From: ASSESSMENT OF SPECIES DIVERSITY IN THE MIXEDWOOD PLAINS ECOZONE
MITES
(Acari)

Ian M. Smith, Evert E. Lindquist
and Valerie Behan-Pelletier

MITE CLASSIFICATION AND HABITS

Mites comprise the taxonomic subclass Acari of the class Arachnida which is also represented in the Mixedwood Plains by the spiders (order Araneae), harvestmen (order Opiliones) and pseudoscorpions (order Pseudoscorpionida). Acari includes 2 orders, Parasitiformes represented in the Ecozone by the suborders Mesostigmata and Ixodides (Metastigmata) and Acariformes with 3 subordinal taxa, namely Prostigmata, Acaridiae and Oribatida, in the Mixedwood Plains. In Acariformes, recent studies indicate that Trombidiformes (including Prostigmata) and Sarcoptiformes (including Acaridiae and Oribatida) may reflect more natural major groupings (OConnor, 1984; Norton et al., 1993). However, the classification presented in Danks (1979) is followed here to facilitate comparisons with data and statements made about the Canadian fauna in that review. The acarine suborders Mesostigmata, Ixodides, Prostigmata and Oribatida are covered in the present document. Data on the Acaridiae will be added at a later date.

Parasitiformes - Mesostigmata

This suborder, also known as Gamasida, includes the supercohorts Monogynaspida and Trigynaspida. Monogynaspida contains 4 cohorts, namely, Sejina, Gamasina, Diarthrophallina and Uropodina; Trigynaspida comprises the 2 cohorts Cercomegistina and Antennophorina. The Canadian fauna of the various cohorts was treated at the family level by Lindquist (1979), Ainscough (1979) and Funk (1979). A summary of expected species diversity of families represented in the Mixedwood Plains is presented in Table 1.

TABLE 1: EXPECTED SPECIES DIVERSITY OF FAMILIES OF MESOSTIGMATA KNOWN FROM CANADA IN MIXEDWOOD PLAINS ECOZONE BASED ON RECORDS OF NAMED AND UNNAMED SPECIES IN CANADIAN NATIONAL COLLECTION OF ACARI

SUPERFAMILYFAMILYHABITAT
OR HOST OF
ADULT
FEEDING
BEHAVIOUR
OF ADULT
# SPECIES
EXPECTED
IN MWP/
CANADA
Monogynaspida - Sejina
SejoideaSejidaeLitterPredaceous1/2
"MicrogyniidaeLitterPredaceous1/2
"IchthyostomatogasteridaeLitterSaprophagous1/1
Monogynaspida -
Gamasina
EpicrioideaEpicriidaeLitterPredaceous1/7
"ArctacaridaeLitterPredaceous0/4
"ZerconiidaeLitterPredaceous15/50
ParasitoideaParasitidaeLitter, Nests,
Carrion
Predaceous20/85
"VeigaiidaeLitterPredaceous10/20
RhodacaroideaRhodacaridaeLitter, NestsPredaceous10/20
"OlogamasidaeLitter,
Compost
Predaceous10/30
"DigamasellidaeLitter, BarkPredaceous20/60
"HalolaelapidaeLitter,
Compost
Predaceous10/30
EviphidoideaEviphididaeLitter, DungPredaceous5/20
"MacrochelidaeLitter, Dung
Compost,
Carrion
Predaceous15/45
"ParholaspidaeLitter,
Compost,
Carrion
Predaceous5/20
"PachylaelapidaeLitterPredaceous3/15
AscoideaAscidaeLitter, Plants,
Bark, Nests
Predaceous,
Fungivorous
70/190
"PhytoseiidaePlants, LitterPredaceous40/100
"OtopheidomenidaeInsectsParasitic2/5
"AmeroseiidaeLitter, Nests,
Fungi
Fungivorous,
Phytophagous
10/25
"PodocinidaeLitterPredaceous1/2
DermanyssoideaLaelapidaeLitter, Nests,
Insects,
Mammals
Predaceous,
Parasitic
35/80
VarroideaHoney BeesParasitic1/1
"HaemogamasidaeMammalsParasitic5/15
"DermanyssidaeBirdsParasitic4/10
"MacronyssidaeSnakes, Birds,
Mammals
Parasitic10/20
"RhinonyssidaeBirdsParasitic15/25
"HalarachnidaeMammalsParasitic3/15
"SpinturnicidaeBatsParasitic2/4
"IxodorhynchidaeSnakesParasitic3/8
"EntonyssidaeSnakesParasitic1/3
"RaillietidaeMammalsParasitic1/1
Monogynaspida -
Diarthrophallina
DiarthrophalloideaDiarthrophallidaeBeetlesParasitic1/1
Monogynaspida -
Uropodina
UropodoideaProtodinychidaeLitterUnknown1/2
"PolyaspididaeLitter, Wood,
Compost
Saprophagous5/30
"DithinozerconidaeLitterPredaceous1/4
"UropodidaeLitter, Wood,
Nests,
Compost
Predaceous,
Saprophagous
50/140
"TrachyuropodidaeLitter, AntsSaprophagous2/6
Trigynaspida -
Cercomegistina
CercomegistoideaCercomegistidaeArthropodsPredaceous1/2
Trigynaspida -
Antennophorina
AntennophoroideaAntennophoridaeAntsSaprophagous1(?)/2
ParantennuloideaParantennulidaeMillipedes,
Beetles
Saprophagous1/2
"Philodanidae  0(?)/1
CelaenopsoideaCelaenopsidaeInsectsSaprophagous1/2
"EuzerconidaeBeetlesPredaceous,
Saprophagous
1/2
"DiplogyniidaeBeetlesPredaceous2/8
FedrizzioideaParamegistidaeMyriapods,
Insects,
Reptiles
Parasitic,
Saprophagous
1/1
TOTALS:
14 Superfamilies
45 Families  398/1118

Monogynaspida.

Sejina - Sejoidea. The only superfamily of Sejina has 3 families represented in the Mixedwood Plains Ecozone. These mites live in litter and decaying wood, often in association with wood-feeding beetles.
Gamasina. This large group includes 6 superfamilies and nearly 30 families represented in the Mixedwood Plains. Most are free-living predators, but some are parasites and some others are fungivores.
Epicrioidea. Species of the family Zerconidae occur commonly in litter and decaying wood in the Mixedwood Plains, and 1 species of the small family Epicriidae has been collected occasionally in similar habitats.
Parasitoidea. These large, predaceous mites are well represented in edaphic habitats in the Mixedwood Plains. Species of the veigaiid genus Veigaia are common in the deciduous forests of the eastern United States and several of them reach the northern limits of their distribution in the Mixedwood Plains and Atlantic-Maritime Ecozones. Many species of Parasitidae (Fig.M-3) are associated with animal nests, carrion and dung and their deutonymphs disperse by phoresy on insects or mammals.

A predatory parasitid mite Parasites sp.

Rhodacaroidea. Species of 4 families occur in the Mixedwood Plains. Members of Rhodacaridae, Ologamasidae and Halolaelapidae, and some species of Digamasellidae are edaphic or nidicolous predators. Other digamasellid species inhabit subcortical insect galleries or accumulations of organic matter such as manure and compost. Many rhodacaroid species disperse as phoretic deutonymphs on their insect associates or on their mammalian nidicolous associates.
Eviphidoidea. This superfamily is also represented in the Ecozone by members of 4 families. Species of Eviphididae, Macrochelidae, Pachylaelapidae and Parholaspidae are predators in edaphic habitats. Many of them are prevalent in compost, manure and carrion, often in association with insects which act as hosts during a phoretic stage of their life history.
Ascoidea. This large taxon is richly represented in the Mixedwood Plains by members of 5 families. Ascidae are mostly predators in edaphic, subcortical, arboreal, nidicolous and subaquatic habitats. Species of this family are often closely associated with insects or mammals, and disperse by phoresy on these hosts as adult females. Fungivorous species of the genus Hoploseius, phoretic on tiny flies of the genus Mycodrosophila , are common on bracket fungi in the southeastern United States; they occur in Canada only in the Mixedwood Plains (Lindquist, 1963). On the other hand, nematophagous members of Arctoseius, (Fig.M-4) phoretic on various nematoceran flies, have boreal distributions, some with southern limits in this Ecozone. Although most species of the ascid genus Proctolaelaps appear to prey on nematodes, members of one species group, represented by two species in
the Mixedwood Plains, are specialized, pollen-feeding associates of bumble bees (Lindquist and Evans, 1965). These mites are phoretic on adult bumble bees, and they develop in the bees' nests, where they feed on pollen. Two species of the ascid genus Mucroseius occur with their sawyer beetle hosts, Monochamus spp., on conifers in the Mixedwood Plains. Female mites are phoretic on adult sawyers, and they may have a tritrophic association with their hosts and the potentially highly destructive pine wood nematode, Bursaphelenchus xylophilus , which the sawyers also carry from infected to healthy conifers (Lindquist and Wu, 1991). Whether these mites prey on the nematodes or the sawyer's immature instars in the galleries of the sawyers has not been determined. Predaceous mites of the relatively well known family Phytoseiidae are the dominant arboreal Mesostigmata in the Mixedwood Plains, and certain species are important regulators of spider mite populations. Several other species occur in litter and edaphic habitats. Two species of the unusual family Otopheidomenidae, parasites in the tympanic cavities of moths, have been reported from the Ecozone. Ameroseiidae are atypical among Mesostigmata in that they feed on fungal spores and pollen grains. Members of several species occur in compost, bracket fungi and nidicolous habitats in the Mixedwood Plains. A single collection of a species Podocinum from a Carolinian woodland habitat in the Lake Erie Lowlands represents the most northern record of the predaceous family Podocinidae in eastern North America (Lindquist and Wu 1987).

A predatory ascid mite, Arctoseius sp.

Dermanyssoidea. This group is also well represented in the Mixedwood Plains. The families Laelapidae and Haemogamasidae include both free living predatory species associated with social and subsocial insects in edaphic and nidicolous habitats and parasitic species (facultative or obligatory) on birds and mammals. The unusual pollen feeding laelapid species Melittiphis alvearius (Berlese) and the parasitic varroid species Varroa jacobsoni (Oudemans) have invaded the Ecozone recently in association with their introduced host, the honey bee Apis mellifera. Parasitic species of 7 other dermanyssoid families have been recorded from reptilian, bird or mammal hosts in the Mixedwood Plains. One of them, Liponyssoides sanguineus (Hirst), attacks mice and rats in the Ecozone and is a potential vector for the rickettsia responsible for rickettsialpox in humans.
Diarthrophallina - Diarthrophalloidea. These highly modified mites live commensally under the elytra of passalid beetles. One species, collected in a Carolinian woodlot in the Lake Erie Lowland, represents the most northern record of the family Diarthrophallidae in eastern North America.
Uropodina - Uropodoidea. Predaceous and saprophagous members of 4 families of tortoise mites occur commonly in edaphic, nidicolous and decaying wood habitats in the Mixedwood Plains (Fig.M-5). Many species are associated with insects and disperse as phoretic deutonymphs on their hosts. At least one species of Uroobovella (Fuscuropoda) is known to be a predatory pest in commercial earthworm cultures (Stone and Ogles,

A uropodiod mite, Dinychus sp.

Trigynaspida. About 10 species representing 8 families and 5 superfamilies of this group are associated with myriapods and insects in the Mixedwood Plains, and all of them reach the northern limit of their distribution in eastern North America in the Ecozone.


Parasitiformes - Ixodides

Ixodides, or Metastigmata, includes the familiar ticks which parasitize birds and mammals. A summary of expected species diversity of families represented in the Mixedwood Plains is presented in Table 2.

TABLE 2: EXPECTED SPECIES DIVERSITY OF FAMILIES OF IXODIDA KNOWN FROM CANADA IN MIXEDWOOD PLAINS ECOZONE BASED ON RECORDS OF NAMED AND UNNAMED SPECIES IN CANADIAN NATIONAL COLLECTION OF ACARI

SUPERFAMILYFAMILYHABITAT
OR HOST OF
ADULT
FEEDING
BEHAVIOUR
OF ADULT
# SPECIES
EXPECTED
IN MWP/
CANADA
IxodoideaArgasidaeBirds,
Mammals
Parasitic1/8
"IxodidaeBirds,
Mammals
Parasitic15/32

Of the eight species in three genera of the family Argasidae known or expected to occur in Canada, only one genus and species, Ornithodoros kellyi Cooley & Kohls, is eastern, reaching its northern limits parasitic on bats in the Mixedwood Plains Ecozone.

Members of 32 species in six genera of the family Ixodidae occur in Canada; of these, 15 species in five genera are known or expected to live in the Mixedwood Plains Ecozone. Two species of Amblyomma and three species of Ixodes reach their northern limits in this region, whereas two species of Dermacentor, two other species of Ixodes, and Rhipicephalus sanguineus (Latreille) extend further north in their more general North American distributions (Fig. M-6).

A tick, Ixodes sp.

As obligate blood feeders, ticks harbor and spread an unusual variety of disease- causing pathogens of their hosts, including bacteria, rickettsia, protozoa and viruses. The deer tick, Ixodes scapularis Say (=I. dammini), known as the most competent North American vector of Lyme Disease, has recently extended its range widely into the Mixedwood Plains Ecozone. Perhaps the most commonly encountered tick on people and pets in this region, Ixodes cookei Packard, is a potential vector of the powassan encephalitis virus. The widespread brown dog tick, R. sanguineus, is a vector of rickettsial human Ehrlichiosis and Canine Haemobartonellosis; the American dog tick, Dermacentor variabilis (Say), and the rabbit tick, Haemaphysalis leporispalustris (Packard), are vectors of bacterial tularemia, rickettsial Q fever and Rocky Mountain spotted fever; and D. variabilis is additionally a potential vector of viral colorado tick fever in this region.

Apart from tick-borne pathogens, some ticks cause severe damage to their hosts just from the effects of their feeding. Under natural conditions, large tick infestations may be the direct cause of anemia in small hosts such as mice and rabbits. Severe damage may be done to moose by the winter tick, Dermacentor albipictus (Packard), particularly during long, late winters, by which time many thousands of these large ticks may be on single hosts. Tick salivary components include anticoagulants whose effects may persist well after feeding; other components may act as allergens which lead to exaggerated inflammations, severe itching, and lesions resulting from scratching by hosts. A few ticks, primarily D. andersoni Stiles in Canada, can cause a potentially lethal paralytic effect from their feeding activity alone. Recovery from tick paralysis occurs after removal of the ticks. In Eastern Canada, including the Mixedwood Plains Ecozone, the closely-related D. variabilis may have a similarly paralytic potential on people, domesticated animals and some wild mammals in restricted areas of its distribution.

Acariformes - Oribatida

This suborder, also known as Oribatei or Cryptostigmata, is divided into 6 cohorts, namely, Palaeosomata, Enarthronota, Parhyposomata, Mixonomata, Desmonomata and Brachypylina, and is represented in Canada by 31 superfamilies, all of which occur in the Mixedwood Plains Ecozone. The Canadian fauna was treated at the family level by Marshall (1979), and published distribution records for species known from Canada were given in Marshall et al. (1987) and Behan-Pelletier (1993a). A summary of expected species diversity of families represented in the Mixedwood Plains is presented in Table 7. Known species diversity of oribatid mites in the Mixedwood Plains Ecozone, based on published records, is presented in Table 8.
In the Mixedwood Plains, as in all terrestrial ecosystems, Oribatida, the so called "beetle" or "box" mites, are actively involved in decomposition of organic matter, in nutrient cycling and in soil formation. All active instars of these mites feed on a wide variety of material including living and dead plant and fungal material, lichens and carrion; some are predaceous, but none is parasitic. Oribatida influence decomposition and soil structure by comminuting organic matter and producing faecal pellets, which provide a large surface area for decomposition and are in turn an integral component of soil structure. They disperse bacteria and fungi, both externally on their body surface, or by feeding, with subsequent survival of spores during passage through their alimentary tracts. Many oribatid species sequester calcium and other minerals in their thickened cuticle. Thus, their bodies may form important 'sinks' for nutrients, especially in nutrient limited environments such as peatlands. (Crossley 1977, Norton and Behan-Pelletier 1991). Recent reviews on the role of Oribatida in decomposition and nutrient cycling include those of Wallwork (1983), Seastedt (1984) and Norton (1986).

TABLE 7:EXPECTED DIVERSITY OF FAMILIES OF ORIBATIDA KNOWN FROM CANADA IN MIXEDWOOD PLAINS ECOZONE BASED ON RECORDS OF NAMED AND UNNAMED SPECIES IN CANADIAN NATIONAL COLLECTION OF ACARI

SUPERFAMILY FAMILY HABITAT
OR HOST OF
ADULT
FEEDING
BEHAVIOUR
OF ADULT
# SPECIES
EXPECTED
IN MWP/
CANADA
Palaeacaroidea Palaeacaridae soil, litter fungivorous,
algivorous
1/3
Ctenacaroidea Aphelacaridae soil fungivorous,
algivorous
1/1
Hypochthonioidea Hypochthoniidae litter fungivore,
saprophagous
1/2
" Eniochthoniidae soil, litter fungivorous 1/2
" Mesoplophoridae litter,
decaying
wood
saprophagous 2/5
Protoplophoroidea Cosmochthoniidae moss, lichen, litter algivorous 2/5
" Arborichthoniidae moss, litter unknown 1/1
Brachychthonioidea Brachychthoniidae moss, soil,
litter, lichens
fungivorous,
algivorous
15/50
Atopochthonioidea Atopochthoniidae soil, litter unknown 2/3
Parhypochthonioidea Parhypochthoniidae soil unknown 1/1
" Gehypochthoniidae soil unknown 1/1
Phthiracaroidea Phthiracaridae decaying
wood, litter
saprophagous
(xylophagous)
10/25
Euphthiracaroidea Oribotritiidae decaying
wood, litter
saprophagous
(xylophagous)
2/10
" Euphthiracaridae decaying,
wood, litter
saprophagous
(xylophagous)
4/10
Eulohmannioidea Eulohmanniidae soil, litter unknown 1/1
Epilohmannioidea Epilohmanniidae litter, moss unknown 2/4
Crotonioidea Nothridae moss, litter saprophagous 4/8
" Camisiidae moss, litter,
canopy,
semiaquatic
saprophagous 8/25
" Trhypochthoniidae moss, litter,
semiaquatic,
aquatic
fungivorous,
algivorous
5/10
" Malaconothridae moss, litter,
semiaquatic
fungivorous,
algivorous
4/10
Nanhermannioidea Nanhermanniidae moss fungivorous 2/10
Hermannioidea Hermanniidae moss fungivorous 1/8
Hermannielloidea Hermanniellidae moss, litter fungivorous,
aprophagous
1/8
" Plasmobatidae moss, litter unknown 1/1
Liodoidea Liodidae moss, canopy saprophagous 1/4
Gymnodamaeoidea Gymnodamaeidae dry litter fungivorous,
saprophagous
2/25
Plateremaeoidea Plateremaeidae dry litter,
moss
unknown 1/2
" Licnodamaeidae moss, litter unknown 1/2
Damaeoidea Damaeidae moss, litter fungivorous 10/50
Polypterozetoidea Podopterotegaeidae litter unknown 1/2
Cepheoidea Cepheidae moss, litter saprophagous 5/20
Microzetoidea Microzetidae litter unknown 2/2
Amerobelboidea Amerobelbidae litter unknown 1/1
" Eremulidae litter unknown 1/2
" Damaeolidae litter unknown 1/1
" Eremobelbidae litter unknown 1/3
Eremaeoidea Eremaeidae litter, moss,
lichen
fungivorous 8/35
" Megeremaeidae litter, moss fungivorous 1/6
Zetorchestoidea Zetorchestidae moss fungivorous 1/1
Gustavioidea Tenuialidae moss unknown 2/10
" Liacaridae moss, litter saprophagous 5/25
" Astegistidae moss, litter fungivorous 5/10
" Peloppiidae moss, litter fungivorous 5/25
" Gustavioidea moss, litter unknown 1/3
" Kodiakellidae moss, litter unknown 0/1
Carabodoidea Carabodidae fungi, litter,
decaying
wood
fungivorous 5/25
Tectocepheoidea Tectocepheidae litter fungivorous 2/4
Oppioidea Oppiidae soil, litter fungivorous 20/80
" Caleremaeidae soil, litter unknown 1/3
" Suctobelbidae soil, litter fungivorous 24/45
" Autognetidae soil, litter fungivorous 4/10
" Thyrisomidae soil, litter,
moss
fungivorous 4/15
" Quadroppiidae soil, litter fungivorous 2/5
Hydrozetoidea Hydrozetidae aquatic algivorous,
fungivorous
2/8
" Limnozetidae aquatic,
semiaquatic
algivorous,
fungivorous
10/20
Ameronothroidea Ameronothridae semiaquatic unknown 1/6
" Tegeocranellidae semiaquatic fungivorous 2/4
Cymbaeremaeoidea Cymbaeremaeidae dry litter fungivorous 5/20
" Micreremidae litter unknown 1/1
Licneremaeoidea Licneremaeidae canopy unknown 1/2
" Passalozetidae dry litter fungivorous 1/4
" Scutoverticidae dry litter unknown 0/5
Oripodoidea Parakalummidae litter saprophagous 6/12
" Scheloribatidae soil, litter,
canopy
fungivorous,
predaceous,
saprophagous
12/40
" Oribatulidae soil, litter,
canopy
fungivorous 20/35
" Haplozetidae litter fungivorous,
omnivorous
12/35
" Mochlozetidae canopy saprophagous 2/5
" Oripodidae canopy fungivorous 2/20
Ceratozetoidea Chamobatidae semiaquatic,
moss
saprophagous 2/6
" Euzetidae semiaquatic saprophagous 1/3
" Zetomimidae aquatic,
semiaquaic
fungivorous,
saprophagous
3/6
" Ceratozetidae litter saprophagous,
fungivorous,
predaceous
20/60
" Mycobatidae moss, litter fungivorous,
saprophagous
10/30
" Humerobatidae canopy fungivorous,
algivorous,
predaceous
2/5
Phenopelopoidea Phenopelopidae litter saprophagous 10/25
" Unduloribatidae litter unknown 1/2
Oribatelloidea Oribatellidae litter, moss saprophagous 5/15
Achipterioidea Achipteriidae litter, moss saprophagous 15/40
" Tegoribatidae litter, moss saprophagous 5/25
Galumnatoidea Galumnatidae litter, moss saprophagous,
predaceous
5/25
TOTALS:
31 Superfamilies
80 Families  344/1080



TABLE 8: LIST OF PUBLISHED SPECIES OF ORIBATID MITES IDENTIFIED FROM THE MIXEDWOOD PLAINS ECOZONE


Cohort - Palaeosomata

Superfamily - Palaeacaroidea

Family - Palaeacaridae
Palaeacarus hystricinus Trägårdh


Cohort - Enarthronota

Unplaced Enarthronota:

Family - uncertain
Gozmanyina majestus (Marshall & Reeves)

Family - Arborichthoniidae
Arborichthonius styosetosus Norton

Superfamily - Hypochthonioidea

Family - Hypochthoniidae
Hypochthonius rufulus C.L. Koch

Family - Eniochthoniidae
Eniochthonius minutissimus (Berlese)

Family - Mesoplophoridae
Archoplophora laevis (Jacot)

Superfamily - Brachychthonioidea

Family - Brachychthoniidae
Brachychthonius berlesei erosus (Jacot)
Brachychthonius jugatus (Jacot)
Eobrachychthonius latior (Berlese)
Liochthonius hystricinus (Forsslund)
Liochthonius sellnicki (Thor)
Mixochthonius pilososetosus (Forsslund)
Poecilochthonius spiciger (Berlese)
Sellnickochthonius lydiae (Jacot)
Sellnickochthonius rostratus (Jacot)
Sellnickochthonius zelawaiensis (Sellnick)
Synchthonius crenulatus (Jacot)


Superfamily - Atopochthonioidea

Family - Atopochthoniidae
Atopochthonius artiodactylus Grandjean

Family - Pterochthoniidae
Pterochthonius angelus (Berlese)


Cohort - Parhyposomata

Superfamily - Parhypochthonioidea

Family - Parhypochthoniidae
Parhypochthonius aphidinus Berlese
Parhypochthonius a. octofilamentis Jacot

Family - Gehypochthoniidae
Gehypochthonius rhadamanthus Jacot


Cohort - Mixonomata

Superfamily - Phthiracaroidea

Family - Phthiracaridae
Atropacarus striculus (C.L. Koch)
Hoplophorella granulata (Banks)
Phthiracarus olivaceus Jacot
Phthiracarus setosellus Jacot
Phthiracarus sphaerulus (Banks)
Phthiracarus sp.

Superfamily - Euphthiracaroidea

Family - Euphthiracaridae
Euphthiracarus flavus (Ewing)
Microtritia minima (Berlese)

Superfamily - Eulohmannioidea

Family - Eulohmanniidae
Eulohmannia ribagai (Berlese)

Superfamily - Epilohmannioidea

Family - Epilohmanniidae
Epilohmannia cylindrica (Berlese)
Epilohmannia sp.


Cohort - Desmonomata

Superfamily - Crotonioidea

Family - Nothridae
Nothrus anauniensis Can. and Fanzago
Nothrus monodactylus (Berlese)
Nothrus truncatus Banks
Nothrus spp.

Family - Camisiidae
Camisia sp.
Camisia segnis (Hermann)
Platynothrus spp.
Platynothrus peltifer (C.L. Koch)

Family - Trhypochthoniidae
Mucronothrus nasalis (Willmann)
Trhypochthonius tectorum (Berlese)
Trhypochthonius spp.

Family - Malaconothridae
Malaconothrus sp.
Malaconothrus mollisetosus Hammer

Superfamily - Nanhermannioidea

Family - Nanhermanniidae
Nanhermannia dorsalis (Banks)
Nanhermannia sp. nov.


Cohort - Brachypylina

Superfamily - Hermannielloidea

Family - Plasmobatidae
Plasmobates sp.

Superfamily - Liodoidea

Family - Liodidae
Platyliodes sp.

Superfamily - Plateremaeoidea

Family - Plateremaeidae
Allodamaeus sp.

Superfamily - Damaeoidea

Family - Damaeidae
Belba sp.
Caenobelba alleganiensis Norton
Damaeus spp.
Epidameus canadensis (Banks)
Epidameus longiseta (Banks)
Epidamaeus sp.

Superfamily - Cepheoidea

Family - Cepheidae
Cepheus corae Jacot
Ommatocepheus clavatus Woolley & Higgins

Superfamily - Microzetoidea

Family - Microzetidae
Berlesezetes appalachicolus (Jacot)

Superfamily - Amerobelboidea

Family - Eremulidae
Eremulus cingulatus Jacot

Family -Eremobelbidae
Eremobelba gracilior Berlese

Superfamily - Eremaeoidea

Family - Eremaeidae
Eremaeus appalachicus Behan-Pelletier
Eremaeus brevitarsus (Ewing)
Eremaeus kevani Behan-Pelletier
Eueremaeus columbianus (Berlese)
Eueremaeus proximus (Berlese)
Eueremaeus tetrosus (Higgins)

Superfamily - Gustavioidea

Family - Tenuialidae
Hafenferrefia sp.

Family - Liacaridae
Adoristes ammonoosuci Jacot
Dorycranosus sp.
Liacarus sp.

Family - Astegistidae
Astegistes sp.
Cultroribula bicultrata (Berlese)
Cultroribula divergens Jacot

Family - Peloppiidae
Ceratoppia bipilis (Hermann)
Ceratoppia spinipes (Banks)

Family - Gustaviidae
Gustavia parvula (Banks)

Superfamily - Carabodoidea

Family - Carabodidae
Carabodes brevis Banks
Carabodes erectus Reeves
Carabodes granulatus Banks
Carabodes niger Banks
Carabodes polyporetes Reeves
Carabodes rugosior Berlese
Carabodes wonalancetanus Reeves

Superfamily - Tectocepheoidea

Family - Tectocepheidae
Tectocepheus velatus (Michael)

Superfamily - Oppioidea

Family - Oppiidae
Microppia minus (Paoli)
Microppia simplissimus (Jacot)
Multioppia carolinae (Jacot)
Oppia minuta (Banks)
Oppia sp.
Oppiella nova (Oudemans)
Oppiella translamellata (Willmann)
Oppiella sp.
Quadroppia sp.
Ramusella (I.) maculata (Hammer)

Family - Suctobelbidae
Allosuctobelba sp.
Suctobelba sp.
Suctobelbella frothinghami Jacot
Suctobelbella hurshi Jacot
Suctobelbella laxtoni Jacot
Suctobelbella longicuspis Jacot

Family - Autognetidae
Autogneta longilamellata (Michael)

Family - Thyrisomidae
Banksinoma spinifera (Hammer)

Superfamily - Hydrozetoidea

Family - Hydrozetidae
Hydrozetes sp.

Family - Limnozetidae
Limnozetes atmetos Behan-Pelletier
Limnozetes borealis Behan-Pelletier
Limnozetes ciliatus (Schrank)
Limnozetes guyi Behan-Pelletier
Limnozetes latilamellatus Behan-Pelletier
Limnozetes lustrum Behan-Pelletier
Limnozetes palmerae Behan-Pelletier

Superfamily - Ameronothroidea

Family - Tegeocranellidae
Tegeocranellus muscorum Behan-Pelletier

Superfamily - Cymbaeremaeoidea

Family - Cymbaeremaeidae
Scapheremaeus palustris (Sellnick)
Scapheremaeus parvula (Banks)

Superfamily - Oripodoidea

Family - Scheloribatidae
Scheloribates lanceoliger (Berlese)
Scheloribates pallidulus (C.L. Koch)
Scheloribates sp.

Family - Oribatulidae
Eporibatula sp.
Lucoppia burrowsii (Michael)
Oribatula pallida Banks
Oribatula sp.
Phauloppia sp.
Zygoribatula exilis (Nicolet)
Zygoribatula pyrostigmata (Ewing)
Zygoribatula sp.

Family - Haplozetidae

Peloribates americanus Jacot
Peloribates sp.
Rostrozetes ovulum (Sellnick)
Xylobates capucinus (Berlese)
Xylobates lophotrichus (Berlese)
Xylobates sp.

Family - Mochlozetidae
Dynatozetes magnus (Banks)
Podoribates sp.

Family - Parakalummatidae
Neoribates aurantiacus (Oudemans)
Neoribates sp.
Protokalumma depressa (Banks)
Protokalumma sp.

Superfamily - Ceratozetoidea

Family - Chamobatidae
Chamobates cuspidatus (Michael)
Chamobates sp.

Family - Ceratozetidae
Ceratozetes cuspidatus Jacot
Ceratozetes gracilis (Michael)
Ceratozetes mediocris Berlese
Ceratozetes virginicus (Banks)
Fuscozetes bidentatus (Banks)
Fuscozetes fuscipes (C.L. Koch)
Trichoribates formosus (Banks)
Trichoribates sp.

Family - Humerobatidae
Humerobates rostrolamellatus Grandjean

Family - Mycobatidae
Mycobates hyaleus Behan-Pelletier
Punctoribates sp.

Superfamily - Phenopelopoidea

Family - Phenopelopidae
Eupelops terminalis (Banks)
Eupelops sp.
Peloptulus sp.

Superfamily - Oribatelloidea

Family - Oribatellidae
Oribatella quadricornuta (Michael)
Oribatella reticuloides Hammer
Oribatella sp.

Superfamily - Achipterioidea

Family - Achipteriidae
Achipteria sp.

Superfamily - Galumnatoidea

Family - Galumnatidae
Galumna nigra (Ewing)
Galumna sp.
Pergalumna emarginata (Banks) 1895
Trichogalumna sp.


Superfamilies with several species represented in the Mixedwood Plains, or those of special interest are discussed below.

COHORT PLALAESOMATA
Palaeacaroidea : This group is represented by only one cosmopolitan species, Palaeacarus hystricinus Trägårdh, in the Mixedwood Plains, a species relatively common in litter throughout temperate North America. The absence of males in collections indicates that P. hystricinus reproduces by thelyokous parthenogenesis (Norton et al. 1993).

COHORT ENARTHRONOTA
Hypochthonioidea: (Fig.M-7) Members of this superfamily live in litter. Hypochthonius rufulus (C.L. Koch) is a species with a holarctic distribution, and has been found as far north as Ellesmere Island in the Canadian arctic archipelago. This species feeds primarily on bacteria (Luxton 1972) and is a common inhabitant of peatlands in Canada and the Palaearctic. The widely distributed Eniochthonius minutissimus (Berlese) lives in Canadian and European peatlands, where it is among the most characteristic oribatid mites. It is very common in this habitat in the Mixedwood Plains Ecozone. Recently, the cuticle of both adults and immatures of this species was shown to be hardened by calcium oxalate, probably absorbed from their food (fungi) (Norton and Behan-Pelletier 1991).

A hypochthonioid mite, Hypochthonius sp.


Brachychthonioidea: (Fig.M-8) Representatives of the Brachychthoniidae are common in the Mixedwood Plains. These small mites, generally less than 250µm in size, inhabit moist soil and litter. They are microphytophagous, feeding on fungi and possibly also bacteria (Luxton 1991).

A brachychthoniid mite, Eobrachychthonius
sp.


Unplaced Enarthronota: Gozmanyina majestus (Marshall and Reeves), a tiny mite, with long decorative setae, and Arborichthonius styosetosus Norton, are related to the Hypochthonioidea and Brachychthonioidea, but cannot be placed in any superfamily as yet because classification in this cohort is unresolved. G. majestus has been collected in moist coniferous forest from the Mixedwood Plains east to the Atlantic Maritime ecozone, whereas A. styosetosus is only known from the type locality, deciduous litter in St. Lawrence Islands National Park, and a single site in the Atlantic Maritimes.

COHORT PARHYPOSOMATA
Parhypochthonioidea: (Fig.M-9) Large numbers of the widely distributed Parhypochthonius aphidinus Berlese were collected from bogs in eastern Canada, including the Mixedwood Plains, during Summer and Fall (Behan-Pelletier and Bissett 1994). This species, though holarctic, has not been recorded from peatland habitats in Europe.

A parhypochthoniid mite, Parhypochthonius sp.


COHORT MIXONOMATA
Phthiracaroidea and Euphthiracaroidea: (Fig.M-10) Species in these superfamilies are primarily macrophytophagous (Luxton 1972), feeding on the woody tissue of plants, a food source well represented in the Mixedwood Plains. They hollow out this woody material leaving the space packed with faecal pellets. Adults and immatures of some species prefer conifer needles, whereas those of other species prefer twigs and rotting branches; others burrow into the woody vascular tissue of leaves. Members of these families can close up their bodies, somewhat like a flick-knife, and when closed look like seeds. The euphthiracarid Rhysotritia ardua (C. L. Koch) is widely distributed in a variety of forested habitats, including peatlands, and extends as far north as the subarctic in Canada.

A phthiracarid mite, Atropacarus sp.

COHORT DESMONOMATA
Crotonioidea: (Fig.M-11) Most identified species of Nothridae and Camisiidae from the Mixedwood Plains are holarctic or worldwide in distribution. In this superfamily, as in Brachychthoniidae and Nanhermanniidae, thelytokous parthenogenesis is the general method of reproduction (Norton et al. 1993). They have been considered to be unspecialized feeders (panphytophages), but Siepel (1990) showed that differences in the activity of carbohydrases in the gut is reflected in differences in feeding preferences. For example, Platynothrus peltifer (C. L. Koch), common in moist habitats, can feed on plant remains, but adults and immatures of this species prefer fungi. Species of Nothrus, which are generalized feeders, are known to prefer biotopes with high humidity, as are species in the genera Trimalaconothrus and Malaconothrus. The aquatic oribatid mite Mucronothrus nasalis (Willmann) which inhabits spring-fed lotic environments and spring seepages, is a globally widespread species, known from every continent except Africa and Antarctica (Norton et al. 1996). Hammer (1965) suggested that the extensive distribution of this species is relictual, and that the species is very old, predating the breakup of the supercontinent Pangaea ca. 200 million years ago. Our knowledge of the biology and ecology of this species is based on the study of a population from a small springbrook near Toronto (Norton et al. 1988).

A nothrid mite, Nothrus sp.


Nanhermannioidea: (Fig.M-12) Species of Nanhermannia Berlese are among the most common and abundant oribatid mites in peatlands in northeastern North America, including those of the Mixedwood Plains (Behan-Pelletier and Bissett 1994). An undescribed species of this genus has been mistaken for N. nana (Nicolet) in both Europe and North America, and it is probable that many published references to N. nana actually pertain to the undescribed species (see Marshall et al. 1987).

A nanhermanniid mite, Nanhermannia sp.


Eremaeoidea. (Fig.M-13) Species in this superfamily primarily inhabit dry habitats where they live in organic litter and in arboreal habitats, such as tree trunks. They are microvores, feeding on fungal hyphae and spores. The Eremaeidae are species rich in Canada and six species are now known from the Mixedwood Plains (Behan-Pelletier 1993b).

An eremaeid mite, Eremaeus sp.


Carabodoidea: (Fig.M-14) Many species in the carabodoid genus Carabodes are more common on tree trunks and in the crown of trees than in litter. They form part of the diverse, but poorly known, arboreal oribatid fauna, which includes species in other superfamilies, e.g., Crotonioidea and Oripodoidea. Carabodes species feed on fungi, especially bracket fungi, and lichens on trunks and branches of trees (Reeves 1988).

A carabodid mite, Carabodes sp.


Oppioidea: (Fig.M-15) Representatives of this superfamily are small (usually 180-300_m as adults) and inhabit litter and soil. The most common species are members of the families Oppiidae and Suctobelbidae; the former are fungal feeders and the latter are probably liquid feeders. Luxton (1991) noted low percentages of immatures of Oppiidae with food boluses and suggested that immatures may be liquid feeders also. Oppiella nova (Oudemans) is one of the most widely distributed terrestrial arthropods and is found in both disclimax systems and complex stable associations.

An oppiid mite, Oppia sp.


Ameronothroidea: (Fig.M-16) Tegeocranellus muscorum Behan-Pelletier is known from temporarily wet habitats at Long Point National Wildlife Area (Behan-Pelletier submitted). This species is distributed throughout eastern USA, but Long Point is the only known habitat in Canada. Until the discovery of both the immatures and adults of this undescribed species the superfamily placement of this genus was unclear (Behan- Pelletier submitted). Species in this genus are well adapted for their semiaquatic habitat by having a series of tecta above legs 2 and 3, which hold an air bubble when the mite is submerged.

A An ameronothrid mite, Tegeocranellus
mite, Tegeocranellus sp.


Hydrozetoidea: (Fig.M-17) Members of the closely related families Limnozetidae and Hydrozetidae are aquatic. All known Limnozetidae are parthenogens, whereas a few species of Hydrozetes Berlese are sexual. Species of Hydrozetes are the most abundant oribatids in the open water of peatland pools. They actively move on or in the water by clinging to the surface film. Limnozetes is the most species-rich oribatid genus in the sphagnum moss of bogs in eastern Canada, including those of the Mixedwood Plains Ecozone. Members of this genus often are the only microarthropods in dripping wet sphagnum and peat layers, and up to 100,000 specimens of Limnozetes have been counted from a cubic metre of wet sphagnum (Bocard 1991).

A limnozetid mite, Limnozetes sp.


Oripodoidea: (Fig.M-18) This is one of the most important superfamilies in the Mixedwood Plains Ecozone. Species of the haplozetid genus Xylobates are found in all forest litter and grassy habitats. The cosmopolitan haplozetid species Rostrozetes ovulum (Sellnick) is widely distributed throughout the nearctic and neotropics. Although a frequent inhabitant of bogs in northeastern North America, including the Mixedwood Plains, it has not been recorded from this habitat in other areas. Species in the scheloribatid genus Scheloribates are found in all terrestrial habitats in the Ecozone.

A scheloribatid mite, Scheloribates sp.


Ceratozetoidea: (Fig.M-19) This is diverse superfamily in the Mixedwood Plains, the fauna being dominated by members of the Ceratozetidae. Species are macrophytophages, microphytophages, fungivores; some feed on carrion. Species of Scheloribatidae, Ceratozetidae and Galumnatidae are among the oribatid mites of veterinary importance as intermediate hosts and vectors of tapeworms.

A ceratozatid mite, Ceratozates sp.

Acariformes - Prostigmata

Prostigmata is a large and extremely diverse group with representatives in virtually every type of habitat and community in the Mixedwood Plains Ecozone. The Canadian fauna was discussed at the family level by Smith and Lindquist (1979). A summary of expected species diversity of families represented in the Mixedwood Plains is presented in Table 3. Water mites (Hydrachnida) and spider mites and their relatives (Tetranychoidea) are probably the best known groups of Prostigmata in the Mixedwood Plains Ecozone, and the former are considered in more detail below.

TABLE 3: EXPECTED SPECIES DIVERSITY OF FAMILIES OF PROSTIGMATA KNOWN FROM CANADA IN MIXEDWOOD PLAINS ECOZONE BASED ON RECORDS OF NAMED AND UNNAMED SPECIES IN CANADIAN NATIONAL COLLECTION OF ACARI

SUPERFAMILYFAMILYHABITAT
OR HOST
OF ADULT
FEEDING
BEHAVIOUR
OF ADULT
# SPECIES
EXPECTED
IN MWP/
CANADA
PachygnathoideaAlicorhagiidaeLitterUnknown2/4
"LordalycidaeLitterUnknown1/2
"SphaerolichidaeLitterUnknown1/2
"NanorchestidaeLitterFungivorous,
Phytophagous
5/10
"PachygnathidaeLitterUnknown4/8
"TerpnacaridaeLitterUnknown2/4
"OehserchestidaeLitterUnknown1/2
NematalycoideaNematalycidaePhreaticUnknown1/5
"MicropsammidaeSoilUnknown0/1
NicoletielloideaNicoletiellidaeLitterPredaceous1/2
EupodoideaEupodidaeLitterFungivorous30/60
"PenthaleidaePlantsPhytophagous1/3
"PenthalodidaeLitterUnknown2/4
"RhagidiidaeLitterPredaceous20/60
TydeoideaTydeidaeLitter,PlantsFungivorous,
Phytophagous
30/60
"EreynetidaeLitter,
Animals
Predaceous,
Parasitic
10/30
BdelloideaBdellidaeLitter, PlantsPredaceous10/20
"CunaxidaeLitter, PlantsPredaceous10/25
HalacaroideaHalacaridaeAquaticUnknown10/100
TarsocheyloideaTarsocheylidaeLitterPredaceous5/10
HeterocheyloideaHeterocheylidaePassalid
Beetles
Parasitic1/1
DolichocyboideaDolichocybidaeInsectsFungivorous5/10
TrochometridioideaTrochometridiidaeLitterFungivorous0/1
"CaraboacaridaeBeetlesUnknown1/1
PyemotoideaPyemotidaeInsectsParasitoid3/10
"AcarophenacidaeInsectsParasitoid4/10
PygmephoroideaSiteroptidaeLitterFungivorous4/10
"PygmephoridaeLitterFungivorous20/60
ScutacaroideaMicrodispidaeLitterFungivorous,
Parasitoid
3/10
"ScutacaridaeLitterFungivorous25/60
TarsonemoideaTarsonemidaeLitter,
Insects,
Plants
Fungivorous,
Parasitic,
Parasitoid,
Predaceous,
Phytophagous
30/100
"PodapolipidaeInsectsParasitic4/10
CheyletoideaCheyletidaeLitter, PlantsPredaceous15/40
"CheyletiellidaeBirds,
Mammals
Parasitic10/25
"CloacaridaeTurtlesParasitic1/2
"MyobiidaeMammalsParasitic10/30
"HarpyrhynchidaeBirdsParasitic10/25
"SyringophilidaeBirdsParasitic250/500
"PsorergatidaeMammalsParasitic20/45
"DemodicidaeMammalsParasitic20/50
RaphignathoideaRaphignathidaeLitterPredaceous20/50
"XenocalligonellidaeLitterUnknown1/1
"CalligonellidaeLitterPredaceous3/8
"CryptognathidaeLitterPredaceous3/8
"EupalopsellidaeLitterPredaceous3/6
"HomocaligidaeLitterPredaceous2/4
"StigmaeidaeLitter, PlantsPredaceous,
Phytophagous
20/40
"CamerobiidaePlants, BarkPredaceous0/1
TetranychoideaTetranychidaePlantsPhytophagous30/55
"TenuipalpidaePlantsPhytophagous10/20
"LinotetranidaePlantsPhytophagous0/1
"TuckerellidaePlantsPhytophagous0/1
EriophyoideaPhytoptidaePlantsPhytophagous30/60
"EriophyidaePlantsPhytophagous300/600
"DiptilomiopidaePlantsPhytophagous25/50
AdamystoideaAdamystidaeLitterPredaceous0/1
"SaxidromidaeLitterPredaceous0/2
CaeculoideaCaeculidaeLitterPredaceous0/2
AnystoideaAnystidaeLitter, PlantsPredaceous10/20
"PseudocheylidaeLitterPredaceous1/2
"PterygosomatidaeInsects,
Reptiles
Parasitic0/2
"PomerantziidaeLitterPredaceous0/2
"BarbutiidaeLitterPredaceous1/1
"TeneriffiidaeLitterPredaceous0/1
"ParatydeidaeLitterPredaceous5/10
CalyptostomatoideaCalyptostomatidaeLitterPredaceous1/1
ErythraeoideaSmarididaeLitterPredaceous6/10
"ErythraeidaeLitter, PlantsPredaceous,
Polleniphagous
25/40
TrombidioideaJohnstonianidaeLitterPredaceous6/10
"TrombellidaeLitterPredaceous1/1
"TrombidiidaeLitterPredaceous30/50
"TrombiculidaeLitterPredaceous25/70
StygothrombidioideaStygothrombididaeFreshwaterPredaceous2/5
HydrovolzioideaHydrovolziidaeFreshwaterPredaceous1/2
EylaoideaPiersigiidaeFreshwaterPredaceous1/2
"LimnocharidaeFreshwaterPredaceous4/4
"EylaidaeFreshwaterPredaceous25/40
HydrachnoideaHydrachnidaeFreshwaterPredaceous25/40
HydryphantoideaHydryphantidaeFreshwaterPredaceous25/50
"HydrodromidaeFreshwaterPredaceous1/2
"RhynchohydracaridaeFreshwaterPredaceous1/2
LebertioideaSperchontidaeFreshwaterPredaceous20/50
"TeutoniidaeFreshwaterPredaceous1/3
"RutripalpidaeFreshwaterPredaceous0/1
"AnisitsiellidaeFreshwaterPredaceous3/10
"LebertiidaeFreshwaterPredaceous20/50
"OxidaeFreshwaterPredaceous10/12
"TorrenticolidaeFreshwaterPredaceous15/50
HygrobatoideaLimnesiidaeFreshwaterPredaceous15/25
"HygrobatidaeFreshwaterPredaceous15/40
"FeltriidaeFreshwaterPredaceous15/35
"UnionicolidaeFreshwaterPredaceous30/40
"PionidaeFreshwaterPredaceous55/75
"AturidaeFreshwaterPredaceous25/75
ArrenuroideaMomoniidaeFreshwaterPredaceous2/7
"NudomideopsidaeFreshwaterPredaceous1/2
"MideidaeFreshwaterPredaceous1/5
"MideopsidaeFreshwaterPredaceous16/25
"ChappuisididaeFreshwaterPredaceous1/5
"AthienemanniidaeFreshwaterPredaceous2/6
"AcalyptonotidaeFreshwaterPredaceous1/4
"NeoacaridaeFreshwaterPredaceous7/10
"LaversiidaeFreshwaterPredaceous1/1
"KrendowskiidaeFreshwaterPredaceous4/4
"ArrenuridaeFreshwaterPredaceous90/150
TOTALS:
33 Superfamilies
105 Families  1558/3332

Pachygnathoidea (or Endeostigmata). Several tiny, soft-bodied species of these mites, representing 7 families, are commonly found in edaphic habitats and moss mats in the Mixedwood Plains (Fig.M-20).
Nematalycoidea. One species of Nematalycidae, a group of highly modified vermiform mites adapted for living in deep soil habitats, has been collected in beach sands along Lake Michigan in Illinois, and probably also occurs in the Mixedwood Plains.
Nicoletielloidea. Though not yet collected in the Mixedwood Plains Ecozone, one species of the nicoletiellid genus Labidostomma (Fig.M-21) probably occurs in extreme southern Ontario. Members of this large, heavily sclerotized, predaceous species are common in the deciduous forests of the northeastern United States.

A pckygnathoid mite, Nanorchestes sp. A predatory nicoletielloid mite, Labidostomma sp, feeding on a springtail


Eupodoidea. Many soft-bodied species of these mites, representing the fungivorous Eupodidae, phytophagous Penthaleidae and predaceous Rhagidiidae, occur in edaphic habitats in the Mixedwood Plains. One species of penthaliid mite, Penthaleus major (Dugès), is a common minor pest of legume crops in the Ecozone.
Tydeoidea. Two families are well represented in the Mixedwood Plains. Species of fungivorous Tydeidae are among the most common mites in soil and litter and members
of pollen-feeding species are often very abundant on foliage. Most species of Ereynetidae found in the Ecozone are predators in edaphic habitats, but some are ectodermal cavity parasites of gastropods, amphibians, birds and mammals.
Bdelloidea. Several species of both Bdellidae and Cunaxidae are among the common predatory mites in litter and herbaceous ground cover habitats in the Mixedwood Plains. A species of the bdellid genus Trachymolgus collected at St. Lawrence Islands National Park is the most northern record of this genus in eastern North America. Some cunaxid species are primarily arboreal and may contribute to the regulation of populations of phytophagous mites.
Halacaroidea. A few species of freshwater Halacaridae occur in benthic and interstitial habitats in both streams and lakes in the Mixedwood Plains. This family is unusual among Acari in that most of its species and clades are marine.
Tarsocheyloidea. Members of about 5 species of the predaceous tarsocheylid genera Hoplocheylus and Tarsocheylus inhabit litter and decaying wood habitats along the north shore of the Great Lakes.
Heterocheyloidea. One species of the heterocheylid genus Heterocheylus is a subelytral parasite of passalid beetles in the Lake Erie Lowland. Both Tarsocheylidae and Heterocheylidae apparently reach the northern limit of their distributions in the Mixedwood Plains Ecozone.
Dolichocyboidea. A small number of fungivorous or possibly parasitoid species of the family Dolichocybidae are associated with insects in the Mixedwood Plains.
Trochometridioidea. Species representing 2 families are associated with insects in the Mixedwood Plains, as fungivorous and possibly parasitoid symbionts. A species of Caraboacaridae represents the most northern record of the family in eastern North America.
Pygmephoroidea. Fungivorous species of the families Siteroptidae and Pygmephoridae are well represented in the Mixedwood Plains in litter habitats and nests of both social insects and mammals. These mites disperse as phoretic adult females on their hosts. Members of the genus Siteroptes are vectors of fungal pathogens that attack stored grain.
Scutacaroidea. Species of Microdispidae and Scutacaridae occur commonly in edaphic habitats and decaying wood in the Mixedwood Plains. Most members of this superfamily have similar habits to pygmephoroid mites, and the phoretic females are frequently found on host insects. A few are thought to be parasitoids of their insect associates (Kaliszewski et al. 1995).
Pyemotoidea. Adult females of Pyemotidae and Acarophenacidae are parasitoids of eggs and larvae of bark- and seed-infesting beetles and of grain- and flour-infesting beetles and moths, among other holometabolous insects. Their parasitoid way of life has led to maximizing the female/male sex ratio at 95 to 99 percent females. The bites of some Pyemotes are so toxic that they can cause severe dermatitis on people handling infested material such as hay.
Tarsonemoidea. Numerous species of the ecologically diverse family Tarsonemidae have been reported from the Mixedwood Plains. Many tarsonemids are fungivorous, and some have interesting tritrophic associations with insects. In the Mixedwood Plains Ecozone, several species of Tarsonemus and Heterotarsonemus are associated with bark beetles in conifers (Lindquist, 1969b, 1970). Adult females of these species are phoretic on the beetles, and both the mites and the beetles carry propagules of fungi, including the blue-staining fungi highly injurious to conifers, upon which the mites depend for sustenance and the beetles for pathogenically weakening their tree hosts (Bridges and Moser, 1986). At least three tarsonemid genera in Canada contain plant feeders capable of causing economic damage. The cyclamen mite, Phytonemus pallidus (Banks) damages a wide variety of herbaceous plants, particularly in greenhouses and nurseries. Members of Steneotarsonemus, including several species in the Mixedwood Plains, feed on monocotyledonous plants (Lindquist 1986). Some species of Dendroptus may prey on eriophyid mites within their galls on deciduous-leaved trees, while some of Acaronemus, including A. destructor (Smiley and Landwehr) in the Mixedwood Plains Ecozone, are known to prey on eggs of tetranychoid mites on conifers (Lindquist and Smiley 1978). At least four species of Iponemus, highly host-specific egg parasitoids of ipine bark beetles, occur in the Mixedwood Plains (Lindquist 1969a); two of these species reach the northern limits of their distribution in the Ecozone. Acarapis woodi (Rennie) has recently been introduced into the Ecozone in association with its host, the honey bee. Members of the family Podapolipidae are all parasites of insects. Of the few species known as yet from Canada, one lives under the elytra of carabid beetles in the Mixedwood Plains Ecozone.
Cheyletoidea. Several species of free- living predaceous Cheyletidae are common in litter and arboreal habitats in the Mixedwood Plains. Numerous species representing 7 other families, all parasites of vertebrates, occur in the Ecozone, but have received very little attention. For example, although at least 250 species of syringophilid quill mites probably parasitize species of birds that reside or breed in the Mixedwood Plains, there are no published records of the family from the Ecozone. Two species of Demodicidae, the follicle mites, Demodex folliculorum (Simon) and D. brevis Akbulatova, infest the facial pores of a substantial percentage of the human population in Canada (Sengbusch and Hauswirth 1986).
Raphignathoidea. About half of the Canadian species of this superfamily, including members of 7 families, inhabit the Mixedwood Plains. Most of them are free-living predators in edaphic and herbaceous ground cover habitats (Fig.M-22), but species of the moderately large stigmaeid genus Eustigmaeus feed on mosses.

A predatory raphignathoid mite, Cryptognathus sp.


Tetranychoidea. The families Tetranychidae (Spider Mites) and Tenuipalpidae (False Spider Mites) are both well represented in the Mixedwood Plains. All species of Tetranychoidea are phytophagous, and several of them reach the northern limit of their distribution along with their host plants in the Carolinian Zone of the Lake Erie Lowland. Some species are important pests of various crops, forest trees and ornamental plants. The two-spotted spider mite, Tetranychus urticae Koch, has a wide range of dicotyledonous hosts and is economically the most important phytophagous mite in Canada. The European Red Mite, Panonychus ulmi (Koch) may cause serious damage to deciduous fruit trees. The Brown Wheat Mite, Petrobia latens (Müller) and Clover Mite, Bryobia praetiosa Koch injure grasses and herbaceous plants (Fig.M-23). Several species of Oligonychus, such as the spruce spider mite, Oligonychus ununguis (Jacobi) on coniferous hosts, can be significant pests of trees in the Ecozone.

A spider mite, Bryobia sp.


Eriophyoidea. Most species of perennial vascular plants in the Mixedwood Plains are host to one or more species of these tiny phytophagous mites (Fig.M-24). Most eriophyoids are highly host specific, and their feeding often causes characteristic symptoms on the host ranging from discolouration of foliage to production of striking galls or erineal excretions on foliage, flowers or fruits (Lindquist et al. 1996) (Fig. M-25). As in the case of spider mites, several species of eriophyoids reach the northern limits of their distribution with their host plant in the Carolinian Zone of the Mixedwood Plains [eg. Aceria nyssae (Trotter) on Black Gum, Tetra liriodendronis Keifer on Tulip Poplar, Aculops gleditsiae Keifer on Honey Locust, etc.]. Some Phytoptidae, especially species of the genera Trisetacus and Nalepella, are significant pests of conifers and may seriously damage plantations throughout the Ecozone. Several Eriophyidae are pests on crop and ornamental plants and infestations of some of these species, such as the recently introduced pest of greenhouse tomatoes Aculops lycopersicae (Tryon), occasionally cause serious economic losses. Diptilomiopidae have been relatively little studied in eastern North America but many species appear to reach their northern distributional limits in the Mixedwood Plains. One species, the big-beaked plum mite Diptacus gigantorhynchus (Nalepa), is frequently encountered on both native and cultivated species of Prunus.

Aneriphyoid mite, Aculus sp. An example of leaf galls caused by an exiophyoid


Anystoidea. The most conspicuous members of this superfamily in the Mixedwood Plains are species of the anystid genus Anystis. These so-called whirligig mites are highly active predators of phytophagous mites and other microarthropods in herbaceous ground cover and arboreal habitats. Members of 3 other families, Pseudocheylidae, Barbutiidae and Paratydeidae, are thought to be predators in edaphic habitats. Species of these families are rarely collected in the Mixedwood Plains and appear to reach the northern limit of their distributions in the Ecozone.
The remaining 11 superfamilies of Prostigmata present in the Mixedwood Plains belong to the subcohort Parasitengona. Mites of this well defined clade share an essentially holometabolous life history pattern with ectoparasitic larvae, active and predaceous deutonymphs and adults, and quiescent protonymphal and tritonymphal instars.
Calyptostomatoidea. One species of the worldwide genus Calyptostoma is commonly found in wet edaphic habitats in the Mixedwood Plains. Larvae are parasitic on adult crane flies (Tipulidae).
Erythraeoidea. Species of Smarididae and Erythraeidae are common in litter and herbaceous ground cover habitats in the Mixedwood Plains. Their larvae typically parasitize various insects, but those of the erythraeid genus Leptus are also frequently found on other arachnids. Larvae of Balaustium are atypical among Parasitengona in having secondarily become predaceous on other mites and aphids, and also feed on pollen grains in arboreal habitats. Most of the species of Smarididae recorded from the Mixedwood Plains reach the northern limit of their known distribution in the Ecozone.
Trombidioidea. Species of 4 families are among the most conspicuous mites in litter and edaphic habitats in the Mixedwood Plains (Fig.M-26). Larvae of Johnstonianidae, Trombellidae and Trombidiidae parasitize a wide variety of insects (Fig.M-27), while those of Trombiculidae are the infamous chigger mites that parasitize vertebrates and occasionally attack humans. Adult johnstonianids and trombellids prefer wet litter and moss habitats near bodies of water, but trombidiids (velvet mites) and trombiculids are found on various substrates in woodlands, savannahs and fields.

A predatory velvet mite, Trombidium sp. A parasitiec larval velvet mite, Allothrombium  sp., on an aphid, host.


Stygothrombidioidea. At least 2 undetermined species of this enigmatic group occur in hyporheic aquatic habitats in the Mixedwood Plains. Larvae parasitize stoneflies (Plecoptera) and the elongate, vermiform deutonymphs and adults are predators in subterranean water. These mites appear to have affinities with both trombidioid taxa and true water mites, considered below, but precise relationships have not yet been worked out.
Water Mites (Hydrachnida or Hydrachnellae) Water mites comprise 7 superfamilies belonging to the cohort Parasitengona, along with Stygothrombidioidea and their terrestrial relatives in the superfamilies Trombidioidea, Calyptostomatoidea and Erythraeoidea. This large and diverse clade is characterized by an essentially holometabolous life history (Fig. M-28). Larval water mites are ectoparasites of aquatic insects (Fig. M-29, Fig. M-30) while the active post-larval instars (deutonymphs and adults) are fully aquatic predators of small invertebrates. The parasitic associations of larvae were discussed by Smith and Oliver (1976, 1986) and B. Smith (1988). A detailed case-study of water mite superfamilies is presented below.

Diagrammatic representation of a water mite life history. Parasitic larval water mites (Aturus sp.) on a chironomid host. Parasitic larval water mites, (Hydrachua sp.) on a giant water bug host (Lethocerus sp.)

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