Fig. 1. Mite Pneumolaelaps costai (Laelapidae) ex Bombus fervidus from Ohio Click here to enlarge

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Genus Pneumolaelaps Berlese, 1920

Hypoaspis (Pneumolaelaps) Berlese, 1920: 151 (part) (type species Iphis bombicolens G. R. Canestrini by original designation); Evans and Till, 1966: 158; Karg, 1982: 243 (part); Karg, 1984: 35; Bregetova, 1977: 514.
Pneumolaelaps: Hunter, 1966: 357 (part.); Hunter and Husband, 1973: 77 (part.).
Qinghailaelaps Gu and Yang, 1984: 371 (type species Qinghailaelaps marmotae Gu et Yang, 1984 by original designation], syn. n.

Species of the genus Pneumolaelaps are common inhabitants of nests of bumblebees. They may also occasionally occur in beehives and in subterranean nests of small mammals (Bregetova, 1977; Haitlinger, 1983; Crozier, 1989). The mites may be found in great numbers in active bumblebee colonies, moving rapidly over the nest floor, nectar pots, and brood cells where they feed on nectar and pollen. Female mites become phoretic on bumblebee queens in old nests, and overwinter with the queens in sheltered sites. Feeding behavior is only known for Pneumolaelaps longanalis (see below). This genus is known from the Holarctic, Oriental and Neotropical regions (Eguaras et al. 1997; Evans and Till, 1966; Hunter and Husband, 1973; Karg, 1982, 1984; Rosario, 1981; Royce and Krantz, 1989).
There are, at present, no comprehensive definitions of this genus. Many authors consider it as a subgenus of the genus Hypoaspis (Bregetova, 1977; Evans and Till; 1966; Karg, 1982) while others consider it as a distinct genus (Casanueva, 1993). According to Evans and Till (1966), this taxon has the following apomorphic character states: the stimata are large and peritremes broad, the opistogaster is usually hypetrichous, and there are two ventral setae on genu IV. Karg (1982) defines it based on the following characters: the idiosomal setae filiform, without extreme difference in lengths, unpaired setae frequently present on the posterior part of the dorsal shield; tectum smooth; rows of the hypognathal groove usually with 2 to 6 denticles; the epigynial shield with one pair of setae; 1-2 pairs of setae between the epigynial and anal shields. The latter concept allows the inclusion of free-living species, making the genus, in our opinion, artificial. We accept the point of view of the former authors restricting the genus only to bumblebee-associated species. With the absence of a robust phylogeny of the genus Hypoaspis and proofs of the Pneumolaelaps monophyly, this is highly speculative, however.
Qinghailaelaps marmotae, the type species of the genus Qinghailaelaps Gu and Yang, 1984 has been described from Marmota himalayana (Hodgson, 1841) (Rodentia: Sciuridae); another species of this genus, Qinghailaelaps gui Bai, 1992, was found phoretic on a bumblebee (Bombus). Because the genus-level characters of Qinghailaelaps correspond to those of Pneumolaelaps, we consider the former as a junior synonym. The following new combinations are proposed: Pneumolaelaps cavicolous (Gu, Liu et Niu, 1997) from Qinghailaelaps; Pneumolaelaps gui (Bai, 1992), comb. n. from Qinghailaelaps; Pneumolaelaps marmotae (Gu et Yang, 1984), comb. n. from Qinghailaelaps; Pneumolaelaps qinghaiensis (Li, Yang et Wang, 1998), comb. n. from Qinghailaelaps.

Key to species of the genus Pneumolaelaps from North America*
females

1	Posterior margin of sternal plate concave ... 2
-	Posterior margin of sternal plate straight or convex ... 6
2(1)	Dorsal plate seta j2 extending approximately to base of seta j4 Pneumolaelaps groenlandicus
-	Seta j2 extending no more than 1/2 the distance between the bases of seta j3 and j4 ... 3
3(2)	Seta j2 extending approximately 1/2 the distance to the base of seta j3 Pneumolaelaps longanalis
-	Seta j2 extending to or slightly beyond the base of seta J3 ... 4
4(3)	Dorsal seta j4 approximately 3 times longer than j5; distance between genital setae bases less than distance between sternal setae st2 Pneumolaelaps sinhai
-	Dorsal seta j4 equal to or only slightly longer than j5; distance between bases of genital setae equal to or slightly exceeding the distance between sternal setae st2 ... 5
5(4)	Dorsal seta j3 extending 1/2 to 2/3 of the distance to seta j4; genital plate without striation pattern Pneumolaelaps aequalipilus
-	Dorsal seta j3 extending almost to base of j4; genital plate with distinct striation pattern Pneumolaelaps patae
6(2)	Dorsal setae j2, j3, and j4 approximately equal in length; seta j4 3-4 times length of seta j5; dorsal seta Z4 and Z5 at least 4 times as long as J5; genital setae extending to posterior margin of genital plate Pneumolaelaps longipilus
-	Dorsal seta j4 less than 3 times length of seta j5; length of seta j2, j3, and J4 variable; setae Z4 and Z5 equal to or only slightly longer than J5; genital setae not extending to posterior margin of genital plate ... 7
7(6)	Dorsal seta j3 extending no more than 1/2 the distance to base of seta j4 ... 8
-	Dorsal seta j3 and j4 extending over 1/2 the distance to base of seta j4 ... 9
8(7)	Genital plate bearing distinct striation pattern; dorsal plate with distinct striation pattern in area of setae J1 to J3 Pneumolaelaps connieae
-	Genital plate without striation pattern; dorsal plate without distinct striation pattern in area of setae J1 to J3 Pneumolaelaps richardsi
9(7)	Anterior marginal dorsal setae (s and r rows) 2 or more times length of seta j6; genital seta extending 2/3 the distance between setal base and posterior margin of genital plate Pneumolaelaps mistipilus
-	Anterior marginal dorsal setae (s and r rows) equal to or only slightly longer than seta j6; genital seta extending no more than 1/2 the distance to posterior margin of plate Pneumolaelaps costai

* modified from Hunter and Husband (1973) Pneumolaelaps groenlandicus (Trägårdth, 1904) Hypoaspis bombicolens (Can.) var. groenlandica Trägårdth, 1904: 34 (part); Hunter, 1966: 359 (part). Pneumolaelaps groenlandica: Hunter and Husband, 1979: Fig. 1A-E (lectotype designation). Hypoaspis (Pneumolaelaps) arctos Karg, 1984: 37, Figs 2, 3a-c, syn. n. Non Hypoaspis bombicolens (Can.) var. groenlandica Trägårdth, 1904: 34, Figs 59-60. Non Hypoaspis (Pneumolaelaps) ?groenlandicus Bregetova, 1977: 519, Fig. 411. Non Hypoaspis (Pneumolaelaps) groenlandica Karg, 1982: 244; Karg, 1984: 36, Fig. 1b. Material. 2 paralectotype females - Greenland: Vestgrønland, Tigssaluk [label reads Tigsaluk, Tlumle], ex Bombus sp. No. 31, 8 Aug 1889, W. Lundbeck. Distribution. Greenland (type locality); also recorded for Russia: Jinretlen (East Chukotka) (Hunter and Husband, 1979; Karg, 1984; Trägårdth, 1904). Biology. Unknown. Hosts. Bombus sp. (type host), Bombus (Alpinobombus) hyperboreus Schönherr, 1809 (Hunter and Husband, 1979; Karg, 1984; Trägårdth, 1904). Notes. Paralectotypes of Hypoaspis bombicolens var. groenlandica are represented by at least three species of Pneumolaelaps: 1) Pneumolaelaps sp. (single female in bad shape; neither resembles the original drawings nor subsequent interpretation of this species by Bregetova (1977) and Karg (1984)); 2) Pneumolaelaps groenlandica sensu Hunter et Husband, 1979 (we studied two paralectotype females, they do not correspond Trägårdth's original drawings); 3) Pneumolaelaps patae Hunter et Husband, 1979 (1 paratype female from Trägårdth's syntypes; probably, this is species that has been depicted by Trägårdth). Because Hunter et Husband (1979) designated a female of the second species as the lectotype for the Trägårdth's variety, it becomes the unique bearer of the name of Hypoaspis bombicolens var. groenlandica Trägårdth, 1904 and the standard for its application (ICZN, 1999, article 74.1). Pneumolaelaps longanalis Hunter et Husband, 1973 Pneumolaelaps longanalis Hunter and Husband, 1973: 81, Fig. 1F-N; Royce and Krantz, 1989: 161; Crozier, 1989: 167. Material. Holotype female - USA: Michigan, Kalamazoo Co., Gull Lake Biological Station, ex petiole of female of Bombus griseocollis, 10 Aug 1964, R. W. Husband (kept in University of Georgia, Athens). 2 females - USA: Ohio Franklin Co., Columbus, Bombus fervidus nest, 6 Jul 2002, J. Wenzel, UMMZ BMOC 02-0706-001; 1 female - USA: New York, Tompkins Co., Ithaca, ex Bombus impatiens worker, 21 May 1974, B.M. OConnor, UMMZ BMOC 74-0521-001. Biology. The following account on the feeding behavior of Pneumolaelaps longanalis is modified from Royce and Krantz (1989). Wandering P. longanalis congregates in considerable numbers on the brood cells, and larvae feeding on pollen grains provided by the adult bumblebees. When an individual pollen grain was chosen, the gnathosoma was depressed ventrally relative to the opisthosoma, and the grain removed from its sticky substrate by the palps and chelicerae. The nectar-coated grain was held in the region of the hypostome under the encircling palpi and rapidly rotated with the help of the chelicerae. During the manipulation, the nectar coating applied earlier by the foraging bee was stripped from the pollen grain surface. The mite probably also removed much of the surface pollenkitt, a lipoidal substance that covers the pollen grain surface and is incorporated to some extent in the outer wall (exine). Pollenkitt, however, does not trigger feeding in the absence of nectar. It appears that the removed nectar and pollenkitt are dissolved by salivary secretions that are directed over the rotating pollen-grain surface, and then recaptured and redirected to the preoral cavity via the fluid transport system formed by the capitular groove and overlying tritosternum. Following feeding, the stripped pollen grain was discarded and another quickly chosen for manipulation. Females were found to process pollen grains twice as rapidly as either males or deutonymphs. The mites may rupture thin-walled pollen grains during the feeding process, and it is possible that nutrients are acquired from the pollen core in this way. Distribution. USA: Kansas, Michigan (type locality), Ohio (first record), Pacific Northwest of the United States; Canada: Alberta (Hunter et Husband, 1973; Royce and Krantz, 1989; our data). Hosts. Apis mellifera Linnaeus, 1758 Bombus (Bombias) nevadensis Cresson, 1874 Bombus (Bombus) affinis Cresson, 1863 Bombus (Bombus) terricola Kirby, 1837 (including occidentalis) Bombus (Fervidobombus) fervidus (Fabricius, 1798) (including californicus) Bombus (Fervidobombus) pensylvanicus (DeGeer, 1773) (as americanorum) Bombus (Fraternobombus) fraternus (Smith, 1854) Bombus (Psithyrus) suckleyi Green, 1860 Bombus (Pyrobombus) bifarius Cresson, 1878 Bombus (Pyrobombus) bimaculatus Cresson, 1863 Bombus (Pyrobombus) frigidus Smith, 1854 Bombus (Pyrobombus) impatiens Cresson, 1863 (first record) Bombus (Pyrobombus) mixtus Cresson, 1878 Bombus (Pyrobombus) ternarius Say, 1837 Bombus (Pyrobombus) vagans Smith, 1854 Bombus (Separatobombus) griseocollis (DeGeer, 1773) (type host) Pneumolaelaps sinhai Hunter et Husband, 1973 Pneumolaelaps sinhai Hunter and Husband, 1973: 82, Fig. 2A-E; Crozier, 1989: 167. Material. 10 paratype females (unprocessed). Biology. Unknown. Distribution. USA: Michigan; Canada: Alberta (type locality); Nova Scotia. Hosts. Apis mellifera Linnaeus, 1758 Bombus (Fervidobombus) fervidus (Fabricius, 1798) (as californicus) Bombus (Psithyrus) insularis (Smith, 1861) Bombus (Psithyrus) suckleyi Green, 1860 (type host) Bombus (Pyrobombus) flavifrons Cresson, 1863 Bombus (Pyrobombus) frigidus Smith, 1854 Pneumolaelaps aequalipilus Hunter, 1966 Pneumolaelaps aequalipilus Hunter, 1966: 360, Fig. 1; Hunter and Husband, 1973: 78, Fig. 4I-J. Material. Multiply slides (unprocessed) Biology. Unknown. Distribution. USA: Georgia (type locality); Florida (first record); Illinois (first record); Lousiana (first record); South Carolina (first record); Taxes (first record); Kansas (first record). Hosts. Bombus (Pyrobombus) impatiens Cresson, 1863. Bombus (Fervidobombus) pensylvanicus (DeGeer, 1773) (type host) Bombus (Psithyrus) variabilis (Cresson, 1872) Pneumolaelaps patae Hunter et Husband, 1973 Hypoaspis bombicolens (Can.) var. groenlandica Trägårdth, 1904: 34 (part), Figs 59-60; Hunter, 1966: 359 (part). Pneumolaelaps patae Hunter et Husband, 1973: 78, Fig. 2F-J. Hypoaspis (Pneumolaelaps) ?groenlandicus Bregetova, 1977: 519 (part). Hypoaspis (Pneumolaelaps) groenlandica Karg, 1982: 244 (part); Karg, 1984: 36 (part). Material. 9 paratype females (unprocessed). Biology. Unknown. Distribution. Canada: Nunavut; Greenland: Vestgrønland (type locality). Hosts. Bombus sp. (type host); Bombus (Alpinobombus) hyperboreus Schönherr, 1809; Bombus (Alpinobombus) polaris Curtis, 1835. Notes. Bregetova's (1977) identified mites of the genus Pneumolaelaps from Yakutiya and Chukotka without a striated sternal shield as Hypoaspis bombicolens var. groenlandica, and this interpretation was subsequently accepted by Karg (1982, 1984). Because all of the Trägårdth's syntypes [paralectotypes] we have seen have a distinctly striated sternal shield, we believe that Hypoaspis (Pneumolaelaps) groenlandicus sensu Bregetova, 1977 is, probably, a new species. Pneumolaelaps connieae Hunter et Husband, 1973 Pneumolaelaps connieae Hunter and Husband, 1973: 84, Fig. 3A-E. Material. 1 holotype and 5 paratype females (unprocessed). Biology. Unknown. Distribution. USA: Michigan (type locality); Georgia. Hosts. Bombus (Bombus) terricola Kirby, 1837 (including occidentalis) Bombus (Fervidobombus) pensylvanicus (DeGeer, 1773) (as americanorum) (type host) Bombus (Pyrobombus) impatiens Cresson, 1863. Bombus (Separatobombus) griseocollis (DeGeer, 1773) Pneumolaelaps richardsi Hunter et Husband, 1973 Pneumolaelaps richardsi Hunter and Husband, 1973: 86, Figs 3F-M. Material. 1 holotype, 8 paratype females and 2 paratype males. Biology. Unknown. Distribution. Canada: Alberta (type locality); USA: Michigan. Hosts. Bombus (Psithyrus) suckleyi Green, 1860 Bombus (Pyrobombus) bifarius Cresson, 1878 (type host) Bombus (Pyrobombus) frigidus Smith, 1854 Bombus (Pyrobombus) huntii Greene, 1860 Bombus (Pyrobombus) mixtus Cresson, 1878 Bombus (Pyrobombus) ternarius Say, 1837 Pneumolaelaps costai Hunter et Husband, 1973 Pneumolaelaps costai Hunter and Husband, 1973: 87, Figs 4A-H. Material. 1 holotype, 5 paratype females, 3 paratype males, 1 paratype hymph; 2 females - USA: Ohio Franklin Co., Columbus, Bombus fervidus nest, 6 Jul 2002, J. Wenzel, UMMZ BMOC 02-0706-001. Biology. Unknown. Distribution. USA: Michigan (type locality); Ohio (first record). Hosts. Bombus (Bombus) terricola Kirby, 1837 (including occidentalis) Bombus (Fervidobombus) fervidus (Fabricius, 1798) (first record) Bombus (Fervidobombus) pensylvanicus (DeGeer, 1773) (as americanorum) (type host) Bombus (Pyrobombus) impatiens Cresson, 1863. Pneumolaelaps mistipilus Hunter, 1966 Pneumolaelaps mistipilus Hunter, 1966: 363, Fig. 2; Hunter and Husband, 1973: 79, Figs 4M-N. Material. 1 female from collection bottle with 5 species of Bombus (unprocessed). Biology. Unknown. Distribution. USA: Georgia (type locality); Michigan. Hosts. Bombus (Bombias) nevadensis Cresson, 1874 Bombus (Bombus) affinis Cresson, 1863 Bombus (Fervidobombus) fervidus (Fabricius, 1798) Bombus (Fervidobombus) pensylvanicus (DeGeer, 1773) (as americanorum) Bombus (Fervidobombus) weisi Friese, 1903 (as Psithyrus laboriosus) Bombus (Fraternobombus) fraternus (Smith, 1854) Bombus (Psithyrus) ashtoni (Cresson, 1864) Bombus (Pyrobombus) bimaculatus Cresson, 1863 Bombus (Pyrobombus) impatiens Cresson, 1863 (type host) Bombus (Pyrobombus) perplexus Cresson, 1863 Bombus (Separatobombus) griseocollis (DeGeer, 1773) Pneumolaelaps longipilus Hunter, 1966 Pneumolaelaps longipilus Hunter, 1966: 365, Fig. 3; Hunter and Husband, 1973: 78, Figs 4K-L. Material. 1paratype female, 6 females (unprocessed). Biology. Unknown. Distribution. USA: Georgia (type locality); Michigan (first record). Hosts. Bombus (Fervidobombus) pensylvanicus (DeGeer, 1773) (as americanorum) (type host); Bombus (Fervidobombus) fervidus (Fabricius, 1798) (as californicus); Bombus (Psithyrus) variabilis (Cresson, 1872). References     B. OConnor and P. Klimov © Created: Apr 19, 2004 Last modified: