Acarological Studies Vol 1 (2): 152-160 RESEARCH ARTICLE Acarine biodiversity associated with bark beetles in Mexico M. Patricia CHAIRES-GRIJALVA 1 , Edith G. ESTRADA-VENEGAS 1,3 , Iván F. QUIROZ-IBÁÑEZ 1 , Armando EQUIHUA-MARTÍNEZ 1 , John C. MOSER 2† , Stacy R. BLOMQUIST 2 1 Colegio de Postgraduados, Carretera México-Texcoco Km.36.5, Montecillo, Texcoco, Estado de México C.P.56230. México 2 Southern Research Station, United States Department of Agriculture, Alexandria Forestry Center, 2500 Shreveport Hwy, 71360 Pineville, Louisiana, USA 3 Corresponding author: [email protected] Received: 10 December 2018 Accepted: 22 March 2019 Available online: 31 July 2019 ABSTRACT: The phloem of dying trees provides habitat for a large number of bark beetles and their associated mites. These mites depend on the scolitids for moving from one place to another, and directly or indirectly for their nutrition. In Mexico, there have been very few works on this topic. The first three studies in Mexico included isolated records of these associations, while the last three refer to new records for several states in the country. A total of 62 mites species were recorded in the present study. The most diverse order was Mesostigmata with 66% of the recorded species, followed by the suborder Prostigmata with 24% and the cohort Astigmatina with 10%. Trichouropoda polytricha (Vitzthum, 1923), Proctolaelaps subcorticalis (Lindquist, 1971), Proctolaelaps dendroctoni Lindquist and Hunter, 1965, Schizosthetus lyri- formis (McGraw and Farrier, 1969) and Dendrolaelaps neodisetus (Hurlbutt, 1967) were the most common species asso- ciated with bark beetles in this study. Dendroctonus frontalis Lindquist and Hunter, 1965 is the bark beetle with the high- est reported number of associated mites in Mexico and worldwide. Among the species mentioned in this study, there was an interesting range of feeding habitats and habits. The different associations among beetles and mites provide an inter- esting topic for future research. Keywords: Coleoptera, Dendroctonus, logs, natural forest, phoretic, symbiosis INTRODUCTION the host, and which represents an adapted state in which it remains until its biological cycle is synchronized with Among the bark beetles species, there are symbiotic in- the host. teractions, many of which are maintained through multi- ple factors, such as climate, predation and the use of re- Several groups of mites are associated with these insects, sources. Mites are important symbiotic organisms associ- forming dynamic, interspecific relationships. The rela- ated with bark beetles. More than one hundred species of tionships of mites and their hosts are defined by distinc- mites have been reported from bark beetles worldwide; tive patterns, which have been shaped by long evolution- some phoretic species are associated with adult beetles ary processes, involving a succession of interactive re- and other invertebrates on infested trees (Hofstetter et sponses, which resulted in different biorelationships al., 2013; Lindquist, 1969). Many of these mites depend (Hoffmann, 1988; Krantz and Walter, 2009). The im- on bark beetles or other insects to be transported inside portance of these relationships is that energy flows are the trees (Moser, 1985, Moser et al., 2010). Other studies created within the trophic networks and they contribute report other microorganisms such as protozoa and nema- to the structuring of the ecosystem. todes in these associations (Perotti and Braig, 2011). The galleries of bark beetles (Coleoptera: Curculionidae: Sco- Mite communities that live in the galleries of bark beetles lytinae) are located under the bark of trees, and all devel- are not only associated with these insects but there are opmental stages live there but other arthropods and mi- species that feed on fungi, leaf litter and other insects croorganisms live and feed there (Lindquist, 1975). (Kaliszewski, 1993; Walter and Proctor, 1999). There is an extensive body of literature, especially related to Den- The bark beetles are host species for a diverse community droctonus frontalis Zimmerman, 1868 (Kinn and of arthropods and microorganisms that are found subcor- Witcosky, 1978; Hofstetter, 2011; Hofstetter et al., 2013, tically, mainly mites. Lindquist (1975) stated that the Hofstetter et al., 2015), D. rufipennis Kirby, 1837 (Cardoza associations between mites and other arthropods can be et al., 2008), Ips typographus (Linnaeus, 1758) (Takov et based on a number of factors, namely: 1) the presence of al., 2009), Pityokteines spp. (Pernek et al., 2008, 2012), a stable habitat that guarantees the supply of food re- and Scolytus spp. (Moser et al., 2010, but species associat- sources and the protection of both organisms; 2) specifici- ed with ambrosia beetles are just beginning to be studied. ty for the habitat, in which the mite shows preference for Mites are good examples of phoretic organisms; mites the habitat and not for the host; 3) the mites specificity on associated with insects are transported to new habitats, the host; 4) specificity of the mites for a site on the host; in which they each play different ecological roles. and 5) synchrony of the life cycle of the mite with the insect, the cycle of the first one being shorter than that of The term phoresy (from the Greek phora: carry, have) MATERIAL AND METHODS defines one of the many kinds of association among ani- mals (Trägårdh, 1943). Mites are the main examples. Review of bark beetles collection. Visits were made to Farish and Axtell (1971) proposed a new definition: different collections of bark beetles and associated mites "Phoresy is a phenomenon in which an animal actively in Mexico and the USA, namely the Universidad Nacional looks for and lives on the outer surface of another animal Autónoma de México (UNAM) (National Collection of for a limited time, during which the linked animal (called Mites), Escuela de Ciencias Biológicas of Instituto Politéc- a phoretic) stops feeding and development, presumably nico Nacional (ENCB-IPN) (Mites collection ENCB-IPN), as a result of dispersal. This process is undoubtedly im- Universidad Autónoma Metropolitana (UAM), Univer- portant in mites’ evolution, which has followed different sidad Autónoma Chapingo (UACh) (Acarology Collection), pathways in the Mesostigmata, Prostigmata and Astigmat- Colegio de Postgraduados (CP) Campus Montecillo (Aca- ina (Cross and Bohart, 1969). The families in which phor- rology Collection), Instituto Nacional de Investigaciones esy occurs are Macrochelidae, Parasitidae, Laelapidae, Forestales, Agrícolas y Pecuarias (INIFAP) Campus Exper- Ascidae, Eviphididae and some Uropodines, and some imental Pabellón (Entomology and Acarology Collection), families from other groups, such as the Scutacaridae and and Entomology collection of Centro Nacional de Investi- Anoetidae (Athias-Binche, 1994; Reynolds et al., 2014) gación Disciplinaria en Conservación y Mejoramiento de Ecosistemas Forestales (CENID-COMEF). The United The bark beetles galleries provide habitat for a large States Forest Service, Southern Research Station in Pine- number of boring insects, mainly beetles and their associ- ville, Louisiana (USDA - SF) (USDA - FS Forest Mite Collec- ated mites. The best known mites are those associated tion) collection was also visited. In all of these collections, with the genera Dendroctonus, Ips, Tomicus, Hylesinus, an exhaustive review of mite species associated with bark Hylastes, Dryocoetes and Scolytus, each having 15 to 20 beetles. species of associated mites, of which 10 or 12 can be common in scolitid habitat (Hofstetter and Moser, 2014). Sampling. In addition, samples of bark and logs showing The relationship between the food webs in the galleries signs of damage by bark beetles (Scolytinae) were ob- and on the bark beetles can be complicated. The feeding tained from 24 states of Mexico from April 2007 to No- habits represented in galleries include fungivory, sa- vember 2017. The samples were from the following prophagy (Tarsonemus spp., Pygmephorus spp., Histio- states: Aguascalientes, Baja California, Ciudad de México, gaster spp., Histiostoma spp., among others), polyphagy Chiapas, Chihuahua, Coahuila, Colima, Durango, Guanajua- and nematophagy, as well as specialized predation in the to, Hidalgo, Jalisco, México, Michoacán, Morelos, Nuevo early stages of bark beetles (Digamasellus spp., Lasioseius Leon, Oaxaca, Puebla, Querétaro, San Luis Potosí, Sinaloa, spp.) or as monophagous or oligophagous parasites of Sonora, Tlaxcala, Veracruz and Zacatecas (Figure 1). The bark beetles larvae (Pyemotes spp., Paracarophenax spp. samples were transported in plastic bags and transferred and Iponemus spp.) (Lindquist, 1970; Cross and Moser, to Forest Entomology Laboratory at Colegio de Post- 1971). graduados, for mite revision under a stereoscopic micro- scope. The logs were placed in emergence chambers to These groups have different degrees of morphological and collect insect adults and then the mites were separated physiological adaptations. They are usually associated from them. with the insect nymphal stage. However, little is known about the biology of phoretic mite and the host. The dis- covery of a phoretic relationship guides us to a better understanding of the ecological role of both species (Binns, 1972). There are records of mites associated with scolytids from 15 countries. Of the total of 178 species of mites, 96 are included in the order Mesostigmata (52%), 55 in the sub- order Prostigmata (32%), 14 in the suborder Oribatida