Chaires-Grijalva, Martha Patricia ¹ ; Estrada-Venegas, Edith Guadalupe ² ; Equihua-Martínez, Armando ³ ; Moser, John C.⁴ ; Blomquist, Stacy R. ⁵ ; Pérez-Silva, Mauricio ⁶ ; Arriola-Padilla, Víctor Javier ⁷ ; Rodríguez-Ortega, Alejandro ⁸ and Kontschán, Jenő ⁹

¹✉ Unidad Académica Multidisciplinaria Mante, Universidad Autonoma de Tamaulipas, Mexico.
²Programa de Entomología y Acarología. Colegio de Postgraduados. Campus Montecillo. Mexico.
³Programa de Entomología y Acarología. Colegio de Postgraduados. Campus Montecillo. Mexico.
⁴Alexandria Forestry Center, Forest Service. United States Department of Agriculture, USA. Deceased.
⁵Alexandria Forestry Center, Forest Service. United States Department of Agriculture, USA.
⁶Instituto de Biología, Universidad Nacional Autonoma de México. Mexico.
⁷Centro Nacional de Investigación Disciplinaria en Conservación y Mejoramiento de Ecosistemas Forestales, Instituto Nacional de Investigaciones Forestales Agrícolas y Pecuarias. Mexico.
⁸Ingeniería en Agrotecnología. Universidad Politécnica de Francisco I. Madero. Mexico.
⁹Széchenyi István University. Győr. Hungary.

2026 - Volume: 66 Issue: 2 pages: 364-376

Original research

Keywords

Abstract

Introduction

Mites of the superfamily Uropodoidea are common inhabitants of soil and forest litter. They are predominantly found in microhabitats rich in decaying organic matter, such as decomposing plant material, manure, and the nests of insects, birds, and mammals (Błoszyk, 1999; Błoszyk et al., 2006; Gwiazdowicz et al., 2015; Karg, 1989; Mašán, 2001; Mašán and Krištofík, 1995; Wiśniewski and Hirschmann, 1993). This diverse group includes 47 families and over 300 currently recognized genera (Halliday, 2016, 2019). With a complete life cycle up to one year, the deutonymph stage is particularly critical for species persistence due to its remarkable resilience to adverse environmental conditions and its capacity for phoresy (Athias-Binche, 1984).

Phoretic interactions between mites and bark-dwelling insects have been documented by several authors (e.g., Gwiazdowicz et al., 2011, 2012; Moser, 1976; Pernek et al., 2012). Notably, some deutonymphs exhibit striking morphological adaptations for phoresy, most spectacularly exemplified by the presence of an anal pedicel. These modified deutonymphs attach to their insect host by pressing their slightly enlarged anal openings against a smooth surface on the host's body or legs. They then expel a sticky, liquid secretion that hardens upon air exposure, forming this pedicel for secure attachment (Evans, 1972).

These relationships can be opportunistic or exhibit strong host specificity, mediated by unique physical, chemical, and behavioral cues (Moser et al., 2005; Walter and Proctor, 2013). Trematurid mites are frequently found associated with or directly on bark beetles (Coleoptera: Curculionidae: Scolytinae) (Chaires-Grijalva, 2013; Chaires-Grijalva et al., 2016; Knee et al., 2012; Quiroz-Ibáñez et al., 2016), which provide the primary means of dispersal for deutonymphs. While the influence of mites on bark beetle populations is not yet fully understood, their high relative abundance and diversity suggest a potential impact on beetle dynamics and the physical condition of their hosts (Chaires-Grijalva, 2013; Halliday, 2019).

The family Trematuridae represents the primary group of phoretic mites associated with bark beetles. This family exemplifies the global nature of these associations, with 20 species recorded from 21 countries (Table 1). Their extensive distribution and phoretic associations underscore the ecological plasticity of these mites across diverse ecosystems. However, much of the foundational research on Uropodina has historically focused on temperate zones of Europe and North America. This geographic bias has created a significant knowledge gap regarding the diversity and ecology of phoretic Trematuridae within tropical and subtropical regions.

Our study, conducted across the diverse climatic range of Mexico, contributes to filling this void by documenting new records and attachment sites in the Neotropical and Nearctic transition zones. The main objective of this study is to assess the diversity of Trematuridae mites associated with bark beetles across various states in Mexico. To ensure taxonomic accuracy, we relied on recent comprehensive revisions (e.g., Halliday, 2016; Knee et al., 2012; Kontschán et al., 2019), particularly regarding the complex historical nomenclature established by Hirschmann and collaborators.

Materials and methods

Study area and sampling

Sampling was conducted across 24 Mexican states between 2009 and 2023. At each site, samples were obtained from both live and dead trees exhibiting signs of bark beetle infestation. The sampling effort covered Aguascalientes, Baja California, Ciudad de México, Chiapas, Chihuahua, Coahuila, Colima, Durango, Guanajuato, Hidalgo, Jalisco, México, Michoacán, Morelos, Nuevo León, Oaxaca, Puebla, Querétaro, San Luis Potosí, Sinaloa, Sonora, Tlaxcala, Veracruz, and Zacatecas. Bark and log samples were transported in plastic bags to the Forest Entomology Laboratory at Colegio de Postgraduados. Bark samples were examined under a Carl Zeiss Stemi DV4 stereomicroscope, while logs were placed in climate-controlled chambers to collect emerging adult insects, which were subsequently processed for phoretic mites. Detailed georeferenced coordinates for all collection sites are provided in the Results section.

Bark beetle identification

Bark beetles and associated arthropods were collected from bark and log samples using forceps and preserved in 70% ethanol. Specimens were identified using the taxonomic keys of Cibrián et al. (1995) and Wood (2007). Identifications were confirmed by Ph.D. Armando Equihua-Martínez and Mauricio Pérez-Silva.

Mite processing and identification

Phoretic mites were carefully removed from each beetle specimen and transferred to a lactic acid solution for clearing. Specimens were then mounted on glass slides using Hoyer's medium for microscopic examination. In addition to phoretic individuals, mites found within bark beetle galleries were also collected. The identification of Trichouropoda species was primarily conducted using the keys provided by Karg (1989), supplemented by specific original descriptions from the complex literature of Hirschmann and collaborators, and confirmed by the late co-author, Dr. John C. Moser^†. It is noteworthy that several species formerly assigned to Trichouropoda, such as Oodinychus hirsuta and O. ovalis, have been reassigned following recent taxonomic revisions (e.g., Halliday, 2016; Kontschán et al., 2019). This study adopts these updated designations to maintain nomenclatural consistency with current uropodid systematics All mite specimens were deposited in the Acarological Collection at Colegio de Postgraduados (COLPOS).

Results

A total of 1,713 bark beetles belonging to nine species were examined. From these, 337 mites of the family Trematuridae (genus Trichouropoda) were collected: 178 were found phoretically attached to the insect bodies, 104 were recovered from the galleries, and 55 were retrieved from the alcohol sediments of the collection vials. Eleven species of Trichouropoda were identified.

Out of the 24 Mexican states sampled, trematurid mites were recorded in ten: Baja California (BC), Chihuahua (Chih.), Estado de México (EMex), Hidalgo (Hgo.), Jalisco (Jal.), Michoacán (Mich.), Puebla (Pue.), Querétaro (Qro.), Tlaxcala (Tlax.), and Veracruz (Ver.) (Table 2).

The distribution of Trichouropoda species across host beetles varied significantly. The highest mite abundance was observed on Dendroctonus rhizophagus (n=44) and D. valens (n=45), which accounted for 25.58% and 18.29% of the total associations, respectively. In contrast, D. mexicanus showed the lowest proportion of associated mites (1.57%) despite having the largest sample size of beetles (n=572).

In terms of host specificity, Trichouropoda polytricha was the most generalist species, associated with six different bark beetle species (D. frontalis, D. mexicanus, D. pseudotsugae, D. rhizophagus, D. valens, and Ips confusus) across eight states. Conversely, species such as T. fallax and T. hondurasae were restricted to a single host and a single state in this study (Veracruz and Querétaro, respectively).

Furthermore, the genus Ips also hosted a considerable number of mites, with I. bonanseai showing a 20.51% association rate, primarily with T. australis (n=24).

Regarding overall species associations, five mite species were collected from two or more bark beetle hosts. In contrast, six species (T. fallax, T. hondurasae, and the undescribed T. sp. 2, T. sp. 3, T. sp. 4, and T. sp. 5) were associated with a single host species, although these were represented by only a few specimens each (Table 2; Figure 1).

Trichouropoda polytricha displayed no clear host preference, as was found on seven of the nine beetle species examined. In contrast, the remaining trematurid species exhibited minimal overlaps in host associations within this study; this suggests relative host specificity, despite previous literature reports characterizing them as generalists (Chaires-Grijalva et al., 2019). Most mites were observed on the ventral surface (29%), gular area (22%), abdominal sternites (18%), elytra (17%), elytral declivity (11%), legs (7%), and pronotum (2%) (Figure 2). Overall, mites were primarily detected on the host body and within the bark beetle galleries (Table 3).

Mites were found in diverse microhabitats associated with their hosts. Of the total specimens collected, 178 were found directly on the bark beetles, 104 within the galleries, and 55 in alcohol sediments. Regarding phoretic attachment, specific body sites were preferred depending on the mite species. For instance, Trichouropoda polytricha and Oodinychus ovalis (the most abundant species) were frequently located in the gular zone (Gz) and on the elytra (Oe). Other species, such as T. australis, showed a preference for abdominal sternites (As) and the pronotum (Pr). Additionally, mites were recovered from both new (Ng) and old (Og) galleries, primarily under humid (H) conditions.

Organisms examined

The specimens analyzed in this study were obtained from 24 states across Mexico between 2009 and 2023. A total of 1,713 bark beetles were examined, from which phoretic trematurid mites were recovered (Figure 3). For each species listed below, the data includes the specific locality, geographic coordinates (where available), collection date, host insect species, and the number of mite individuals (deutonymphs) collected. All specimens were collected by the authors and have been deposited in the Acarological Collection of the Colegio de Postgraduados (COLPOS). The taxonomic arrangement follows recent nomenclature, including the reassignment of certain species to the genus Oodinychus.

Oodynichus ovalis (Koch, 1939)

Notaspis ovalis Koch, 1839

Chihuahua: La Laja, Bocoyna, Pinus arizonica, 2452 masl, 27.931916 N, 107.5985 W; Jun. and Jul. 2008 to 2011, 19 DN ex. on gular zone of D. rhizophagus, 9 DN ex. in new and humid galleries of D. rhizophagus; Cuesta Prieta, Ejido San Juanito, Bocoyna, Pinus arizonica, 2450 masl, 27.930555 N, 107.595999 W, 16 DN ex. on gular zone of D. valens, 18 DN ex. galleries under bark. Estado de México: Experimental Station Zoquiapan, Universidad Autónoma de Chapingo, Ixtapaluca, P. hartwegii, 3600 masl. 19.282225 N, 98.670701 W, May 2006, 1 DN ex. on elytra of D. adjunctus. Tlaxcala: Highway Texcoco-Calpulalpan, 2842 masl., 19.5601027 N., 98.6944138 W., Jun. 16th, 2012. 5 DN ex. on elytral declivity of D. valens. colls. M.P. Chaires-Grijalva,

Trichouropoda adjuncti Wisniewski & Hirschmann, 1988

Chihuahua: Mesa de Parra, Municipality Madera, Pinus engelmannii, 2 481 masl, 29.204424 N, 108.184462 W, 1DN ex. elytral declive Dendroctonus valens. Jalisco: Rancho los Zuno, Gómez Farias, 2 200 msnm, 19.886659 N, 103.409291 W.; 3DN ex. on elytral declive of D. valens. Veracruz: Ejido Forestal Los Pescados, Municipality Perote; 3 128 masl. 19.528888 N, 97.123333 W; May 2004 to Apr. 2005; 4 DN ex. on elytra of D. adjunctus, 2 DN on legs and 2 DN ex. old and dry galleries of D. adjunctus. colls. M.P. Chaires-Grijalva, A. Rodríguez.

Trichouropoda australis Hirschmann, 1972

Estado de México: Experimental Station Zoquiapan, Universidad Autónoma de Chapingo, Municipality Ixtapaluca, P. hartwegii, 3600 masl. 19.282225 N, 98.670701 W; 7DN ex. on abdominals sternites of D. frontalis, 2 DN ex. new and humid galleries of D. frontalis; 24 DN ex. abdominal sternites of Ips bonanseai, 1♀, 12 DN ex. galleries under bark. Jalisco: Rancho los Zuno, Gómez Farias, 2200 msnm, 19.886659, 103.409291; 2DN ex. on abdominals sternites of Ips calligraphus, coll. M.P. Chaires-Grijalva.

Trichouropoda fallax (Vitzthum, 1926)

Trichoobscura fallax Hirschmann & Zirngiebl-Nicol, 1961

Uropoda fallax Vitzthum, 1926

Estado de México: Experimental Station Zoquiapan, Universidad Autónoma de Chapingo, Municipality Ixtapaluca, P. hartwegii, 3600 masl. 19.282225 N, 98.670701 W, May 2006, 2♀ ex. old and humid galleries of D. adjunctus. Veracruz: Ejido Forestal Los Pescados, Municipality Perote; P. hartwegii, 3128 masl. 19.528888 N, 97.123333 W; May 2004 to Apr. 2005; 8 DN ex. on elytra of D. mexicanus, 3 DN ex. galleries of D. mexicanus.

Trichouropoda hondurasae (Hirschmann & Wisniewski, 1986)

Ipiduropoda hondurasae Hirschmann & Wisniewski, 1986

Querétaro: Rio Blanco, Municipio de Peñamiller, 1720 masl, 21.208056 N, 99.737500 W; Nov 26th, 2010; 5 DN ex. on elytra of D. frontalis.

Trichouropoda polytricha (Vitzthum, 1923)

Ipiduropoda polytricha Hirschmann 1986

Uropoda polytricha Vitzthum, 1923

Baja California: Parque Nacional Sierra de San Pedro Mártir, Ensenada. Pinus jeffreyi, 2830 masl, 31.027794 N, 115.467846 W., Sept. 2006, 3 DN ex. ventral sternites of D. pseudotsuga. Chihuahua: La Laja, Bocoyna, P. arizonica, 2452 masl, 27.931916 N, 107.5985 W.; Jun. and jul. 2008 to 2011, 8 DN ex. on ventral sternites of D. rhizophagus; Cuesta Prieta, Ejido San Juanito, Bocoyna, P. arizonica, 2450 masl, 27.930555 N, 107.595999 W., 5 DN ex. on gular zone of D. rhizophagus; Mesa de Parra, Madera, P. arizonica, 2300 masl., 29.187027, -108.171007, 6 DN ex. on ventral sternites of D. rhizophagus. Jalisco: Parque Nacional Nevado de Colima, San Gabriel, Pinus hartwegii, 3300 masl, 19.626806, 103.562139, 8 DN ex. on elytral declivity of Ips calligraphus. Michoacán: Zirahuén Lake, Salvador Escalante, Pinus patula, 2187 masl, 19.466111 N, 101.736336 W, Jan. 14th, 2010, 13 DN ex. on ventral sternites of D. valens, 3♀, 1♂, 27 DN, 9 PN ex. old and humid galleries of D. valens. Puebla: El puerto, Tetela de Ocampo, P. patula, 2120 masl, 19.805075 N, 97.779233 W., 1 DN ex. on ventral sternites of D. frontalis. Querétaro: Rio Blanco, Peñamiller, P. patula, 1720 masl, 21.208056 N, 99.737500 W; Nov 26th 2010; 1 DN ex. on ventral sternites of D. mexicanus. Veracruz: Ejido Forestal Los Pescados, Perote; P. hartwegii, 3128 masl. 19.528888 N, 97.123333 W; May 2004 to Apr. 2005; 1 DN ex. on ventral sternites of D. frontalis, 5 DN ex. on ventral sternites of D. valens. colls. M.P. Chaires-Grijalva, A. Rodríguez.

Trichouropoda sp. 1

Estado de México: Experimental Station Zoquiapan, Universidad Autónoma de Chapingo, Ixtapaluca, P. patula, 3600 masl. 19.282225 N, 98.670701 W; 7 DN ex. on elytra of D. frontalis. Tlaxcala: Highway Texcoco-Calpulalpan, 2842 masl., 19.5601027 N., 98.6944138 W., Jun. 16th, 2012, 2 DN ex. on elytral declivity of D. valens.

Trichouropoda sp 2

Chihuahua: La Laja, Bocoyna, P. arizonica, 2452 masl, 27.931916 N, 107.5985 W.; Jun. and Jul. 2008 to 2011, 6 DN ex. on ventral sternites of D. rhizophagus*.

Trichouropoda sp 3

Estado de México: Experimental Station Zoquiapan, Universidad Autónoma de Chapingo, Ixtapaluca, P. patula, 3600 masl. 19.282225 N, 98.670701 W, May 2006, 1 DN ex. on elytra of D. frontalis. Hidalgo: Presa el Tejocotal, Acaxochitlan, 2260 masl., 20.133055 N, 98.129749 W, Oct 10th, 2022; 4 DN ex. old and humid galleries of D. valens.

Trichouropoda sp 4

Jalisco: Rancho los Zuno, Gómez Farias, 2200 msnm, 19.886659 N, 103.409291W, Nov 13th, 2023; 12 DN ex. in old and humid galleries of Ips calligraphus, coll. M.P. Chaires-Grijalva; 7 DN ex. on elytra of Hylurgops incomptus.

Trichouropoda sp 5

Veracruz: Ejido Forestal Los Pescados, Perote; P. hartwegii, 3128 masl. 19.528888 N, 97.123333 W; May 2004 to Apr. 2005, 7 DN ex. on ventral surface of Hylurgops incomptus.

Discussion

Biogeography and host plasticity. Six of the eleven mite species identified in this study are also present in Europe and North America, suggesting a high capacity for co-invasion with their host insects. This distribution pattern likely stems from the native status of both beetles and mites across these continents. For instance, Trichouropoda australis has been documented on 17 species of Dendroctonus and Ips across the United States and Canada (Hofstetter, 2008; Knee et al., 2012), while T. fallax is known to associate with seven species of Dendroctonus, Hylastes, and Hylurgops in Europe and North America (Hirschmann and Wiśniewski, 1989; Knee et al., 2012).

Notably, these mites are not obligate associates of bark beetles. O. ovalis has been recorded in diverse habitats, including bird nests in Poland and Scandinavia (Błoszyk et al., 2006), reflecting broad ecological plasticity. Similarly, T. polytricha (the most generalist species in our study, found on seven different bark beetle species) is a cosmopolitan mite previously recorded in Mexico (Moser et al., 1989). Its success across eight Mexican states suggests that a lack of host-sex preference and high host plasticity broaden the range of available carriers, increasing the probability of successful dispersal and colonization of new habitats (Bajerlein et al., 2023).

Mechanisms of phoresy and attachment sites. The selection of specific attachment sites by phoretic deutonymphs is a strategic adaptation to minimize active eviction by the host or passive dislodgment during flight (Chaires-Grijalva, 2013). Unlike most mites, uropodines utilize a flexible and resistant anal pedicel for anchorage, significantly reducing the likelihood of dislodgment (Bajerlein et al., 2013). Our results show a preference for the ventral surface (29%) and the gular area (22%), consistent with findings by Hunter (1993), who noted that uropodids typically attach to areas posterior to coxa III.

These sites likely offer protection from the beetle's grooming behavior. However, when primary sites become saturated, deutonymphs adhere to other areas, such as the elytra or pronotum. We observed that uropodine deutonymphs tend to occupy setae-uncoated body parts not yet colonized by other mite taxonomic groups (Bajerlein et al., 2013). It is worth noting that some mites found in alcohol sediments likely originated from these sites but were dislodged during sampling, a common limitation in phoretic studies.

The gallery as a micro-ecosystem. Mites found within bark beetle galleries are integral components of the associated fauna. Each gallery represents a complex micro-ecosystem where mites find essential resources, including nematodes, fungi, and bark beetle eggs or larvae (Moser, 1975; Kontschán et al., 2019). Humid, newly formed galleries support the mites' entire biological cycle by providing stable temperatures and refuge from natural enemies (Chaires-Grijalva et al., 2013).

The high prevalence observed in Dendroctonus rhizophagus (25.58%) compared to D. mexicanus (1.57%) is particularly striking. This disparity may be linked to the specialized biology of D. rhizophagus, which attacks the roots and base of seedlings; the increased humidity and relative stability of the subterranean environment may favor higher mite densities and more successful phoretic transfers compared to the more exposed subcortical environments of trunk-attacking species like D. mexicanus. According to Wilson's (1980) association/predation index, a closer phoretic relationship often correlates with a reduced predatory threat to the host (Moser, 1995). Consequently, the high abundance of certain Trichouropoda species suggests a well-established symbiotic equilibrium within the Mexican conifer forests.

Conclusions

The present study reveals a significant diversity of trematurid mites associated with bark beetles in Mexico, identifying eleven species within the genera Trichouropoda and Oodinychus. The presence of six species previously recorded in Europe and North America highlights a broad biogeographic distribution and suggests a robust capacity for co-invasion alongside their host insects.

Our findings demonstrate two distinct ecological strategies among the identified mites:

1. Host Plasticity: T. polytricha stands out as a highly successful generalist, exhibiting no clear host preference and a wide geographic range across ten Mexican states.
2. Specialization and microhabitat: Most species showed restricted host associations and specific phoretic attachment sites, particularly in protected areas like the gular zone and ventral surface. These locations are likely selected to minimize dislodgment and avoid host grooming behaviors.

Furthermore, the high prevalence observed in D. rhizophagus (25.58%) underscores the importance of the host's specific ecology—such as its subterranean habitat—in determining mite population densities. The consistent presence of these mites within both new and old galleries confirms that these structures serve as essential micro-ecosystems, providing the humidity, stable temperature, and food resources (nematodes, fungi, and larvae) necessary for the mites' entire biological cycle.

In conclusion, this research provides a fundamental baseline for understanding the symbiotic networks within Mexico's coniferous forests. Future studies should focus on the functional role of these mites—whether as commensals, predators, or competitors—to better assess their impact on bark beetle population dynamics and forest health.

Acknowledgements

To Dr. Guillermo Sánchez Martínez (INIFAP Campus Pabellón) and, Dr. Gerardo Zúñiga Bermúdez (IPN ENCB) for their help in providing the biological material for this study.

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