope</td><td>Tunkas</td><td>Yucatán</td><td>21.00472</td><td>-88.7316</td><td>04 Dec. 2013</td><td>9</td><td>0</td><td>0</td><td>0</td></tr><tr><td>San Felipe Carrillo/Atlantic Slope</td><td>Bokoba</td><td>Yucatán</td><td>21.00139</td><td>-89.14388</td><td>02 Dec. 2013</td><td>1</td><td>0</td><td>0</td><td>0</td></tr><tr><td>10 de mayo/Atlantic Slope</td><td>Bokoba</td><td>Yucatán</td><td>21.015</td><td>-89.14472</td><td>02 Dec. 2013</td><td>2</td><td>0</td><td>0</td><td>0</td></tr><tr><td>Dzodzil/Atlantic Slope</td><td>Bokoba</td><td>Yucatán</td><td>21.00583</td><td>-89.16083</td><td>02 Dec. 2013</td><td>3</td><td>0</td><td>0</td><td>0</td></tr><tr><td><strong>Chepilme/Pacific Slope</strong></td><td><strong>San Pedro Pochutla</strong></td><td><strong>Oaxaca</strong></td><td><strong>15.7696</strong></td><td><strong>-96.44615</strong></td><td><strong>25 Jun. 2014</strong></td><td><strong>3</strong></td><td><strong>2</strong></td><td><strong>0.66</strong></td><td><strong>114</strong></td></tr><tr><td><strong>Chepilme/Pacific Slope</strong></td><td><strong>San Pedro Pochutla</strong></td><td><strong>Oaxaca</strong></td><td><strong>15.7696</strong></td><td><strong>-96.44615</strong></td><td><strong>26 Jun. 2014</strong></td><td><strong>1</strong></td><td><strong>1</strong></td><td><strong>1</strong></td><td><strong>11</strong></td></tr><tr><td>Riscos Cave/Center Mexico</td><td>Pinal de Amoles</td><td>Querétaro</td><td>21.19389</td><td>-99.51389</td><td>01 Jul. 2014</td><td>17</td><td>0</td><td>0</td><td>0</td></tr><tr><td><strong>Barra de Navidad/Pacific Slope</strong></td><td><strong>Santa María Colotepec</strong></td><td><strong>Oaxaca</strong></td><td><strong>15.81169</strong></td><td><strong>-97.01121</strong></td><td><strong>21 Aug. 2014</strong></td><td><strong>3</strong></td><td><strong>2</strong></td><td><strong>0.66</strong></td><td><strong>18</strong></td></tr><tr><td><strong>Barra de Navidad/Pacific Slope</strong></td><td><strong>Santa María Colotepec</strong></td><td><strong>Oaxaca</strong></td><td><strong>15.81169</strong></td><td><strong>-97.01121</strong></td><td><strong>22 Aug. 2014</strong></td><td><strong>3</strong></td><td><strong>2</strong></td><td><strong>0.66</strong></td><td><strong>91</strong></td></tr><tr><td><strong>Mandingas, Botanic garden UMAR/Pacific Slope</strong></td><td><strong>San Pedro Mixtepec (Juquila)</strong></td><td><strong>Oaxaca</strong></td><td><strong>15.91504</strong></td><td><strong>-97.07805</strong></td><td><strong>22 Oct. 2014</strong></td><td><strong>7</strong></td><td><strong>3</strong></td><td><strong>0.43</strong></td><td><strong>36</strong></td></tr><tr><td>Mandingas, Botanic garden UMAR/Pacific Slope</td><td>San Pedro Mixtepec (Juquila)</td><td>Oaxaca</td><td>15.91504</td><td>-97.07805</td><td>23 Oct. 2014</td><td>10</td><td>0</td><td>0</td><td>0</td></tr><tr><td><strong>Limones/Pacific Slope</strong></td><td><strong>San Pedro Mixtepec (Juquila)</strong></td><td><strong>Oaxaca</strong></td><td><strong>15.91120</strong></td><td><strong>-97.08287</strong></td><td><strong>22 Nov. 2014</strong></td><td><strong>10</strong></td><td><strong>1</strong></td><td><strong>0.1</strong></td><td><strong>36</strong></td></tr><tr><td><strong>Limones/Pacific Slope</strong></td><td><strong>San Pedro Mixtepec (Juquila)</strong></td><td><strong>Oaxaca</strong></td><td><strong>15.91120</strong></td><td>-97.08287</td><td>23 Nov. 2014</td><td>10</td><td>0</td><td>0</td><td>0</td></tr><tr><td><strong>Hojitales/Pacific Slope</strong></td><td><strong>San Pedro Mixtepec (Juquila)</strong></td><td><strong>Oaxaca</strong></td><td><strong>15.90914</strong></td><td><strong>-97.08422</strong></td><td><strong>22 May. 2015</strong></td><td><strong>4</strong></td><td><strong>1</strong></td><td><strong>0.25</strong></td><td><strong>49</strong></td></tr><tr><td><strong>Hojitales/Pacific Slope</strong></td><td>San Pedro Mixtepec (Juquila)</td><td>Oaxaca</td><td>15.90914</td><td>-97.08422</td><td>23 May. 2015</td><td>4</td><td>0</td><td>0</td><td>0</td></tr><tr><td><strong>Limones/Pacific Slope</strong></td><td>San Pedro Mixtepec (Juquila)</td><td>Oaxaca</td><td>15.91120</td><td>-97.08287</td><td>21 Aug. 2015</td><td>13</td><td>0</td><td>0</td><td>0</td></tr><tr><td><strong>Lucerna/Pacific Slope</strong></td><td>San Pedro Mixtepec (Juquila)</td><td>Oaxaca</td><td>15.91120</td><td>-97.08287</td><td>21 Aug. 2015</td><td>1</td><td>0</td><td>0</td><td>0</td></tr><tr><td><strong>Lucerna/Pacific Slope</strong></td><td><strong>San Pedro Mixtepec (Juquila)</strong></td><td><strong>Oaxaca</strong></td><td><strong>15.90025</strong></td><td><strong>-97.08727</strong></td><td><strong>22 Aug. 2015</strong></td><td><strong>5</strong></td><td><strong>2</strong></td><td><strong>0.4</strong></td><td><strong>7</strong></td></tr></tbody></table>

Notes: NB: Number of bats collected; IH: Infested hosts; P: Prevalence; NP: Number of parasites (*Alabidocarpus ricardoguerreroi* n. sp.) collected. Bold: Localities and regions where were found specimens of *Alabidocarpus ricardoguerreroi* n. sp.
**Material examined for comparative analysis**

*Alabidocarpus furmani* 1 ♀, 1 ♂, ex *Carollia perspicillata* ♂, ?? *Molossus ater* 485, Surinam
Expedition, 1971, Zool. Lab. Nijmegen. Loc Moeroekreek, Surinam, 16-9-71, Lukoschus &
Kok leg 484 (A. Fain Collection) [RBINS]. *A. guyanensis* Holotype ♂, Allotype ♀, ex *Artibeus
cinereus* ♀, Loc Rochambeau Frans Guyana, 12-10-71, Lukoschus & Kok leg 665 (A. Fain
Collection) [RBINS]. *A. saccopteryx* Holotype ♂, 2 ♂♂ Paratypes, ex *Saccopteryx bilineata*,
Lelydorp Surinam, 26-2-70 Lukoschus leg 486 (Inst. Méd. Trop. Anvers; A. Fain Collection)
[RBINS].

Drawings were prepared using the ZA2 with an attached drawing tube, and photographs
were taken with AIA2, equipped with an AxioCam MRc and AxioVision 4.8.2 software
(Göttingen, Niedersachsen, Germany). Morphological features and nomenclature were
assessed using taxonomic guides of Fain (1972), Guerrero (1992), and Bochkov (2010), with
additional details on idiosomal chaetotaxy following Guerrero (1992), Bochkov (2010) and
Griffiths *et al*. (1990). All measurements are given in micrometers.

Redefinition of the *Alabidocarpus* diagnosis, was made using the morpho-taxonomic
features from the original genus description by Ewing (1929) and additional morphological
analysis made by Lawrence (1948), Pinichpongse (1963), McDaniel (1970), Fain (1971, 1973),
and Guerrero (1992).

**Results**

Of the 173 vampire bats collected (Oaxaca n = 74, Querétaro n = 37, Yucatán n = 62), we recovered 362 *Alabidocarpus* mites, which were exclusively located on the dorsal side of the wings, particularly in the propatagium on 14 bats from Oaxaca (14/74) (Pacific slope). A differential prevalence of these mites was observed in some populations of the vampire bats (Table 2), with an average prevalence of 18.9%. However, no *Alabidocarpus* mites were found in the Atlantic slope (Yucatán) or the central region of México (Querétaro) (Table 2).

All active stages were described based on 26 specimens (type series); while the remaining
specimens were considered additional voucher material.

**Alabidocarpus Ewing 1929**

*Alabidocarpus* Ewing, 1929: 188

*Alabidocarpus*, Lawrence, 1948: 374; Pinichpongse, 1963: 266; McDaniel, 1970: 804; Fain, 1971: 140; Fain, 1973: 115; Guerrero, 1992: 31

Type species. *Labidocarpus megalonyx* Trouessart, 1895

**Diagnosis**

Males and females. Idiosoma laterally compressed. Prescapular shield conspicuous, extending posteriorly to or beyond the level of legs II, with its lateral margins extending ventrally between coxae I and II. Setae *si* and *se* short appearing as microsetae, vestigial or alveoli. Setae *c2* short, appearing as microsetae, vestigial or alveoli located at the level of or slightly anterior or posterior to the level of the base of the setae *cp*. Legs III and IV equal or sub-equal, with four articulated segments; tarsi III and IV without ambulacral discs. Tarsus III with apical long claw-like not bifid seta *f* and with 2 thickened striated setae: *s* and *w*; additionally has 2 to 3 simple thin setae: *d*, *e*, and/or *r*. Tarsus IV with apical long claw-like not bifid seta *f* and with only one thick striated seta *w*, additionally has 2 to 3 simple thin setae: *d*, *e*, and/or *r*.

***Alabidocarpus ricardoguerreroi*** *Morales-Malacara and Vázquez-Xicoténcatl*
***n. sp.***

Zoobank: 7BFE93BD-CD35-45E6-9779-2C9D11714A40

(Figures 2-9)
Diagnosis

Males and females with prescapular shield, posterior border rounded and slightly sinuous; setae *si* and *se* vestigial inconspicuous; seta *c3* medium size, seta *cp* very long, seta *c2* vestigial, and setae *e1* and *e2* as alveoli. Males with 3 pairs of opisthosomal setae, *h1* minute, *h2* short and *h3* medium. Females with only 2 pairs of opisthosomal setae, *h1* absent, *h2* very long, and *h3* long. Coxa IV in males and females with very fine and setiform seta *4a*, and seta *e* on tarsus IV absent. Males with seta *d* on tarsus IV, four times longer than in females.

Male

<figure><img src="image_4.png" /><figcaption>**Figure 2** *Alabidocarpus ricardoguerreroi* n. sp. Male holotype, DIC photograph (including chaeto-taxy nomenclature). Scale bar: 100 µm.</figcaption></figure>

Idiosoma short (Figures 2 and 3A). Gnathosomal shield conical in lateral view. Chelicera medium sized, with dorsal border of the fixed digit with three serrated spines directed backward, and inner ventral border as a broad knife-like edge, and with frontal border with broad pointed edge, and movable digit articulated, with inner dorsal border with two denticles directed backward and with distal edge pointed (Figure 3B). Prescapular shield rounded and slightly sinuous, with its dorsal border medium size (40–43), and in the midline longer (50–57). Unarmed cuticle with 32–34 striations. Idiosomal setae: *si* and *se* vestigial inconspicuous (3–4); *c3* medium size (34–54) and *cp* very long (128–167), both located over coxa III; *c2* vestigial (2–3); *d1* vestigial (2–3); *e1* alveoli not well defined, setae *d2* and *e2* alveoli (Figure 3A). Opisthosomal shield from the bottom to dorsal border with its posterior edge sinuous, with narrowed length (27–31), Opisthosomal concavity conspicuous with a pair of rounded paranal suckers (Figure 3C). Setal pair *h1* minute (4–5) located paraxial inside the opisthosomal concavity at level of *h2*; *h2* short (15–24) and *h3* medium (26–36), both located at the opisthosomal concavity border (Figure 3C). Legs I and II modified, with segments trochanter–genua reduced, and integrated to the pro-lateral region of idiosoma; with small
tibiae, with one medium-sized curved postero-lateral solenidion φ (18–24). With tarsi I and II modified as an articulated and wide fan-like tarsus, with small ventro-lateral solenidion ω1 (11–17) (Figure 3A). Coxal III seta 3a short (17–18). Tibia III with anterior seta kt short (23–34), and posterior solenidion φ short (22–23). Tarsus III with both thickened seta w and s striated and short (15 and 12–14 respectively); apical claw-like seta f medium size (29–35); seta d short (12–15), setae e and r very short (5–6 and 7–10 respectively) (Figure 4A). Coxal IV seta 4a setiform medium size (33–48); setal pair g minute (4–6) and located at the base of the aedeagus. Tibia IV with solenidion φ medium size (26–30); Tarsus IV with thickened seta w medium size (22–26), apical claw-like seta f long size (58–64); seta d very long (73–92), seta r short (11–15), and seta e is absent (Figure 4B). Aedeagus located between legs IV, in lateral view it seems medium size (20–21) in some paratypes and curved (Figure 3D).

<figure><img src="image_5.png" /><figcaption>**Figure 3** *Alabidocarpus ricardoguerreroi* n. sp. Male holotype (including chaetotaxy nomenclature). A – Idiosome, B – Chelicera, C – Opistosomal shield, D – Aedeagus. Scale bars: A: 100 µm; B: 10 µm; C: 30 µm; D: 20 µm.</figcaption></figure>

*Measurements*. Holotype ♂ (followed in parentheses by range and mean of 4 Paratype ♂♂ and the Holotype [exceptions noted]): Idiosoma length in lateral view (gnathosoma included) 329 (298–330, 317), width 111 (105–128, 115); length gnathosomal triangle shield at midline in lateral view 49 (48–51, 49); prescapular shield at dorsal edge 43 (40–43, 41.5); prescapular shield length at midline in lateral view 57 (49–57, 53.6); opisthosomal shield length in lateral view at midline, 27 (27–31, 28). Aedeagus length in lateral view (not well seen in the holotype
male and some other paratypes), – (20–21,18.7 [n = 2]); seta g besides aedeagus, 6 (4–6, 5.4).
Setal lengths: *si*, 3 (3–4, 3.1); *se*, 4 (3–4, 3.3); *c3*, 52 (50–54, 48); *cp*, 163 (128–167, 151); *c2*,
2 (2–3, 2.4); *dl*, 2 (2–3, 2.4 [n = 4]); *h1*, 5 (4–5, 4.1); *h2*, 19 (17–24, 20); *h3*, 36 (26–36, 33);
Legs setae: φ tibia I, 22 (21–24, 23); ω1 tarsus I, 15 (13–16, 14.7); φ tibia II, 20 (18–20, 19);
ω1 Tarsus II, 14 (11–17, 13.8); 3a coxa III, 17 (17–18, 17.6); kt tibia III, 29 (23–34, 28); φ tibia
III, 22 (22–23, 22.6); w tarsus III, 15 (15–15, 15); s tarsus III, 13 (12–14, 13.2); d tarsus III, 12
(12–15, 13.4); e tarsus III, 5 (5–6, 5.6); r tarsus III, 7 (7–10, 8); f tarsus III, 35 (29–35, 32); 4a
coxa IV, 33 (33–48, 39); φ tibia IV, 26 (26–29, 27.6); w tarsus IV, 22 (22–26, 23.7); d tarsus IV,
91 (73–92, 84); r tarsus IV, 13 (11–15, 13.3); f tarsus IV, 63 (56–64, 60.5).

<figure><img src="image_6.png" /><figcaption>**Figure 4** *Alabidocarpus ricardoguerreroi* n. sp. Legs III–IV tibia and tarsus and chaetotaxy of male: A – leg III, B – leg IV; legs III–IV tibia and tarsus and chaetotaxy of female: C – leg III, D – leg IV; E – Leg IV tibia and tarsus and chaetotaxy of male protonymph. Scale bars: A–E: 20 µm.</figcaption></figure>

Female

Same as male, except as noted. Idiosoma longer (Figures 5 and 6). Prescapular shield rounded and slightly sinuous, with its dorsal border medium size (42–48), and in the midline longer (54–57). Unarmed cuticle with 54–58 striations. Idiosomal setae: *si* and *se* vestigial inconspicuous, located very near to the posterior border of prescapular plate; *c3* medium size (33–42) and *cp* very long (134–145), both located near each other and over coxa III; *c2* vestigial; seta *e1* alveoli
<figure><img src="image_7.png" /><figcaption>**Figure 5** *Alabidocarpus ricardoguerreroi* n. sp. Idiosome of pregnant female paratype, DIC photograph (including chaetotaxy nomenclature; L: developing larva inside the female). Scale bar: 50 µm.</figcaption></figure>

not well defined; setae *d2* and *e2* only distinguishable by the alveoli; *h2* very long (118–150) and *h3* long (81–99), both located at the opisthosomal border (Figure 6). Coxal III seta *3a* short (13–16), Tibia III with anterior seta *kt* short (19–33), and posterior solenidion *φ* short (16–21); Tarsus III with both thickened seta *w* and *s* striated and short (12–15 and 11–13 respectively), apical claw-like seta *f* medium size (31–33); tarsus III setae *d* short (12–14) and setae *e* minute (4–6) and *r* very short (8–10) (Figure 4C). Coxal IV setae *4a* setiform short (18–28). Tibia IV with solenidion *φ* medium size (20–25). Tarsus IV with thickened seta *w* short (19–22), apical claw-like seta *f* long size (58–62); setae *d* medium size (16–20); *r* short (12–15), and setae *e* is absent (Figure 4D).

<figure><img src="image_8.png" /><figcaption>**Figure 6** *Alabidocarpus ricardoguerreroi* n. sp. Female allotype (including chaetotaxy nomenclature). Scale bar: 100 µm.</figcaption></figure>

*Measurements*. Allotype ♀ (followed in parentheses by range and mean of 4 Paratype ♂♂ and the Allotype [exceptions noted]): Idiosoma length in lateral view (gnathosoma included) 494 (456–494, 477), width 137 (114–137, 126); length gnathosomal triangle shield at midline in lateral view 52 (49–52, 50); prescapular shield at dorsal edge 48 (42–48, 44); prescapular shield length at midline in lateral view 57 (54–57, 55). Setal lengths: *si*, 3 (3–3, 2.6 [n = 4]); *se*,
3 (2–3, 2.8); c3, 42 (34–41, 38); cp, 137 (132–145, 139 [n = 4, because 1 broken]); c2, 3 (2–3, 2.3); d1, 3 (2–3, 2.6 [n = 2, because on three just seen as alveoli]); h2, 150 (118–150, 128); h3, 84 (81–99, 87). Legs setae: φ tibia I, 20 (19–21, 20); ω1 tarsus I, 16 (13–16, 14); φ tibia II, 18 (16–19, 16.4); ω1 tarsus II, 13 (11–13, 12.3); 3a coxa III, 14 (13–16, 15); kt tibia III, 23 (19–33, 24); φ tibia III, 21 (16–21, 18.5); w tarsus III, 14 (12–14, 13.4); s tarsus III, 13 (11–13, 12.2); d tarsus III, 12 (12–14, 12.8); e tarsus III, 5 (4–6, 4.3); r tarsus III, 9 (8–10, 9.6); f tarsus III, 32 (31–33, 32); 4a coxa IV, 28 (18–28, 21); φ tibia IV, 25 (20–25, 23); w tarsus IV, 20 (19–22, 20); d tarsus IV, 20 (16–20, 18); r tarsus IV, 15 (12–15, 13); f tarsus IV, 60 (58–62, 59).

<figure><img src="image_9.png" /><figcaption>Male larva</figcaption></figure>

**Figure 7** *Alabidocarpus ricardoguerreroi* n. sp. Larvae, DIC photographs (including chaetotaxy nomenclature): A – male larva paratype (arrow: caudal region with normal striated cuticle), B – female larva paratype (PN: female protonymph developing inside its body) (arrow: smooth cuticle on dorso-caudal region). Scale bar: 50 µm.

Smaller than male (Figure 7A). Gnathosomal triangle shield and prescapular shield similar to male, but slightly smaller in dimensions. Unarmed cuticle with 62 striations including the
caudal region of opisthosoma with normal striated cuticle. Idiosomal setae, *si* and *se* vestigial inconspicuous as in adult male (2-3); *c3* medium size (12) and *cp* long (69), both located over coxa III; *c2* vestigial (3); setae *d1* absent or not seen, *d2* and *e2* as alveoli. Without opisthosomal shield. Legs I and II as in male, and leg III, normally developed.

*Measurements.* Paratype male L (followed in parentheses by range and mean of a total of 2 Paratype male LL [exceptions noted]): Idiosoma length in lateral view (gnathosoma included) 277 (277-373, 339), width 95 (95-119, 110); prescapular shield at dorsal edge 27 (27-31, 30); prescapular shield length at midline in lateral view 35 (35-37, 36). Setal lengths: *si*, 3 (3-3, 3); *se*, 3 (2-3, 2.5); *c2*, 3 (2-3, 2); *c3*, 12 (12-16, 14); *cp*, 69 (66-69, 68); *d1*, – (2 [n = 1]) *h2*, 99 (80-99, 90 [n = 2]).

## Female Larva

Same as in male larva, except as noted. Unarmed cuticle with 29-32 striations and interrupted smooth cuticle without regular striations on the dorso-caudal region, with a few small irregular and perpendicular lines (Figure 7B). Idiosomal setae *c3* and *h2* are slightly smaller compared to the male larva. All other features as in male larva.

*Measurements.* Paratype female L (followed in parentheses by range and mean of a total of 3 Paratype female LL [exceptions noted]): Idiosoma length in lateral view (gnathosoma included) 225 (225-259, 237), width 86 (86-93, 89); prescapular shield at dorsal edge 29 (28-30, 29); prescapular shield length at midline in lateral view 32 (32-35, 34). Setal lengths: *si*, 2 (2-2, 2); *se*, 2 (2-2, 2); *c2*, 2 (2-2, 2); *c3*, 8 (8-10, 9); *cp*, 67 (67-74, 70 [n = 2]); *h2*, 56 (56-67, 62).

## Male protonymph

As in adult male, except as noted. Idiosoma slightly bigger than the adult male (334-365). Unarmed cuticle with about 56-58 striations. Gnathosomal triangle shield and prescapular shield similar to male, but slightly smaller in dimensions. Idiosomal setae, *si* and *se* vestigial inconspicuous as in adult male (2-3); *c3* medium size (17-23) and *cp* long (84-97), both located over coxa III; *c2* vestigial (2-3); setae *d1* vestigial (2-3), *d2* and *e2* as alveoli. Without opisthosomal shield. Legs I and II as in male, and Leg III, normally developed. Seta *h1* absent; Seta *g* Coxa IV absent; solenidion *φ* tibia IV absent; seta *e* tarsus IV absent (Figure 4E).

*Measurements.* Paratype Male PNN (followed in parentheses by range and mean of a total of 3 Paratype Male PNN [exceptions noted]): Idiosoma length in lateral view (gnathosoma included) 334 (334-365, 351), width 113 (97-123, 111); prescapular shield at dorsal edge 31 (31-34, 33); prescapular shield length at midline in lateral view 40 (40-43, 42). Setal lengths: *si*, 2 (2-4, 3 [n = 2]); *se*, 2 (2-4, 3 [n = 2]); *c2*, 2 (2-3, 2); *c3*, 23 (17-23, 19); *cp*, 91 (84-97, 91); *d1*, 3 (2-3, 2.5 [n = 2]); *h2* 99 (99-112, 105); *h3*, 66 (56-66, 62); Legs setae: *3a* coxa III, 8 (8-10, 9); *kt* tibia III, 17 (17-24, 20); *φ* tibia III, 13 (11-13, 12); *w* tarsus III, 12 (12-12, 12); *s* tarsus III, 10 (9-10, 9.5); *d* tarsus III, 9 (9-10, 9.5); *e* tarsus III, 4 (4-5, 4.5); *r* tarsus III, 6 (5-7, 6); *f* tarsus III, 26 (25-27, 26). *4a* coxa IV, 9 (9-11, 10.5); *w* tarsus IV, 11 (11-12, 11.8); *d* tarsus IV, 9 (9-10, 9.2); *r* tarsus IV, 6 (6-7, 6.7); *f* tarsus IV, 48 (42-48, 44.8).

## Female protonymph

Elongated idiosome, almost all its length with striations (41 striations). The fronto-dorsal region moderately rounded with unstriated cuticle. The chelicera lobes in the anteriormost region are almost imperceptible, and the ventrolateral area is covered with arm-like lobulated palps medium-sized. With four legs I-IV just represented with very small underdeveloped stump, that seems constituted with two short segments (Figure 8A). Legs I and II stump with 2 tiny setae (*φ* and *ω*), leg III stump with 3 tiny setae (*w*, *s* and *f*), and leg IV stump with 2 tiny setae (*w* and *f*). Ventro-caudal region with one pair of tiny setae *h2* (3).

*Measurements.* Paratype Female PN: Idiosoma length in lateral view (gnathosoma included) 207, width 82. Setal lengths: *h2*, 3. Legs stump setae: *φ* tibia I, 1.33; *ω1* tarsus I, 2; *φ* tibia II, 1.46; *ω1* tarsus II, 2; *w* tarsus III, 3; *s* tarsus III, 3; *f* tarsus III, 5; *w* tarsus IV, 2; *f* tarsus IV, 5.
<figure><img src="image_10.png" /><figcaption>**Figure 8** *Alabidocarpus ricardoguerreroi* n. sp. A – Detail of legs I–IV stumps of female protonymph. B – Idiosome and detail of legs I–IV stumps of female tritonymph. Scale bars: A–B (legs stumps): 10 µm; B (Idiosome): 50 µm.</figcaption></figure>

Male tritonymph

As in female, exceptions noted. Idiosoma slightly smaller (298–401). Unarmed cuticle with 58 striations. Without opisthosomal shield. Idiosomal setae smaller: *si* and *se* vestigial inconspicuous (2–4); *c3* medium (18–27) and *cp* moderately long (102); *c2* vestigial (2); setae *d1* (3), *e1* not seen or absent; *d2* and *e2* alveoli. Setae *h2* very long (123–127) and *h3* long (75–79). Coxal III seta *3a* small (9–10). Tibia III with anterior seta *kt* medium sized (16–20), and posterior solenidion φ short (12). Tarsus III with both thickened seta *w* and *s* striated and short (11–12 and 11 respectively); apical claw-like seta *f* medium size (25–28); seta *d* short (9–10), setae *e* and *r* very short (4–5 and 6–8 respectively). Coxal IV seta *4a* medium size (21–26); setal pair *g* present and short (9–11) and located behind the setae *4a* (Figure 9). Tibia IV with solenidion φ short (10–11); tarsus IV with thickened seta *w* medium size (14), apical claw-like seta *f* moderately long size (41–43); seta *d* short (10–11) and seta *r* short (8), and seta *e* is present and diminutive (4–5).

*Measurements.* Paratype Male TN (followed in parentheses by range and mean of a total of 3 Paratype Male TNN [exceptions noted]): Idiosoma length in lateral view (gnathosoma included) 347 (298–401, 349), width 112 (86–139, 112); prescapular shield at dorsal edge 35 (35–35, 35); prescapular shield length at midline in lateral view 44 (39–44, 42). Setal lengths: *si*, 4 (2–4, 3); *se*, 3 (2–3, 3); *c2*, 2 (2–2, 2); *c3*, 18 (18–27, 22); *cp*, – (100–102, 101 [n = 2]); *d1*, – (2–3, 2.5 [n = 2]); *h2* 127 (122–128, 126); *h3*, 79 (75–80, 78). Legs setae: *3a* coxa III, 9 (9–13, 11); *kt* tibia III, 20 (16–20, 18); φ tibia III, 12 (12–15, 13); *w* tarsus III, 11 (11–12, 11); *s* tarsus III, 11 (9–11, 10); *d* tarsus III, 9 (9–10, 10); *e* tarsus III, 4 (3–5, 4); *r* tarsus III, 6 (6–8, 7); *f* tarsus III 28 (25–28, 27). *4a* coxa IV, 26 (18–26, 22); *g*, 11 (9–11, 10); φ tibia IV, 11
<figure><img src="image_11.png" /><figcaption>**Figure 9** *Alabidocarpus ricardoguerreroi* n. sp. Coxa IV region with detail of setae 4a and g of male tritonymph paratype. Scale bar: 20 µm.</figcaption></figure>

(10–11, 11); *w* tarsus IV, 14 (14–16, 15); *d* tarsus IV, 11 (10–12, 11); *e* tarsus IV, – (4–5, [n = 2]); *r* tarsus IV, 8 (8–9, 8); *f* tarsus IV, 43 (41–47, 44).

## Female tritonymph

As in female protonymph, exceptions noted. Elongated idiosome, almost all its length with striations (36–46 striations). With four legs I-IV just represented with very small underdeveloped stump, that seems constituted with two short segments (Figure 8B). Legs I and II stump with 2 tiny setae ($\varphi$ and $\omega 1$), leg III stump with 4 tiny setae ($\varphi$, *w*, *s* and *f*), and leg IV stump with 3 tiny setae ($\varphi$, *w* and *f*). Ventro-caudal region with one pair of tiny setae *h2* (2-3).

*Measurements.* Paratype Female TN (followed in parentheses by range and mean of a total of 3 Paratype Female TNN [exceptions noted]): Idiosoma length in lateral view (gnathosoma included) 210 (210–352, 279), width 107 (94–120, 107). Setal lengths: *h2*, 3 [n = 1]. Legs stump setae: $\varphi$ tibia I, 2 (1.4–2, 1.7); $\omega 1$ tarsus I, 2.8 (2.8–3.14, 2.9); $\varphi$ tibia II, – (1.51–1.54, 1.52 [n = 2]); $\omega 1$ tarsus II, – (2.11–2.45, 2.28 [n = 2]); $\varphi$ tibia III, 1.13 (1.13–2.1, 1.46); *w* tarsus III, 2.8 (2.8–3.18, 3.26); *s* tarsus III, 2.48 (2.09–2.48, 2.22); *f* tarsus III, 2.09 (2.09–3.87, 3.26); $\varphi$ tibia IV, – (1.85–3.71, 2.78 [n = 2]); *w* tarsus IV, – (2.88–3.31, 3.09); *f* tarsus IV, 2.98 (2.98–4.52, 3.90).

## Remarks

The morpho-structural similarity and phenotypic characteristics of the proterosoma of male larvae, female larvae (Figure 7), male protonymphs, male tritonymphs, as well as adult males (Figure 2), and females (Figure 5) show similar morphological features of the gnathosoma and the prescapular shield. They also share similarities in idiosomal chaetotaxy, except that in males, there are variations in the opisthosomal area, which includes the opisthosomal shield, opisthosomal concavity with paranal suckers, and differences in the number and characteristics of the opisthosomal setae. These features distinguish males from adult females and contribute to their conspicuous sexual dimorphism.

Another sexually dimorphic characteristic observed in males was the seta *d* of tarsus IV, which was four times longer than in females (Figure 4B). Also, additional dimorphic characteristics were evident in both developmental lines (male and female).
Additionally, the pronounced dimorphism in female protonymphs and female tritonymphs stands out (Figure 8). Their morphology is drastically altered to assume almost entirely apodous states, except for the presence of stumps of legs I-IV, the extreme reduction of the gnathosoma area, and the absence or reduction of sclerotization in the prodorsum region.

Furthermore, in *Alabidocarpus ricardoguerreroi* n. sp. some other traits were observed that evidenced sexual dimorphism. In male larvae, the dorso-caudal opisthosomal region showed normal striae (Figure 7A). In contrast, in female larvae, the dorso-caudal opisthosomal region displayed a smooth area with a few irregular lines (Figure 7B), indicating that they are precopulatory female larvae.

## Type Series

Holotype ♂, ex *Desmodus rotundus* ♂, Chepilme, San Pedro Pochutla, Oaxaca, MÉXICO, 25-VI-2014, C. García-Estrada (M7V001.35) [MM]. Allotype ♀, ex *D. rotundus* ♂, same data (M7V003.27). 2 Paratype ♀♀, 2 Paratype LL♀, 1 Paratype PN♂, 2 Paratype TNN♂, ex *D. rotundus* ♂, same data (M7V001.11, .40, .46, .49, .56, .58, .77) [MM]. 2 Paratype ♂♂, 2 Paratype ♀♀, 1 Paratype L♂, 1 Paratype PN♂, 1 Paratype TN♂, ex *D. rotundus* ♂, same data (M7V003.24, .26, .29-.30, .46, .50, .61) [MM]. 1 Paratype L♂, ex *D. rotundus*, same data (M7V004.2) [MM]. 1 paratype L♂, 2 Paratype TNN♀, ex *D. rotundus* ♀, Barra de Navidad, Santa María Colotepec, Oaxaca, MÉXICO, 22-VIII-2014, C. García-Estrada (M7W005.9, .15, .34) [MM]. 1 Paratype ♂, 1 Paratype L♀, ex *D. rotundus*, same data (M7W006.16-.17) [MM]. 1 Paratype ♂, 1 Paratype TN♀, ex *D. rotundus* ♂, Juquila San Pedro Mixtepec, Mandingas, Jardín Botánico, UMAR, Oaxaca, MÉXICO, 22-X-2014, C. García-Estrada (M7X001.10a,b) [MM]. 1 Paratype PN ♀, ex *D. rotundus*, Juquila, San Pedro Mixtepec, Los Limones, 22-XI-2014, C. García-Estrada (M7Y007.6a) [MM]. 1 Paratype PN♂, ex *D. rotundus*, Rancho La Lucerna, San Pedro Mixtepec, Oaxaca, MÉXICO, 22-V-2015, C. García-Estrada (M8B004.12) [MM].

## Deposition of types

The Holotype ♂ (M7V001.35), Allotype ♀ (M7V003.27), 1 Paratype L♂ (M7V003.26), 1 Paratype L♀ (M7V001.11), 1 Paratype PN♂ (M7V001.58), 1 Paratype TN♂ (M7V003.46), 1 Paratype TN♀ (M7W005.9) at Colección Nacional de Ácaros, Instituto de Biología, Universidad Nacional Autómoma de México, Mexico City (CNAC); 1 Paratype ♂ (M7V003.24), 1 paratype ♀ (M7V003.50), 1 Paratype L♀ (M7V001.77) at The Natural History Museum (British Museum), London, England (BM); 1 Paratype ♂ (M7W006.17) [RBINS/IG: 34844], 1 Paratype ♀ (M7V001.40) [RBINS/IG: 34844] at Royal Belgium Institute of Natural Sciences, Brussels, Belgium (RBINS); the remaining paratypes: 2 Paratype ♂♂ (M7V003.61, M7X001.10a), 2 Paratype ♀♀ (M7V001.56, M7V003.29), 2 Paratype LL♂ (M7V004.2, M7W005.34), 1 Paratype L♀ (M7W006.16), 2 Paratype PNN♂ (M7V003.30, M8B004.12), 1 Paratype PN♀ (M7Y007.6), 2 Paratype TNN♂ (M7V001.46, M7V001.49), 2 Paratype TNN♀ (M7W005.15, M7X001.10b) at Morales-Malacara Collection, housed at Laboratorio de Espeleobiología y Acarología, Unidad Multidisciplinaria de Docencia e Investigación, Facultad de Ciencias, UNAM, campus Juriquilla, Querétaro, México (MM).

## Additional Material Examined

MÉXICO: OAXACA: 12 ♂♂, 23 ♀♀, 1 L♂, 10 LL♀, 1 TN♂, ex *Desmodus rotundus*, Chepilme, San Pedro Pochutla, 26-VI-2014, C. García Estrada (M7V001.1, .33,.34,.36-.39, .41-.45, .47-.48, .50-.51, .54-.55, .57, .59-.76, .78-.87). 17 ♂♂, 30 ♀♀, 1 L♀, 2 TN♂♂, 1 TN♀, ex *D. rotundus*, same data, 27-VI-2014, C. García Estrada (M7V003.16-.23, .25, .28, .31-.45, .47-.49, .51-.60, .62-.71a,b-.72-.73). 2 ♂♂, 5 ♀♀, 1 L♀, 2 PNN♂, ex *D. rotundus*, same data, 27-VI-2014, C. García Estrada (M7V004.1, .26-.32, .34-.35). 3 ♂♂, 9 ♀♀, 3 PNN♂, 2 TNN♂, ex *D. rotundus*, Barra de Navidad, Santa María Colotepec, 21-VIII-2014, C. García Estrada (M7W002.2-.4, .10a,b-.11a,b, .13a,b-.14, .16-.17a,b, .18-.21). 1 ♂, ex *D. rotundus*,
same data, 21-VIII-2014, C. García Estrada (M7W003.3). 15 ♂♂, 29 ♀♀, 2 LL♂, 2 LL♀, 2 PNN♂, 3 TNN♂, 3 TNN♂, 3 TNN♀, ex *D. rotundus*, same data, 22-VIII-2014, C. García Estrada (M7W005.2a,b–3a,b, .4–.6, .7a,b–.9a, .10–.12a,b–.13a,b–.14a,b–.15a, .16–.21a,b – .22–.23, .25–.33a,b, .35–.36a,b–.37–.41a,b–.42–.43a,b–.44a,b–.45a,b,c). 5 ♂♂, 16 ♀♀, 4 LL♂, 3 LL♀, 2 TNN♂, ex *D. rotundus*, same data, 22-VIII-2014, C. García Estrada (M7W006.1a,b–.2a,b–.3–.4a,b–.5a,b–.6–.7a,b–.8a,b–.9a,b, .12a,b–.13a,b–.14a,b–.15a,b,c, .18a,b–.19a,b–.20). 5 ♂♂, 1 L♀, 5 TNN♀, ex *D. rotundus*, Juquila, San Pedro Mixtepec, Mandingas, Jardín Botánico, 22-X-2014, C. García Estrada (M7X001.4a,b–.5a,b,c, .11a,b–.12a,b–.13a,b). 1 ♀, ex *D. rotundus*, same data, 22-X-2014, C. García Estrada (M7X002.1). 3 ♀♀, 1 L♀, ex *D. rotundus*, same data, 22-X-2014, C. García Estrada (M7X006.2a,b,c–.3). 2 ♂♂, 1 ♀, 2 LL♀, 1 TN♀, ex *D. rotundus*, Juquila, San Pedro Mixtepec, Los Limones, 22-XI-2014, C. García Estrada (M7Y007.3–.4a,b–.5a,b–.6b). 2 ♂♂, 6 ♀♀, 2 LL♂, 4 LL♀, 1 PN♂, 3 TNN♂, ex *D. rotundus*, Rancho La Lucerna, San Pedro Mixtepec o Los Hojitales, 23-V-2015, C. García Estrada (M8B004.4–.5a,b,c–.6, .8a,b–.9a,b–.10a,b–.11a,b,.13a,b–.14a,b). 1 ♀, ex *D. rotundus*, Juquila, Rancho La Lucerna, 21-VIII-2015, C. García Estrada (M8C005.3).

## Specimens preserved in alcohol

18 specimens, ex *D. rotundus*, Juquila, San Pedro Mixtepec, Mandingas, Jardín Botánico, 22-X-2014, C. García Estrada (M7X001). 29 specimens, ex *D. rotundus*, Juquila, San Pedro Mixtepec, Los Limones, 22-XI-2014, C. García Estrada (M7Y007). 30 specimens, ex *D. rotundus*, Rancho Los Hojitales, San Pedro Mixtepec, 23-V-2015, C. García Estrada (M8B004). 6 specimens, ex *D. rotundus*, Juquila, Rancho La Lucerna, 21-VIII-2015, C. García Estrada (M8C006).

## Etymology

The species name is dedicated to Dr. Ricardo Guerrero, a good friend and an enthusiastic researcher on Vertebrate endoparasites and ectoparasites, in recognition of his academic and research trajectory.

## Discussion

*Alabidocarpus ricardoguerreroi* n. sp. is a permanent ectoparasitic mite that completes its entire life cycle on its host, making it a monoxenous species associated with *Desmodus rotundus*. As such, it would be expected to be distributed across all populations of its host throughout its geographical range, which extends from the state of Tamaulipas in eastern Mexico, westward to Sonora, southward through Central America, the Antilles in Trinidad, and into South America, including Uruguay, Northern Argentina, and central Chile (Greenhall *et al.* 1983).

However, this fur mite species, *Alabidocarpus ricardoguerreroi n. sp.*, was found only in association with vampire bats from the Pacific slope, specifically in localities from the state of Oaxaca, México (Table 2). Additionally, *A. ricardoguerreroi n. sp.* shows a differential prevalence from very low to high (10% – 100%), with an average prevalence of 18.9% across different collecting sites on the Pacific slope. This prevalence is comparable to other monoxenous mite species, such as *Periglischrus herrerai* Machado-Allison (Mesostigmata: Spinturnicidae), which is associated with *Desmodus rotundus* in some regions of the neotropics, where the average prevalence is moderate, around 27% (Morales-Malacara *et al.* 2018). Therefore, this prevalence could be considered a stable minimum for the survival of monoxenous ectoparasites associated with bats.

*Alabidocarpus ricardoguerreroi n. sp.* is phenetically similar to *A. longipilus* in that the posterior border of the prescapular plate is slightly curved and sinuous, without posterolateral projections. However, *A. ricardoguerreroi n. sp.* can be easily distinguished from *A. longipilus* by the fine, setiform seta 4a on Coxa IV (in *A. longipilus*, seta 4a is short and thickened), and the medium to large size of setae c3 (in *A. longipilus*, seta c3 is short). Furthermore, the two species are associated with different families and genera of bats. *A. ricardoguerreroi n. sp.* is
— Female with 2 pairs of opisthosomal setae, *h2* very long, *h3* long. Coxa IV with very fine and setiform seta 4a. Seta *c3* medium sized.................. *A. ricardoguerreroi* n. sp.

Note: The male of *A. longipilus* remain unknown.

## Acknowledgments

The senior author expresses his appreciation to Dr. Wouter Dekoninck (Royal Belgium Institute of Natural Sciences, Brussels) who allowed to the first author to review type and voucher specimens from the A. Fain Collection. The senior author also would like to thank to Camille Locatelli (Royal Belgium Institute of Natural Sciences, Brussels) who helped with the use of the Leica equipment, and thank Dr. Ninon Robin for help with accommodating image acquisition on site with the use of Zeiss Axionplan2 microscope. We also thank for their help in different field collections of parasitic mites to Juan M. Córdova Lara and Daniel I. Córdova Aldana, Ariadna B. Eb Córdova, Esther G. Marroquín Lavadores, Juan M. Canto Osorio, Violeta Jiménez Parejas, Liliana Trujillo Pahua, Claudia M. Chávez Ramírez, Adriana Espino del Castillo, Itzel Sigala Regalado, Miguel A. Hernández Patricio, Ricardo Paredes León, Edith Vázquez Ayala, and Lorenzo Ruiz Santos. We also thank Laura Del Castillo Martínez (Laboratorio de Acarología, Facultad de Ciencias, Universidad Nacional Autónoma de México) Laura Daniela Bonilla Hernández (Laboratorio de Espeleobiología y Acarología, Unidad Multidisciplinaria de Docencia e Investigación, Facultad de Ciencias, Campus Juriquilla) for their helpful assistance in mounting some additional material on microspopic slides. To Leon E. Ibarra Garibay (Unidad Multidisciplinaria de Docencia e Investigación, Facultad de Ciencias, Universidad Nacional Autónoma de México, campus Juriquilla) for his assistance with photographs edition. We also thank Dirección General de Vida Silvestre (Secretaría del Medio Ambiente y Recursos Naturales) who give us the field collecting bats permits numbers SGPA/DGVS/13206 and SGPA/DGVS/05181. For their comments on a draft of the manuscript, we express our appreciation to Ricardo Guerrero (Laboratorio de Biología de Vectores y Parásitos, Centro de Ecología y Evolución, Instituto de Zoología y Ecología Tropical, Facultad de Ciencias, Universidad Central de Venezuela), Margarita Ojeda (Laboratorio de Ecología y Sistemática de Microartrópodos, Facultad de Ciencias, Universidad Nacional Autónoma de México), and Ángela Nava Bolaños (Unidad Multidisciplinaria de Docencia e Investigación, Facultad de Ciencias, Universidad Nacional Autónoma de México, campus Juriquilla, Querétaro). We are grateful to the anonymous referees and the Editor who suggesting essential details that helped improve this paper. Financial assistance was provided by DGAPA, UNAM, grants No. IN226010, and IN219113 to JBMM. Additionally, This work was supported by Universidad Nacional Autónoma de México (UNAM), Programa de Apoyos para la Superación del Personal Académico (PASPA), Dirección General de Asuntos del Personal Académico (DGAPA), for the financial assistance to the first author during his sabbatical leave (at RBINS: 2024).

## ORCID

Juan B. Morales-Malacara ID https://orcid.org/0000-0002-9828-5176

Naomi Vázquez-Xicoténcatl ID https://orcid.org/0009-0002-7421-6720

Carlos García-Estrada ID https://orcid.org/0000-0002-4965-9426

Helisama Colín-Martínez ID https://orcid.org/0000-0002-7426-7264

Cenia E. Almazán-Marín ID https://orcid.org/0000-0003-0367-4492

Hugo A. Valdes-Saucedo ID https://orcid.org/0009-0003-8804-5456

Hugo A. Ruiz-Piña ID https://orcid.org/0000-0002-3080-1752

Alan Cuxim-Koyoc ID https://orcid.org/0000-0002-5449-8954

Enrique Reyes-Novelo ID https://orcid.org/0000-0001-9526-5033
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