Bee Mites: family Parasitidae (Acari): Parasitus, Parasitellus, Pergamasus Vulgarogamasus

mite Parasitellus talparum (Acari: Parasitidae) ex Bombus flavifrons from Wyoming

Fig. 1. Deutonymph of Parasitellus talparum (=Parasitus favus) (Acari: Parasitidae) ex Bombus flavifrons from Wyoming. Click here to enlarge

Bee Mites : Acari : Parasitiformes

Family Parasitidae Oudemans, 1901

This family is distributed worldwide and includes about 400 species grouped in 2 subfamiles: Parasitinae (9 genera) and Pergamasinae (20 genera) (Johnston, 1982; Tichomirov, 1977). Parasitid mites are essentially predatory and feed upon other microarthropodes, including their eggs, and on nematodes. They live in moss, forest litter, soil, dung, rotting seaweed, decaying organic substances, caves, and nests of small mammals and insects. These mites disperse during the deutonymph stage, usually on insects of the orders Coleoptera and Hymenoptera. All species of the genus Parasitellus are obligatory associates of bumblebees (Bombus) in the Holarctic region. Several occasional records of different species of the genus Parasitus and Vulgarogamasus are known from bumblebee nests and honeybee hives in North America and Europe (see below). Pergamasus crassipes (Linnaeus, 1758), Pergamasus diversus Halbert, 1915 have been recorded from the honeybee, Apis mellifera L. in Europe (Chmielewski W. 1991; De Jong et al., 1982; Haragsim et al., 1987), but these records probably are also accidental. We give accounts for the genera Parasitellus, Parasitus, and Vulgarogamasus, but not for Pergamasus, which is not found so far on North American bees. This genus, however, is included in the key below.

Key to genera of the family Parasitidae found on bees or bee nests
deutonymphs and adults

1 Females with entire holodorsal shield. Setae of dorsal hexagon homogeneous (subfamily Pergamasinae). Idiosoma elongate, not rounded. Holodorsal shield of female not fused with ventral shield ... Pergamasus
- Females with 2 dorsal shields, occasionally shields partially fused. Setae z5 may differ from other setae of dorsal hexagon. (subfamily Parasitinae) ... 2
2(1) Opistogaster with more than 40 pairs of setae ... Parasitellus
- Opistogaster with less than 30 pairs of setae ... 3
3(2) Setae of dorsal hexagon almost similar in form. Base of male tritosternum never closely associated with genital orifice. (Deutonymphs of a single North American species associated with bees has biramous tectum)... Vulgarogamasus
- Setae of dorsal hexagon different in form, usually setae z5 stout and pilose and j5 and j6 simple. Base of male tritosternum closely associated with genital orifice. Tectum usually triramous. ... Parasitus

Genus Parasitellus Willmann, 1939

Parasitellus Willmann, 1939: 534 (Type species Eugamasus (?) ferox Tragardh, 1910 = Acarus fucorum De Geer, 1778 by original designation); Hyatt, 1980: 327 (his synonymy).
Parasitus (Parasitus) non Latreille, 1795: Tichomirov, 1977: 60.
Parasitus non Latreille, 1795: Richards, 1976: 732 (part.); Karg, 1985: 525 (part.).

This genus includes 11 species that inhabit nests of bumblebees (Bombus) in the Holarctic region (Davydova, 1988; Hyatt, 1980; Karg, 1985; Richards, 1976). Occasionally they occur in beehives or borrows of small mammals (Hyatt, 1980; Crozier, 1989). Deutonymphs are commonly phoretic on the adult bumblebees or cuckoo bumblebees. Since bumblebee colonies are annual and only young queens overwinter, mite deutonymphs are able to distinguish between queens and other castes (workers, males). Although the mites may disperse on all castes of bumblebees, they prefer queens, and never move from a queen to a male. Mites dispersing on workers and males may try to switch to queens later, either during copulation or on flowers, where bumblebees forage (Huck et al., 1998; Schwarz and Huck, 1997).
The species of Parasitellus, although only associated with bumblebees, are not specific to a particular species of host, with species often co-occurring in individual Bombus nests. The lack of host specificity in this group may be the result of host switching in flowers by cockoo bumblebees, or by queens that visit old nests with overwintered deutonymphs. In the nests of a single bumblebee species, mites can disperse due to queen supersedure or invasion of workers or queens from different nests (Richards and Richards, 1976; Tichomirov, 1969a).
The exact nature of the association between these mites and their bumblebee hosts is uncertain, although predatory behavior toward acarid mites and other parasitid mites was suggested (Richards and Richards, 1976). If these mites feed preferentially on potentially damaging acarid mites, they may be beneficial to colony health.
Variation in many species of Parasitellus involves asymmetry in the shape of the tectum.

Key to species of the genus Parasitellus from North America*
deutonymphs
1. Tarsus I not divided ... Parasitellus inquilinobombus
- Tarsus I divided ... 2
2(1). Opisthonotal shield with more than 20 pairs of setae. Tectum with tapered middle point. Apex of tarsus II with 2 lateral spines. Basitarsus IV with 1 long seta (about twice as long as other tarsal setae) ... Parasitellus talparum
- Opisthonotal shield with less than 16 pairs of setae. Tectum with rounded or slightly tapered middle point. Apex of tarsus II with filiform setae. Basitarsus IV with or without long seta ... 3
3(2) Basitarsus IV without long setae. Metapodal shields comma-shaped. Sternal setae smooth. ... Parasitellus hobbsi
- Basitarsus IV with 1 long seta (about twice as long as other tarsal setae). Metapodal shields oval or pear-shaped. Sternal setae serrate or smooth. ... 4
4(3) Opisthonotal shield with rounded posterior end and slightly convex lateral edges. Sternal shield narrower (Fig. 1), without longitudinal striations. Sternal setae serrate, sternal setae ST1-3 not inflated at bases ... Parasitellus perthecatus
- Opisthonotal shield triangular. Sternal shield broad, with scarce longitudinal striations. Sternal setae smooth, sternal setae ST1-ST3 distinctly inflated at bases ... Parasitellus sp ex Bombus diligens (Mexico)

* Crozier (1989) lists four tentatively identified species from in bee hives in Nova Scotia (Canada): Parsitus near crinitus, Parasitus near favus, Parasitus near perthecathus, and Parasitus sp5. Davis and McRoy (1987) reported "Parasitus sp. that is not one of the native species" for Canada. This material is not included in the key because it has not been yet studied.

Parasitellus talparum (Oudemans, 1913)

Parasitus talparum Oudemans, 1913: 333.
Parasitellus talparum: Hyatt, 1980: 337, Figs 48-49 (his synonymy); Klimov, 1998: 18.
Parasitus favus Richards, 1976: 735, Figs 8-14; Richards and Richards, 1976: 1, syn. n.

Material examined. 1 deutonymph - USA: Alaska, "upper Amsler river", ex Bombus, 11 May 1995, K. Ross, BMOC 04-0326-001; 6 deutonymphs - USA: Wyoming, Albany Co., Laramie, 1 Jun 1999, ex Bombus (Pyrobombus) flavifrons Cresson, 1863, S. Shaw & R. Nunamaker, BMOC 99-0624-001; 2 males, 2 deutonymphs - USA: Ohio, Franklin Co., Columbus, nest of Bombus (Fervidobombus) fervidus (Fabricius, 1798), 6 Jul 2002, J. Wenzel, BMOC 02-0706-001.
Biology. This species was originally described from a nest of the common vole, Talpa europea L., 1758 in the Netherlands. Numerous subsequent records indicate that this species is actually associated with different species of Bombus (Hyatt, 1980). The biology of Parsitellus talparum (=Parasitus favus) was studied by Richards and Richards (1976). In a bee nest, several generations of mites can be accomplished in a single season. Males copulate with females several times. Female lays 0-1 eggs in laboratory settings. Larvae hatch in 5 days and molt into protonymphs in six days. Protonymphs constantly die in cultures. The deutonymphs captured from bumblebee nests were able to survive longer in culture than any other developmental stages. Usually the sex ratio is biased toward females. The feeding preferences are not known, although feeding on nidicolous microarthropods is possible. Deutonymphs and females were observed to be cannibalistic. Deutonymphs usually attach themselves to hairs of metasoma and propodeum using their chelicerae. If their numbers are great, they will also crowd around the neck. Usually the mites occur on queens before the bee starts to rearrange nesting material in a prospective nest, and more rarely on workers or males. The number of deutonymphs increases with queen production. The highest number of Parasitus deutonymphs on a single bee queen can be up to 210, but usually they are less numerous. The mites are easily excited, and if disturbed will run over the entire body, but seldom leave the bee.
Distribution. USA (first record): Alaska, Wyoming, Ohio; Canada: Alberta, Northwest Territories; Greenland; British Isles; Sweden; Germany; Netherlands (type locality); Switzerland; former Czechoslovakia; Russia: Moskovskaya Oblast', Western and Eastern Siberia, Kuril Islands; Turkmenistan; China: Qinghai Prov. (Hyatt, 1980; Gu et al., 1987; Klimov, 1998; Micherdzinski, 1969; Richards, 1976; Richards and Richards, 1976; Tichomirov, 1977).
Hosts. North America:
Bombus (Alpinobombus) polaris Curtis, 1835
Bombus (Bombias) nevadensis Cresson, 1874
Bombus (Bombus) terricola Kirby, 1837 (as occidentalis)
Bombus (Cullumanobombus) rufocinctus Cresson, 1863
Bombus (Fervidobombus) fervidus (Fabricius, 1798) (as californicus)
Bombus (Psithyrus) fernaldae (Franklin, 1911)
Bombus (Psithyrus) insularis (Smith, 1861)
Bombus (Psithyrus) suckleyi Green, 1860
Bombus (Pyrobombus) bifarius Cresson, 1878
Bombus (Pyrobombus) flavifrons Cresson, 1863
Bombus (Pyrobombus) frigidus Smith, 1854
Bombus (Pyrobombus) huntii Greene, 1860
Bombus (Pyrobombus) lapponicus (Fabricius, 1793) (as sylvicola)
Bombus (Pyrobombus) melanopygus Nylander, 1848
Bombus (Pyrobombus) mixtus Cresson, 1878
Bombus (Pyrobombus) ternarius Say, 1837
Bombus (Pyrobombus) vagans Smith, 1854
Bombus (Subterraneobombus) appositus Cresson, 1878
(Richards, 1976; Rrichards and Richards, 1976)

Eurasia (only bumblebee hosts are listed):
Bombus (Bombus) albocinctus Smith
Bombus (Bombus) florilegus Panfilov, 1956
Bombus (Bombus) terrestris (Linnaeus, 1758)
Bombus (Megabombus) tichenkoi (Skorikov, 1925)
Bombus (Melanobombus) lapidarius (Linnaeus, 1758)
Bombus (Thoracobombus) pascuorum (Scopoli, 1763) (as agrorum (F.))
(Hyatt, 1980; Klimov, 1998; Micherdzinski, 1969)

Parasitellus perthecatus (Richards, 1976), comb. n.

Parasitus perthecatus Richards, 1976: 738, Figs 22-31; Richards and Richards, 1976: 1.

Material examined. 2 deutonymphs - USA: Alaska, "upper Amsler river", ex Bombus, 11 May 1995, K. Ross, BMOC 04-0326-001; 1 deutonymph - USA: Ohio, Franklin Co., Columbus, nest of Bombus (Fervidobombus) fervidus (Fabricius, 1798), 6 Jul 2002, J. Wenzel, BMOC 02-0706-001; 2 deutonymphs - USA: Colorado, Boulder Co., Roosevelt National Forest, Indian Peaks Wilderness, Niwot Ridge, trough, 3667 m., malaise trap, 8 Aug 1993, J. Cooley, BMOC 94-0125-005.
Biology. Female lays 0-8 eggs per day in laboratory settings. Larvae hatch in 3-6 days (average 3.9) and in 3-6 days molt to protonymphs. The highest number of the deutonymphs on a single bee queen can be up to 210. Otherwise the biology is similar to that of P. talparum (Richards and Richards, 1976).
Distribution. USA (first record): Alaska, Ohio, Colorado; Canada: Northwest Territories, Alberta (type locality).
Hosts.
Bombus (Alpinobombus) polaris Curtis, 1835
Bombus (Bombias) nevadensis Cresson, 1874
Bombus (Bombus) terricola Kirby, 1837 (as occidentalis)
Bombus (Cullumanobombus) rufocinctus Cresson, 1863
Bombus (Fervidobombus) fervidus (Fabricius, 1798) (as californicus)
Bombus (Psithyrus) fernaldae (Franklin, 1911)
Bombus (Psithyrus) insularis (Smith, 1861)
Bombus (Psithyrus) suckleyi Green, 1860
Bombus (Pyrobombus) bifarius Cresson, 1878
Bombus (Pyrobombus) centralis Cresson,1864
Bombus (Pyrobombus) flavifrons Cresson, 1863
Bombus (Pyrobombus) frigidus Smith, 1854
Bombus (Pyrobombus) huntii Greene, 1860
Bombus (Pyrobombus) lapponicus (Fabricius, 1793) (as sylvicola)
Bombus (Pyrobombus) melanopygus Nylander, 1848
Bombus (Pyrobombus) mixtus Cresson, 1878
Bombus (Pyrobombus) ternarius Say, 1837
Bombus (Pyrobombus) vagans Smith, 1854
Bombus (Subterraneobombus) appositus Cresson, 1878 (type host)
(Richards, 1976; Rrichards and Richards, 1976)

Parasitellus sp.

Material examined. 1 deutonymph - MEXICO: Michoacán de Ocampo, 12 mi. W. Zamora de Hidalgo [label reads Zamora], on Hwy 5, 5000 ft, on sides of propodeum of Bombus (Fervidobombus) diligens Smith, 1861 (Apidae), 4 Aug 1962, S. Graham, UMMZ BMOC 83-0428-002.
Biology. Unknown.
Distribution. Mexico: Michoacán de Ocampo.
Hosts. Bombus (Fervidobombus) diligens Smith, 1861.

Parasitellus hobbsi (Richards, 1976), comb. n.

Parasitus hobbsi Richards, 1976: 733, Figs 1-7; Richards and Richards, 1976: 3; Crozier, 1989: 167.

Biology. Similar to that of other Parasitellus. Occurs less frequently than P. talparum and P. perthecatus. In Canada (Nova Scotia) was found in beehives.
Distribution. Canada: Alberta (type locality); Nova Scotia (Crozier, 1989; Richards, 1976).
Hosts.
Bombus (Bombias) nevadensis Cresson, 1874
Bombus (Bombus) terricola Kirby, 1837 (as occidentalis)
Bombus (Cullumanobombus) rufocinctus Cresson, 1863
Bombus (Fervidobombus) fervidus (Fabricius, 1798) (as californicus)
Bombus (Psithyrus) fernaldae (Franklin, 1911)
Bombus (Psithyrus) insularis (Smith, 1861)
Bombus (Pyrobombus) bifarius Cresson, 1878 (type host)
Bombus (Pyrobombus) flavifrons Cresson, 1863
Bombus (Pyrobombus) frigidus Smith, 1854
Bombus (Pyrobombus) huntii Greene, 1860
Bombus (Pyrobombus) mixtus Cresson, 1878
Bombus (Pyrobombus) ternarius Say, 1837
Bombus (Pyrobombus) vagans Smith, 1854
Bombus (Subterraneobombus) appositus Cresson, 1878
(Richards, 1976; Rrichards and Richards, 1976)

Parasitellus inquilinobombus (Richards, 1976), comb. n.

Parasitus inquilinobombus Richards, 1976: 736, Figs 14-21; Richards and Richards, 1976: 9.

Material examined. 1 deutonymph - USA: Michigan, Mackinac Co., Millecoquins River, at US 2, T43N,R9W,530, killing jar debris (with Bombus vagans, Bombus ternarius, Bombus terricola), T. W. Porter, 30 Aug 1965 [not processed].
Biology. Similar to that of other Parasitellus. Occurs less frequently than P. talparum and P. perthecatus.
Distribution. USA (first record): Michigan; Canada: Alberta (type locality).
Hosts.
Bombus (Bombus) terricola Kirby, 1837 (as occidentalis)
Bombus (Cullumanobombus) rufocinctus Cresson, 1863
Bombus (Fervidobombus) fervidus (Fabricius, 1798) (as californicus)
Bombus (Psithyrus) fernaldae (Franklin, 1911)
Bombus (Psithyrus) insularis (Smith, 1861)
Bombus (Pyrobombus) bifarius Cresson, 1878 (type host)
Bombus (Pyrobombus) flavifrons Cresson, 1863
Bombus (Pyrobombus) frigidus Smith, 1854
Bombus (Pyrobombus) huntii Greene, 1860
Bombus (Pyrobombus) mixtus Cresson, 1878
Bombus (Pyrobombus) ternarius Say, 1837
Bombus (Pyrobombus) vagans Smith, 1854
Bombus (Subterraneobombus) appositus Cresson, 1878
(Richards, 1976; Rrichards and Richards, 1976)

Genus Parasitus Latreille, 1795

Parasitus Latreille, 1795: 19 (Type species Acarus coleoptratorum Linnaeus, 1758); Richards, 1976: 732 (part.); Hyatt, 1980: 256 (his synonymy).
Parasitus (Coleogamasus) Tichomirov, 1969b: 1470 (type species Gamasus celer C. L. Koch, 1835 by original designation); Tichomirov, 1977: 61.

Species of this genus live in manure, dung, compost or other decaying organic substances. Many of them have broad geographic ranges. Usually, deutonymphs are phoretic on dung beetles (Hyatt, 1980).
Several occasional records are available from bumblebee nests: Parasitus fimetorum (Berlese, 1904) in North America and Europe (Hyatt, 1980; Richards, 1976); Parasitus beta Oudemans et Voigts, 1904, Parasitus coleoptratorum (Linnaeus, 1758), Parasitus cornutosimilis (Schweizer, 1949), and Parasitus loricatus (Wankel, 1861) in Europe (Hyatt, 1980; Karg, 1971; Micherdzinski, 1969). Parasitus coleoptratorum was also found in honeybee hives in Europe (De Jong et al., 1982), and we have seen a single female of Parasitus aff. loricatus from a mammal nest in a bee hive (New York, Tompkins Co., Ithaca, 14 Dec 1976, V. Lazar, BMOC 76-1214-001).
We give an account on Parasitus fimetorum, because it frequently occurs in the nests of bumblebees in the United States and Canada (Richards, 1976; Richards and Richards, 1976). Athias-Henriot (1979) created a new genus for this species, Phorytocarpais Athias-Henriot, 1979, however, we accept here generic designation of Hyatt (1980).

Parasitus fimetorum (Berlese, 1904)

Gamasus fimetorum Berlese, 1904: 238.
Parasitus fimetorum: Hyatt, 1980: 271, Figs 12-14 (his synonymy); Richards, 1976: 741; Richards and Richards, 1976: 1.
Parasitus (Coleogamasus) fimetorum: Tichomirov, 1977: 86, Figs 36,9; 37,9.
Parasitus fimestorum [lapsus pro fimetorum]: Gu et al., 1987: 41.
Phorytocarpais fimetorum: Hennesey and Farrier, 1988: 10, Figs 4-6.
Gamasus posticatus Banks, 1910: 137, Fig. 10, 3,8 (synonymized by Hennesey and Farrier, 1988).

Material examined. 4 deutonymphs (2 slides) from USA (Michigan) [not processed].
Biology. Parasitus fimetorum is a common inhabitant of compost, rotting vegetation, manure, dung, and subterranean nests of small mammals, and also occasionally bumblebee and bird nests. Deutonymphs are phoretic on coprophagous and necrophagous beetles, Aphodius, Geotrupes (Coleoptera: Scarabaidae), Nicrophorus (Silphidae), and Atholus (Histeridae). In the United States, Canada, and Europe, this species was found in bumbleebee nests, but not on the bees. Thus, it likely does not have any specific relationships to bumblebees (Hyatt, 1980; Richards, 1976).
Distribution. USA; Michigan; Canada: Alberta; Iceland, British Isles, Netherlands, Belgium, Germany, Austria, Switzerland, Italy (type locality); Poland; former USSR, including Western Siberia; China: Yunnan Prov.; Australia (Gu et al., 1987; Richards 1976; Wallace and Holm, 1985).
Hosts (only North American records).
Bombus (Bombus) terricola Kirby, 1837 (as occidentalis)
Bombus (Cullumanobombus) rufocinctus Cresson, 1863
Bombus (Fervidobombus) fervidus (Fabricius, 1798) (as californicus)
Bombus (Fervidobombus) pensylvanicus (De Geer, 1773) (as americanorum) (first record)
Bombus (Pyrobombus) bifarius Cresson, 1878
Bombus (Pyrobombus) flavifrons Cresson, 1863
Bombus (Pyrobombus) frigidus Smith, 1854
Bombus (Pyrobombus) huntii Greene, 1860
Bombus (Pyrobombus) mixtus Cresson, 1878
Bombus (Pyrobombus) ternarius Say, 1837
Bombus (Pyrobombus) vagans Smith, 1854
Bombus (Subterraneobombus) appositus Cresson, 1878
(Richards and Richards, 1976)

Genus Vulgarogamasus Tichomirov, 1969

Parasitus (Vulgarogamasus) Tichomirov, 1969a: 1335 (Type species Parasitus (Vulgarogamasus) burchanensis Oudemans, 1903 = Parasitus burchanensis Oudemans, 1903 by original designation); Tichomirov, 1977: 61.
Vulgarogamasus: Hyatt, 1980: 290; El-Banhawy and Nasr, 1986.

This genus includes 39 described species distributed in the New and Old World (Bai et al., 1995; Bai et al., 1991; El-Banhawy and Nasr, 1986; Hyatt, 1980; Gu and Guo, 1997; Gu and Huang, 1993; Gu et al., 1987; Karg, 1998; Ma, 1987; Ma, 1990; Ma et al., 2002; Ma-Liming and Wang-Shenron, 1996; Tichomirov, 1969a; Tseng, 1995; Ye and Ma, 1996; Ye et al., 1996). Many species of the genus are habitat generalists living in dung, decaying seaweed, forest litter, poultry litter, moss, and nests of small mammals and birds. One species, Vulgarogamasus sphecophilus (Cooreman, 1945), is restricted to the nests of wasps. Vulgarogamasus pollinerus El-Banhawy et Nasr, 1984 has been previously known only from a bee hive in Ithaca, New York. We found this species on bumblebees and in their nests.

Vulgarogamasus pollinerus El-Banhawy et Nasr, 1984

Vulgarogamasus pollinerus El-Banhawy et Nasr, 1984: 76, Figs 1-10.

Material examined. 7 slides from R. W. Husband's collection (not processed).
Biology. A few deutonymphs have been originally found in a bee hive. We found seven deutonymphs phoretic on bumblebees and inside their nests. There is no evidence, however, that V. pollinerus is strictly associated with bees. Probably, this species, like other Parasitidae, preys on small arthropods in a variety of habitats, including bee nests.
Distribution. USA; Indiana (first record); Michigan (first record); Montana (first record), New York (type locality).
Hosts. Apis mellifera Linnaeus, 1758 (type host)
Bombus (Bombias) auricomus (Robertson, 1903) (first record)
Bombus (Bombus) terricola Kirby, 1837 (first record)
Bombus (Fervidobombus) pensylvanicus (De Geer, 1773) (as americanorum) (first record)
Bombus (Pyrobombus) bimaculatus Cresson, 1863 (first record)
Bombus (Pyrobombus) impatiens Cresson, 1863 (first record)

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