Bee Mites - Family Cheyletidae: Cheletophyes, Nodele, Eucheyletia

mite Cheletophyes panamensis (Cheyletidae) female ex nest of Xylocopa frontalis

Fig. 1. Cheletophyes panamensis (Cheyletidae) female: dorsal view, nest of Xylocopa frontalis in Costa Rica. Click here to enlarge

Bee Mites : Acari : Acariformes : Trombidiformes

Family Cheyletidae Leach, 1815

This cosmopolitan family includes more than 370 described species and 72 genera (Bochkov and Fain, 2001a; Bochkov, 2003, pers. comm.). Some species are predators occurring in soil, forest litter, under tree bark and on foliage, in spore tubes of polypore fungi, nests of birds, mammals or insects, quills of birds, grain storages, and house dust. Several species are ectoparasites of different terrestrial vertebrates.
Females of some cheyletid genera were found phoretic on different insects, including bees: Cheletophyes Oudemans, 1914 on Xylocopa spp. (Hymenoptera: Apidae), Nodeleon Aztecanthidium tenochtitlanicum (Hymenoptera: Megachilidae), Samsinakia on Coleoptera: Tenebrionidae, Hypopicheyla Volgin, 1969 on Aradus (Hemiptera: Aradidae) and Tenebrionidae, Cheyletia Haller, 1884, Cheletophanes Oudemans, 1904, and Cunlifiella Volgin, 1964 on Aradus, and Paracaropsis Volgin, 1969 on Laphria and Pagidolaphria (Diptera: Asilidae) (Bochkov and Fain, 2001a; Bochkov and Mironov, 1998; Klimov, 1997; Volgin, 1969; our data).
There are three genera associated with bees in North America: Cheletophyes, Nodele, and Eucheyletia. Species of the genus Cheletophyes are obligatory associated with large carpenter bees (Xylocopa), whereas the level of association of Nodele sp. and Eucheyletia bakeri (found in a bumble bee nest) with their hosts is unknown.

Genus Cheletophyes Oudemans, 1914

Cheletophyes Oudemans, 1914: 101 (Type species Cheletophyes vitzthumi Oudemans, 1914 by monotypy); Baker, 1949: 287 (part.); Volgin, 1969: 160; Summers and Price, 1970: 40; Fain and Bochkov, 2001a: 61.
Cheletes: Vitzthum, 1919: 2 (part.).
Cheyletus: Baker, 1949: 275 (part.); Volgin, 1969: 73 (part.); Summers and Price, 1970: 23 (part.).

The genus Cheletophyes is distributed in the Oriental, Afrotropic, and Neotropic realms with eight species obligatory associated with bees of the genus Xylocopa (several records from other hosts are probably accidental) (Fain and Bochkov, 2001a,b). All stages of these mites are predators of different microarthropods feeding on provision in the bees' nests (Eickwort, 1994). Females of Cheletophyes disperse in mesosomal acarinaria of adult bees. According to the hypothesis of OConnor (1993), the relationships between Cheletophyes and their bee hosts are mutualistic, and the bees have developed mesosomal acarinaria to transfer these predacious mites controlling nest cleptoparisites. Our study (Klimov, Bochkov, OConnor, in press) suggests that despite the fact that the mites do not directly depend on resources provided by the bees, they are highly specific to their hosts.
One species, Cheletophyes panamensis, has been recorded for North America.

Cheletophyes panamensisKlompen, Méndez et Lukoschus, 1984

Cheletophyes panamensis Klompen et al., 1984: 249, Figs 1-5; Fain and Bochkov, 2001a: 63.

Material. 1 female - Costa Rica, nest of Xylocopa frontalis, UMMZ BMOC 03-1228-001.
Biology. The species was originally found in the nest of Xylocopa frontalis constructed in the tree Luehea seemannii Triana & Planch., 1862 (Malvales: Tiliaceae).
Distribution. Panama (type locality), Costa Rica (Klompen et al., 1984; our data).
Hosts. Xylocopa (Megaxylocopa) frontalis (Olivier, 1789) (Klompen et al., 1984).

mite Nodele (Aztecocheyletus) conquistador

Fig. 2. Nodele sp. (Cheyletidae): female, dorsal view, ex Aztecanthidium tenochtitlanicum (Hymenoptera: Megachilidae) from Mexico. Click here to enlarge

Genus Nodele Muma, 1964

Nodele Muma, 1964: 252 (type species Nodele calamondin Muma, 1964 by original designation); Volgin, 1969: 161; Fain and Bochkov, 2001a: 54.
Neocheletophyes Volgin, 1965: 296 (type species Cheletophyes philippinensis Baker, 1949 by original designation). Synonymized Volgin,1969.
Cheletophyes: Baker, 1949: 287 (part.).

This genus includes five species collected from forest litter, under bark, on bostrichid beetles, galleries of bark beetles, nest of sparrow (Passer), and grain in the Holarctic, Oriental, and Afrotropic realms (Fain and Bochkov, 2001a, Kuznetsov, 1977; Thewke and Enns, 1968; Wafa and Soliman, 1968).
Biology is only known for Nodele calamondin. It is a predator of other mite species and of small insect larvae. Unmated females produced only male offspring (arrhenotoky), which subsequently mated with their mothers to produce female as well as male offspring. Like other representatives of the family, males have one nymphal stage in their development and females have two nymphal stages. Males guarded the quiescent deutonymph females and mating took place as soon as the females emerged as adults. Unmated females lived more than twice as long as mated females and laid about 89% of the number of eggs laid by their mated sisters. The average duration of a generation was 62 days at 25 degrees C and 34 days at 29.5 degrees C. Net reproduction rate (ratio of female births in two successive generations) was 20 at 25 degrees C and 34 and 29.5 degrees C (after Barker, 1992).
We have a single record of three females of an undescribed species of Nodele collected from propodeum of the megachilid bee Aztecanthidium tenochtitlanicum Snelling, 1987. The mites have several substantial differences from the nominal genus. A new subgenus will be created for the new species.

Nodele sp.

Material. 3 females MEXICO: Jalisco, Chamela, Estacion Biologica, ex propodeum of Aztecanthidium tenochtitlanicum Snelling, 1987 (Hymenoptera: Megachilidae) on Croton sp. (Euphorbiales: Euphorbiaceae), 10-14 July 1989. Coll. T. Griswold BMOC 03-1003-043.
Description. Two pairs of setae of series E, e1 and e2. All dorsal idiosomal setae clavate. Protegmen bears a pair of well-developed lateral teeth. Propodonotal shield transversely serrate-striated. Tibia I relatively short. Tarsus I, excluding pretarsus/tibia I 2.3 : 1.0. Solenidion w1 short, 4 times shorter than seta ft, situated far from guard seta.
Biology. We believe that relationships of our species with the bee, like in other species of the genus Nodele, depend simply on availability of a host in the mite environment. In other words, it is not host specific.
Distribution. Mexico: Jalisco.
Hosts. Aztecanthidium tenochtitlanicum.

Fig. 3. Eucheyletia bakeri (Cheyletidae): female, dorsal view (left) and gnathosoma, dorsal view (right) (modified from Baker, 1949)

Genus Eucheyletia Baker, 1949

Eucheyletia Baker, 1949: 294 (type species Eucheyletia bishoppi Baker, 1949 by original designation); Volgin, 1969: 120; Fain and Bochkov, 2001a: 48.

This genus includes ten species distributed in the Holarctic and Oriental realms. It is primarily associated with nests of rodents. The genus also has been found in nests of other small mammals and birds, on oat seeds, barn and house dust (Fain and Bochkov, 2001; Volgin, 1969). One species, Eucheyletia bakeri, has been described from the nest of a bumble bee (Baker, 1949; Volgin, 1963).

Eucheyletia bakeri Volgin, 1963

Eucheyletia bakeri Volgin, 1963: 62, Figs 3, 5-7; Volgin, 1969: 149, Figs. 143-145; Fain and Bochkov, 2001a: 49.
Eucheyletia harpyia: Baker, 1949: 294, Figs. 76-79.

Biology. Three females of this species have been collected from the nest of a bumble bee (Baker, 1949). This record is probably accidental because it has not been confirmed and this type of habitat is unusual for the genus. On the other hand, there are mite genera (Kuzinia, Parasitellus) that occur both in rodent and bumble bee nests. This may be explained by the fact that bumble bees often construct their nests in borrows of rodents.
Distribution. USA: Alaska (Baker, 1949).
Hosts. Bombus sp. (Baker, 1949).

References

Baker E. W. 1949. A review of the mites of the family Cheyletidae in the United States National Museum. Proceedings of the United States National Museum. 99(3238): 267-320.

Barker P. S. 1992. Bionomics of Nodele calamondin Muma (Acarina: Cheyletidae) fed on Lepidoglyphus destructor (Schrank) (Acarina: Glycyphagidae) at two constant temperatures. Canadian Journal of Zoology. 70(12): 2333-2337.

Bochkov A. V., Mironov S. V.1998. Samsinakia trilobitus sp. n., a new cheyletid mite from South India (Acari: Cheyletidae). Entomologishe Mitteilungen aus dem zoologishen Museum Hamburg, 12(157): 265-268.

Bochkov A., Fain A. 2001. Phylogeny and system of the Cheyletidae (Acari: Prostigmata) with special reference to their host-parasite associations. Bulletin de l'Institut royal des sciences naturelles de Belgique. Entomologie. 71: 5-36.

Eickwort G. A. 1994. Evolution and life-history patterns of mites assciated with bees. In: Mites: Ecological and Evolutionary Patterns (ed. by M. A. Houck), Chapman & Hall, New York, London. pp. 218-251.

Fain A., Bochkov A. V. 2001a. A review of some genera of cheyletid mites (Acari: Prostigmata) with descriptions of new species. Acarina. 9(1): 47-95.

Fain A., Bochkov A. V. 2001b. A review of the genus Cheyletus Latreille, 1776 (Acari: Cheyletidae). Bulletin de l'Institut royal des sciences naturelles de Belgique. Entomologie. 71: 83-114.

Klimov P. B. 1997. [Data on the biology and systematics of Paracaropsis travisi (Acari, Cheyletidae), a new mite for Russia] / Fundamental'nye problemy okhrany okruzhayushchey sredy. Vol. 1. 9-10 Dec 1997, Vladivostok. P. 9-11. [in Russian]

Klimov P. B., Bochkov A. V., OConnor B. M. Host specificity and multivariate diagnostics of cryptic species in predacious cheyletid mites of the genus Cheletophyes (Acari: Cheyletidae) associated with large carpenter bees. Systematic Entomology [in press]

Klompen J. S. H., Méndez E., Lukoschus F. S. 1984. A new species of the genus Cheletophyes Oudemans, 1914 (Prostigmata: Cheyletidae) from the nest of a carpenter bee in Panama. Acarologia. 25(3): 249-251.

Kuznetsov N. N. 1977. [Cheyletid mites (Acariformes, Cheyletidae) of Crimea]. Entomologicheskoe Obozrenie. 56(4): 923-928. [in Russian, translated: Kuznetzov N. N. 1977(1978). The cheyletid mites (Acariformes: Cheyletidae) of the Crimea. Entomological Review. 77(4): 149-152.]

Muma M. H. 1964. Cheyletidae (Acarina: Trombidiformes) associated with Citrus in Florida. The Florida Entomologist. 47(4): 239-253.

OConnor, B.M. 1993. The mite community associated with Xylocopa latipes (Hymenoptera: Anthophoridae: Xylocopinae) with description of a new type of acarinarium. International Journal of Acarolology 19(2): 159-166.

Oudemans A. C. 1914. Acarologische Aanteekeningen LIV. Entomologische Berichten. 4(78): 101-103.

Summers F. M., Price D. W. Review of the mite family Cheyletidae. University of California Publications in Entomology. 61: 1-153.

Thewke S. E. , Enns W. R. 1968. A new species of predacious mite (Acarina: Cheyletidae) from galleries of bark beetles in Missouri. Acarologia. 10(2): 215-219.

Vitzthum H. G. 1918 (1920). Acarologische Beobachtungen. Zweite Reihe. Archiv für Naturgeschichte. 84A(6): 1-40.

Volgin V. I. 1963. [Materials on systematics of predatory mites of the family Cheyletidae (Trombidiformes)]. Parazitologicheskiy Sbornik. 21: 44-68.

Volgin V. I. 1965. [On the artificial nature of the genus Cheletophyes Oudms. (Acarina, Cheyletidae)]. Trudy Zoologicheskogo instituta. 35: 288-299. [in Russian]

Volgin V. I. 1969. Kleshchi semeystva Cheyletidae mirovoy fauny. Nauka, Leningrad. 1-432. [in Russian, English translation: Volgin V. I. 1987. Acarina of the Family Cheyletidae of the World. Amerind Publishing Co., New Delhi. 1-532].

Wafa A. K., Soliman Z. R. 1968. Five genera of family Cheyletidae (Acarina) in the U.A.R. with a description of four new species. Acarologia. 10(2): 220-229.