Morales-Malacara, Juan B. ¹ ; Vázquez-Xicoténcatl, Naomi ² ; García-Estrada, Carlos ³ ; Colín-Martínez, Helisama ⁴ ; Almazán-Marín, Cenia E. ⁵ ; Valdes-Saucedo, Hugo A. ⁶ ; Ruiz-Piña, Hugo A. ⁷ ; Cuxim-Koyoc, Alan ⁸ and Reyes-Novelo, Enrique ⁹

¹✉ Laboratorio 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, Querétaro, México.
²Laboratorio 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, Querétaro, México.
³Universidad 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.
⁴Universidad 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.
⁵Laboratorio 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, Querétaro, México.
⁶Laboratorio 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, Querétaro, México.
⁷Centro de Investigaciones Regionales "Dr. Hideyo Noguchi" Universidad Autónoma de Yucatán. Av. Itzaes por 59 No. 490 Centro, C.P. 97000, Mérida, Yucatán, México.
⁸Centro de Investigaciones Regionales "Dr. Hideyo Noguchi" Universidad Autónoma de Yucatán. Av. Itzaes por 59 No. 490 Centro, C.P. 97000, Mérida, Yucatán, México.
⁹Centro de Investigaciones Regionales "Dr. Hideyo Noguchi" Universidad Autónoma de Yucatán. Av. Itzaes por 59 No. 490 Centro, C.P. 97000, Mérida, Yucatán, México.

2025 - Volume: 65 Issue: 3 pages: 736-755

ZooBank LSID: CCBBD646-AF2E-481F-9E81-B42AD4A3EE43

Original research

Keywords

Abstract

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 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 saturatus Miller, 1897 from California, USA. Pinichpongse (1963) himself noted some morphological differences between the two species. However, later McDaniel (1970), after obtaining specimens of M. nigricans (Schinz, 1821) from Nicaragua, conducted a morphological 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. eptesicus, indicating undetermined or undescribed species, from the vespertilionid bats Eptesicus 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:

1. 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).
2. Alabidocarpus noctilio Fain, 1970, associated with Noctilio labialis (Kerr, 1792) from Suriname (Fain 1970b; 1973), is now Pseudoalabidocarpus noctilio (Fain, 1970) (Bochkov 2010).
3. 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 38^th species described in the genus, the 7^th 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.

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.

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

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).

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); d1, 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).

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 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 seta e is absent (Figure 4D).

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).

Male larva

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 PN (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.

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 diminute (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 (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 (φ and ω1), leg III stump with 4 tiny setae (φ, w, s and f), and leg IV stump with 3 tiny setae (φ, 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: φ tibia I, 2 (1.4–2, 1.7); ω1 tarsus I, 2.8 (2.8–3.14, 2.9); φ tibia II, – (1.51–1.54, 1.52 [n = 2]); ω1 tarsus II, – (2.11–2.45, 2.28 [n = 2]); φ 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); φ 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 Siences, 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 Colletion, 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–.36,a,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 associated with the Phyllostomid common vampire bat D. rotundus, whereas A. longipilus is found on Vespertilionid bats of the genus Myotis in the Americas.

It is noteworthy that no evidence of this chirodiscid species was found on the Atlantic slope in the state of Yucatán, México, or in the central region of México in the state of Querétaro, México, creating a paradox: Why?

A possible explanation is that there may be evidence of a disjunction in the ancient dispersal routes of at least two different populations of D. rotundus in the neotropical region of México, e.g., disjunct variations have been observed in the bat genus Chiroderma on both the Atlantic (Eastern Mexico) and Pacific slopes (Western Mexico) (Garbino et al. 2020). While A. ricardoguerreroi n. sp. is present on vampire bats from the Pacific slope, it is absent in populations of D. rotundus from the Atlantic or Gulf slopes, as well as from the central region of México.

Another potential explanation for this paradox could involve the close morphological similarities between A. ricardoguerreroi and A. longipilus, which could reflect certain evolutive affinity. Considering that some studies have occasionally found that D. rotundus and Myotis spp. share roosts in other regions of the country (Conde-Pérezprina et al. 2012), which could facilitate horizontal transmission of mites between both species of bats. Given that chirodiscid mites are permanent ectosymbionts, a speciation process may have begun based on the morphological characteristics of their host.

However, to confirm these hypotheses, it will be necessary to collect common vampire bats from other localities to determine whether Alabidocarpus ricardoguerreroi n. sp. is present or absent.

Key to species of the genus Alabidocarpus in the American continent

Females and Males

1. Legs IV abnormally long or very long, clearly larger than legs III, with rectangular tibiae longer than broad
...... 2

— Legs III and IV subequal, tibiae nearly square
...... 4

2. Posterolateral angle of prescapular plate, with a slightly rounded border with a short projection in the lower angle
...... A. nicaraguae

— Posterolateral angle of prescapular plate, with a conspicuous and large triangular projection directed towards seta c2
...... 3

3. Coxa IV with very long seta 4a almost the same size as the principal claw-like seta (f). Male with 2 pairs of opisthosomal setae
...... A. guyanensis

— Coxa IV with medium sized seta 4a shorter than principal claw-like seta (f). Male with 3 pairs of opisthosomal setae
...... A. jonesi

4. Posterior border of prescapular plate slightly straight or with a faint concavity and with small or short posterolateral projections, one near the dorsal edge and one in the lower angle
...... 5

— Posterior border of prescapular plate slightly curved, and generally sinuous, and without any posterolateral projections
...... 6

5. Prescapular plate on its posterior edge with very narrow width behind the base of the leg I. Seta c3 small to medium sized. Male with 4 pairs of opisthosomal setae
...... A. saccopteryx

— Prescapular plate on its posterior edge with wider width behind the base of leg I, that reaches beyond the middle level of the base of leg II. Seta c3 minute or inconspicuous. Male with 2 pairs of opisthosomal setae
...... A. furmani

6. Female with 3 pairs of opisthosomal setae, h1 minute, h2 long and h3 medium sized. Coxa IV with short, thick and spiniform seta 4a. Seta c3 minute
...... A. longipilus

— 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).

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