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Important Acarine Families

Abstract
The subclass Acari (mites and ticks) belong to the assemblages of terrestrial
chelicerates in class Arachnida. They are quite small in size: the smallest are
among the plant mites (about 80 μm long), and the largest are among the preda-
tors (about 13 mm long). Most mite species are 400–800 μm long in adult stage.
Their varied modes of life influenced diversified morphology and functions.
Among arachnids, mites are the only phytophagous and parasitic. More than
55,000 species of different Acari have been described under 124 superfamilies
and 540 families. Among Acari, new species of order Ixodida are found only
infrequently; however, greatest increases in new species are among the Astigmata,
Mesostigmata, Oribatida and Prostigmata. Suborder Opilioacarida is compara-
tively less diverse and lesser known. Not more than 10 % of the total Acari are of
significant economic importance. Diagnostic characters of different families
with the help of figures are discussed in more than 100 families mentioning
important species under each family, but details of most important mites are
given under different relevant chapters. However, general account of ticks, tet-
ranychid mites and eriophyid mites is discussed in this chapter as these groups
needed more attention considering their utmost economic importance.

Keywords
Astigmata • Mesostigmata • Oribatida • Prostigmata • Trombidiformes

The Subclass Acari belong to the assemblages of terrestrial chelicerates in


Arachnida, whose representatives have secondarily adapted to an aquatic habitat.
Mites are quite small in size: the smallest are plant mites (about 80 μm long), and
the largest are predators (about 13 mm long). Most mite species are 400–800 μm
long in adult stage. Their varied modes of life influenced diversified morphology
and functions. Mites are the only phytophagous and parasitic arachnids. More than
55,000 species of different Acari in 540 families and 124 superfamilies have been

© Springer Science+Business Media Singapore 2016 73


M.S. Dhooria, Fundamentals of Applied Acarology,
DOI 10.1007/978-981-10-1594-6_6
74 6 Important Acarine Families

described, and many new species are being described routinely from different geo-
graphic regions. Among Acari, new species of Ixodida are found only infrequently.
But greatest increases in new species are among the Astigmata, Mesostigmata,
Oribatida and Prostigmata. Suborders Opilioacarida and Holothyrida are compara-
tively less diverse and less known too. Not more than 10 % of the total Acari are of
significant economic importance (Krantz and Walter 2009; Walter and Proctor
2013). In this chapter brief characteristics of different families to which the impor-
tant species belong are discussed under different superfamilies. Economically
important species along with their hosts/habitats are given under each family, but a
detailed account of each most important species is given under relevant chapters.
However, ticks, tetranychid mites and eriophyid mites are being discussed in more
detail in this chapter as they needed more attention considering their utmost eco-
nomic importance.

6.1 Superorder Parasitiformes

6.1.1 Order Holothyrida

6.1.1.1 Superfamily Holothyroidea


Adults are fully sclerotized with dome-shaped holodorsal shield with short, broad
peritremes above legs. Tritosternum is flagelliform, if present; hypostome with cor-
niculi; chelicerae three segmented; palpi five segmented. They are relatively large
mites (>2 mm). The superfamily Holothyroidea consists of three families, namely,
Allothyridae, Holothyridae and Neothyridae. Adult holothyrids are large (2–7 mm),
heavily sclerotized and found in leaf litter, in mosses and under stones in moist
forests. Some species are scavengers and even predaceous in habit.

6.1.2 Order Opilioacarida

6.1.2.1 Superfamily Opilioacaroidea


They have leathery cuticle with purple or bluish markings but without distinct plates
and four pairs of dorsolateral spiracles with no peritremes; females have well-­
developed extrusible ovipositor terminating in two to three lobes, paired rutella and
discoid or biramous With’s organ lateroventrally. Leg coxae free; trochanters III–IV
and tarsi I–IV are divided in tritonymphs and adults. Opilioacarids are covered in
only one family, Opilioacaridae. They are 1500–2300 μm in length, with long legs,
having leathery cuticle with distinctive blue or purple pigment patterns on the body
and legs. They live in caves, litter or under rocks in both semiarid and forest litter in
tropical and warm temperate regions. Opilioacarids have been reported to feed on
pollen, fungal spores and fragmented arthropod remains. Caribeacarus, Neoacarus
and Phalangiacarus are important genera.
6.1 Superorder Parasitiformes 75

6.1.3 Order Ixodida

6.1.3.1 Superfamily Ixodoidea


Idiosoma with a few or no sclerites but with a thick cuticle, with or without a
podonotal shield. Integument folded or striate, leathery or mammillated/tubercu-
lated. Festoons may or may not be present on the posteromarginal region.
Gnathosoma anterior or anteroventral in position. Hypostome modified as holdfast
organ; palpi simple, telescoped or normally extended with three to four segments
but without palpal apotele. Postlarval stages with dorsolateral stigmatic plates
between III and IV or behind coxae IV ((Sonenshine 1993; Sonenshine et al. 2002);
Sonenshine and Roe 2014). Haller’s organs on the distidorsal aspect of tarsus I; all
tarsi with claws. Divided into three families, namely, Ixodidae, Argasidae and
Nuttalliellidae (ER10 and ER11 and ER12).

Family Ixodidae (Fig. 6.1a, b) The members are commonly known as ‘hard ticks’.
Scutum (shield) present in female larvae, nymphs and adults is small, while in males
it is large. Mouthparts (capitulum) are visible dorsally in larvae, nymphs and adults.
Palps are club shaped; chelicerae have denticulate sheaths and no coxal glands. It is
generally accepted that the Ixodidae are divided into two lineages, the Prostriata
which consists of a single genus Ixodes and the Metastriata (all other genera). Brief
account of different genera belonging to hard ticks is given below (ER13).

Amblyomma Anal groove lies below anus, palpi longer than wide, ornate ticks, eyes
present, festoons present, rectangular basis capituli and oval idiosoma. A. variega-
tum Fabricius and A. hebraeum Koch cause major economic losses in cattle
industry.

Dermacentor Palpi wider than long, ornate, eyes present, festoons present and
basis capituli rectangular. D. marginatus Sulzer infests cattle and sheep and is
worldwide in distribution.

Haemaphysalis Anal groove below anus, eyes are absent, palpi wider than long, not
ornate and festoons present. H. punctata (Canestrini and Fanzago) is an important
pest of sheep.

Hyalomma Eyes present, palpi longer than wide, not ornate and festoons present.
Important species are H. aegyptiacum Linnaeus (tortoise ticks) and H. marginatum
Pomerantsev (Mediterranean ticks).

Ixodes Anal groove above anus, eyes absent, palpi longer than wide, not ornate and
no festoons. Ixodes ricinus Linnaeus (castor bean tick) is the most important and is
found worldwide. It parasitizes sheep, deer, horses and humans.
76 6 Important Acarine Families

Rhipicephalus Anal groove below anus, palpi wider than long, eyes present, not
ornate, festoons present and basis capituli hexagonal. R. appendiculatus Neumann
(brown ear ticks) and R. sanguineus Latreille (brown dog tick) are important
species.

Family Argasidae (Fig. 6.1c, d) The members included in this family are com-
monly known as ‘soft ticks’. Scutum (shield) is absent; mouthparts (capitulum) are
not visible dorsally in nymphs and adults but are seen in larvae. Palps of soft ticks
look like legs. Chelicerae in these ticks have smooth sheaths; coxal organs are pres-
ent. Brief characteristics of important genera of soft ticks are given below.

Antricola Dorsal integument tuberculated, hypostome scoop-like and exclusively


associated with cave dwelling bats. The capitulum of these ticks cannot be seen in
dorsal view because it lies within a groove or depression called a camerostome. A.
marginatus (Banks) is a parasite of bats.

Fig. 6.1 Ixodidae: (a) Dorsum. (b) Venter, Argasidae: (c) Dorsum. (d) Venter
6.1 Superorder Parasitiformes 77

Argas Provided with an obvious marginal sutural line separating dorsal and ventral
body surfaces. A. persicus Oken is an economically important pest of fowls.

Ornithodoros Dorsal integument mammillated and hypostome usually with teeth


but never scoop-like. Mainly associated with bats. O. coriaceous Koch infests
mainly ears of cattle, but also attacks human beings.

Otobius Nymphs with dorsal integument beset with spines and adults with granular
integument and vestigial hypostomes. O megnini Duges, also known as spinose ear
tick, is an important species feeding within the ears of many mammals.

Family Nuttalliellidae (Fig. 6.2) This is a monotypic family characterized by fea-


tures, namely, intermediate to those of the two major tick families reported earlier.
Only one species, Nuttalliella namaqua Bedford, is reported from nests of swallows
from South Africa and Tanzania. These ticks have a leathery, mammillated or granu-
lar integument and lack a dorsal scutum. Hypostome is short with few denticles;
palpi consist of three segments.

Important Website on Ticks Considering the great economic importance of ticks


as most important parasites and disease transmitters in human beings, veterinary
animals, reptiles and birds, affecting their health badly, a website TicksBase (http://
www.icttd.nl/) has been initiated at Utrecht University, the Netherlands. This web-
site provides taxonomic catalogue of all known ticks (Acari: Ixodida) of the world
and is facilitated by the international consortium on ticks and tick-borne disease

Fig. 6.2 Nuttalliellidae: (a) Dorsum. (b) Venter


78 6 Important Acarine Families

(ICTTD). Currently, it comprises 889 valid species (702 hard ticks, 186 soft ticks
and one nuttalliellid tick) including their synonyms, distribution data and common
names.

Tick Life Cycle Ticks have complex life cycles involving several blood meals with
the same or different animal hosts. Soft ticks are usually associated with nests, dens,
burrows or roosts of their animal hosts; they usually mate when they are not on a
host. Hard ticks are usually not associated so closely with their hosts but instead are
free-ranging and come in contact with animal hosts only for blood feeding. Hard
ticks usually mate when they are on a host. Because soft ticks take small blood
meals for a short feeding period, the female lays only a few hundred eggs during her
lifetime, with eggs laid at intervals. Hard ticks take large blood meals and lay 6000
or more eggs at one time (ER6 and ER10).
Ticks have four generalized life cycles which are known as one-host, two-host,
three-host and multi-host life cycle. Ticks are often referred to by the kind of life
cycle they have, for example, the American dog tick is a three-host tick. The three-­
host life cycle has been adopted by most of the ixodid ticks, but two-host and one-­
host life cycle is also found in some species of hard ticks. However, the multi-host
life cycle is characteristic of nearly all species of soft ticks. In multi-host life cycle,
many nymphal moults occur and these moults are called intermediate stages.
Individual nymphs feed and moult several times before moulting to the adult stage.
The adult ticks may feed many times. The female ticks may lay small batches of
eggs after each blood feeding.

Tick Removal To remove feeding ticks, dab them with alcohol. If feeding has just
started, and mouthparts are not cemented in, ticks sometimes pull their mouthparts
out. Try covering them with a dollop of menthol shaving cream or ‘Vicks VapoRub’.
It is generally best to take tweezers, grasp tick at the skin level and pull steadily until
the tick is removed. If mouthparts are left in the skin, it causes wound if left
untreated. So always treat with an antiseptic to prevent secondary infection (ER12).

Attachment and Feeding A fully engorged tick can deposit 100–18,000 eggs on
the ground. Normally thousands of tiny six-legged larvae hatch from the batch of
eggs and crawl randomly up grasses, weeds, twigs or low vegetation or walking
over the ground to await or search for a host in the surrounding area. These ticks,
called seed ticks, suck blood. The engorged seed ticks drop off and moult to eight-
legged nymph and again find a host. After several days of feeding, the engorged
nymph drops off its host and moults again. Females of hard ticks may feed from
several days to more than a week (ER6).
The tick slides its pair of slender teeth painlessly into the hosts’ skin and so the
feeding attachment begins. The central holdfast organ (hypostome), covered with
6.1 Superorder Parasitiformes 79

recurved teeth or ridges, is then inserted. Blood sucking begins; secretions from the
ticks’ salivary glands are then painlessly injected into the wound; these secretions
form around the holdfast organ and glue it in place. At this point, the tick cannot
voluntarily detach until its feeding ceases and the secretions stop. The strength of
the holdfast organ helps the ticks resist being scratched or pulled off. When feeding
is complete, the engorged female drops off the host, lays eggs and then dies.

Losses Caused Ticks are extremely dangerous external parasites of warm-­blooded


animals, birds, amphibians, reptiles, veterinary animals and man. Their bites are not
only annoying and occasionally painful but may result in localized skin inflamma-
tion, secondary infection and possible introduction of disease-causing pathogens.
More than 65 disease-causing pathogens are known to be transmitted by ticks. Some
ticks are venomous and produce very painful bites, and some ticks even cause tick
paralysis and lameness in people and animals (ER13).
Both the hard and soft ticks are vectors of over 30 diseases to man, pets, cattle,
sheep, goats and livestock. Many domestic and wild animals are killed by tick-­
borne diseases, such as fowl spirochaetosis, Texas tick fever, relapsing fever, piro-
plasmosis and anaplasmosis. Many other animals are so weakened that they
succumb to other diseases. Argas persicus causes nervousness, weight loss, lowered
egg production and even death in poultry birds due to blood loss. Many species of
hard ticks are significant human disease vectors and are responsible for spread of
Lyme disease and Rocky Mountain spotted fever (RMSF). Tick-borne encephalitis
(TBE) can lead to meningitis and, in serious cases, results in paralysis and deaths.

Management of Ticks No single strategy can control or manage ticks. So the fol-
lowing options need to be tried for their successful management:

• Ticks prefer still, damp, shady areas, so turn on the lights and dehumidifier and
fans.
• Look for ticks where pets (dogs, cats, etc.) are living and sleeping. Search areas
under the edge of rugs, under furniture, and in cracks around baseboards, win-
dows and door frames, or dog boxes.
• Since most ticks only slowly climb to the top of the grass, medium height vegeta-
tion and/or low bushes to await a passing host. So cultivate the land or at least
keep your grass and bush cut low especially around walks, paths, fences, sheds,
trees, shrubs, play areas and other potentially dangerous locations. Keeping the
grass short in tick-infested areas increases tick desiccation during hot weather.
• Protect yourself by wearing hats, long-sleeved pants and shorts, and tuck shirts
into light coloured pants and pants into socks or boots when going into tick-­
infested or suspect areas.
• Repellents. Use any chemical recommended as repellent, permethrin
(Permanone) – a synthetic pyrethroid pesticide, azadirachtin or neem oil and/or
menthol or sulphur powder. Since these chemicals are also toxic, use them
cautiously.
80 6 Important Acarine Families

• Monitoring and Surveying for Ticks. Examine the volunteer persons who wear
protective clothing and walk through tick-infested areas, for the attached ticks.
Also monitor by dragging white cloth over relatively open ground or ‘flagging’
low-level vegetation (i.e. moving the cloth in a waving motion over and through
vegetation) in densely bushy ground. Ticks that are lining up, waiting or questing
for passing hosts cling to the cloth and can be removed for identification and for
counting. Sampling should be done when vegetation is not wet and when ambi-
ent temperature is above 10 °C. Dry ice (solid form of carbon dioxide) can also
be used for most efficient collection of ticks. CO2 vaporizing from the dry ice
attracts ticks onto a white cloth panel on which they are easily visible and can be
removed periodically, or onto a platform lined with double-sided sticky tape on
which they get trapped.
• Dusting. Dust the open grass lawn when it is 6 cm or less with powdered sulphur,
lime, talcum powder or ground up chalk or calcium carbonated, or treat the area
as a last resort with an appropriate labelled pesticide poison, e.g. wettable pow-
der and/or microencapsulated formulations.
• Biological Control. Releases of chalcid parasitic wasp, Ixodiphagus hookeri (=
Hunterellus hookeri), have been found somewhat effective. Female wasp lays
eggs inside an engorged nymphs of the tick and parasitize them. Many species of
spiders and ants reported to eat ticks if they can find them. Several species of
fungi and nematodes are also reported feeding nymphal ticks while they are
moulting on the soil.
• Mechanical Controls. Increase areas of open lawn and sunlight penetration.
Keep lawn mower to a height of 7.5 cm or less as it lowers the humidity at ground
level. Get rid of bush, weeds, leaf litter, and other debris since these vegetations
can attract ticks and their host. Use wood chips or gravel to create a barrier
between wooded areas (where ticks are common) and your lawn.
• Habitat Management. Controlled burning of habitat may also help reduce tick
numbers. Promote areas that receive lots of direct sunlight as these are not con-
ducive to high tick populations. A 45 cm perimeter border of gravel or limestone
may prevent movement of ticks from grass areas into buildings. Cracks and crev-
ices around the base of buildings should be sealed with caulk. In the indoors,
routinely thorough vacuum cleaning of cracks, furniture, ceilings, floors/carpets,
wall behind pictures, bookshelves and drapes should be done. Spot treatment
with talcum powder or medicated body powder, and also crack and crevice treat-
ment should be done with residual dusts or silica gel or food grade diatomaceous
earth.

6.1.4 Order Mesostigmata (Krantz and Walter 2009)

6.1.4.1 Superfamily Ascoidea

Family Ameroseiidae (Fig. 6.3) Females measure 300–500 μm. The dorsal shield
is entire and usually bears 29 pairs of setae. The sternal shield of female bears two
to three pairs of setae; the genital shield is usually wedge shaped. Podal sperma-
6.1 Superorder Parasitiformes 81

Fig. 6.3 Ameroseiidae

theca is present. The corniculi are conspicuous and usually forked distally. They are
found in cracks, on the surface of baled hay, or feed on moulds which grow under
damp situations. Kleemania plumosa (Oudemans) is a cosmopolitan inhabitant of
stored food and in nests of social insects where it feeds on fungi.

Family Melicharidae (Fig. 6.4) Dorsal shield is entire; fixed cheliceral digit with
pilus dentilis modified into a hyaline flap, found as predators. Many species have
adapted to feed on fungi, pollen and nectar. Melichares agilis (Hering) is commonly
found in cultivated soil and in insect-infested stored grains and on dried fruits.

Family Ascidae (Fig. 6.5) Members are small to medium sized (300–500 μm) and
pale yellow to brown in colour. Chelicerae are dentate, serrate or rarely with
regressed fixed digit. Palp tarsus possesses two-tined apotele. The idiosoma has one
shield or is divided into two shields. Genital opening in males is found at the base
of tritosternum. The legs are long terminating in a pair of claws. They are preda-
ceous on soil mites and plant mites. Some species are very good biocontrol agents
of soil-inhabiting pests in greenhouses. Lasioseius althiasae (De Leon) is a good
predator of soil arthropods and nematodes. Blattiscocius tarsalis (Berlese) is
reported to feed on eggs of moth, Ephestia kuehniella, in flour mills.

6.1.4.2 Superfamily Phytoseioidea

Family Otopheidomenidae (Fig. 6.6) Otopheidomenidae is closely related to the


family Phytoseiidae. They have a reduced fixed digit of chelicerae, a terminally
located anus and a reduced number of setae and are found in the ­thoraco-­abdominal
and wing axillary areas of noctuid and sphingid moths. Otopheidomenis spp. and
82 6 Important Acarine Families

Fig. 6.4 Melicharidae

Fig. 6.5 Ascidae


6.1 Superorder Parasitiformes 83

Fig. 6.6 Otopheidomenidae

Noctuiseius spp. are found in tympanic recess of moths. Heavy infections of


Hemipteroseius indicus (Evans) sometimes may paralyze red cotton bugs.

Family Podocinidae (Fig. 6.7) Measures 430–600 μm. Leg I greatly elongated
with genual, tibial and tarsal segments attenuate, subequal; palp tarsal claw usually
three tined, with one to two apical, whip-like seta. They possess a phytoseiid-­type
spermathecal system. They inhabit humus and litter habitats throughout the tropics.
Podocinum pacificum Berlese is reported from rodent nests, ant nests and grain field
debris. P. sagax (Berlese) is sometimes reported voraciously feeding on
collembolans.

Family Blattiscocidae (Fig. 6.8) Fixed cheliceral digit with setiform pilus dentilis;
movable cheliceral digit lacking a ventral mucro, peritrematic shield broadly fused
posteriorly to exopodal plate curving behind coxae IV; epigyneal shield usually
truncate posteriorly. Members have adopted to a broad spectrum of terrestrial, arbo-
real and sub-aquatic habitats. Blattisocius species are common predators of acarid
mites and insect eggs and larvae in stored food and in nests of small mammals and
birds. B. keegani Fox is reported feeding on bagworm pupae.

Family Phytoseiidae (Fig. 6.9) They are medium-sized (250–400 μm) mites with
relatively long legs. Many species are pale, yellowish, orange or dark brown.
84 6 Important Acarine Families

Fig. 6.7 Podocinidae

Idiosoma is covered by a single shield (rarely two shields) with not more than 20
pairs of setae. Leg I terminates in well-developed ambulacra. The stigma opens
between legs III and IV. The male chelicerae bear a spermatodactyl, the shape of
which has diagnostic value.

Life History and Biology The life cycle consists of egg, larva, protonymph,
deutonymph and adult stages. Eggs generally require very high humidity for hatch-
ing. Phytoseiid mites develop faster than spider mites; they can reproduce through-
out the year in tropical and subtropical conditions and in greenhouses under
temperate areas. Under field conditions in temperate areas, phytoseiids overwinter
in protected habitats as fertilized females. Phytoseiid mites feed on a variety of food
and have developed different feeding habits. They are known to be common inhabit-
ants of leaf domatia. Neoseiulus, Amblyseius, Galendromus, and Phytoseiulus are
the well-known genera. Most phytoseiids feed on a variety of products and small
arthropods (including thrips, whiteflies, scale crawlers). Phytoseiid mites are mainly
predators of spider mites; some species have also been reported feeding on nema-
todes, fungus and pollen. Some species are reported useful as predators of thrips
particularly under greenhouse conditions. They are fast-running mites. Phytoseiids
are the best known predators among the Acari and are being commercially used for
6.1 Superorder Parasitiformes 85

Fig. 6.8 Blattiscocidae: (a) Dorsum. (b) Venter

biological control of spider mites and are sold by many insectaries particularly for
control of T. urticae in greenhouse crops (cucumbers, peppers, tomatoes, strawber-
ries and some ornamentals).

In general, phytoseiid mites have a high intrinsic rate of increase (rm). Species
like Phytoseiulus persimilis are usually specialized predators of Tetranychus spe-
cies. These predators have been most widely used in applied biological control of
spider mites. They have relatively low food requirements for development and
reproduction, and this quality accounts for their efficiency even at low prey
densities.

6.1.4.3 Superfamily Dermanyssoidea

Family Dermanyssidae (Fig. 6.10) Elongate second cheliceral segment of female


far exceeding basal cheliceral segment I length. In case of male chelicerae, second
segment of chelicerae is with normal length. Idiosoma is broadly rounded posteri-
orly. Dermanyssids are parasites of mammals and birds. Dermanyssus gallinae (De
Geer) and Ornithonyssus sylviarum (Canestrini and Fanzago) are important pests of
poultry.
86 6 Important Acarine Families

Fig. 6.9 Phytoseiidae: (a)


Dorsum. (b) Venter

Family Laelapidae (Fig. 6.11) Members of this family have dorsal shield with
more than 23 pairs of setae; female genital shield is flask shaped. Stigma is located
between legs III and IV, without straight peritremes. Leg I is usually with ambula-
crum. Soilborne species, Androlaelaps and Stratiolaelaps, serve as natural control
of root worms. Another soilborne predator Hypoaspis aculeifer (Canestrini) is
reported feeding on plant nematodes and scarabaeid beetles. Tropilaelaps clareae
Delfinado and Baker is an important ectoparasite of honeybees.

Family Varroidae (Fig. 6.12a) Fixed digit of chelicerae absent; peritremes of


female short, looped medially or apically and confined to level of coxae III or III–
IV. Varroa destructor Anderson and Trueman is an important parasite of honeybees,
Apis mellifera, and is discussed in detail in a separate chapter concerning bee para-
sitic mites.

Halarachnidae (Fig. 6.12b) Epigyneal shield absent or rudimentary; sternal shield


usually present; and stigmata are located ventrally or are lateroventral in position.
They are parasites of respiratory tracts of terrestrial and marine mammals. Many
members of the family, particularly Pneumonyssus simicola Banks and P. caninum
Chandler and Ruhe, infest the sinuses and nasal passages of monkeys and dogs.
6.1 Superorder Parasitiformes 87

Fig. 6.10 Dermanyssidae: (a) Dorsum. (b) Venter

Fig. 6.11 Laelapidae: (a) Dorsum. (b) Venter


88 6 Important Acarine Families

Fig. 6.12 (a) Varroidae


(b) Halarachnidae

Family Spinturnicidae (Fig. 6.13) Tritosternum absent or represented by tritoster-


nal base remnant; and epigyneal shield narrowed or otherwise reduced. Members
are parasites of bats’ wing membrane. Platyrrhinus and Sturnia are important
genera.

Family Entonyssidae (Fig. 6.14) Sternal and epigyneal shield well developed but
often weakly sclerotized; stigmata are lateral in position. Entonyssus asiaticus Fain
is a lung parasite of snakes.
6.1 Superorder Parasitiformes 89

Fig. 6.13 Spinturnicidae: (a) Venter. (b) Dorsum

Fig. 6.14 Entonyssidae:


(a) Venter. (b) Dorsum.
(c) Chelicerae
90 6 Important Acarine Families

Fig. 6.15 Rhinonyssidae

Fig. 6.16 Macronyssidae: (a) Dorsum. (b) Venter


6.1 Superorder Parasitiformes 91

Family Rhinonyssidae (Fig. 6.15) Sternal shield reduced or absent but with distinct
sternal setae; epigyneal shield well developed or reduced; and stigmata are dorsal in
position. Members are parasites of respiratory tracts of birds. Ptilonyssus mimi George
and Sternostoma tracheacolum (Lawrence) are important nasal parasites of birds.

Family Macronyssidae (Fig. 6.16) Members have edentate but elongate chelic-
erae. Corniculi are membranous; palp trochanter often with a raised mediolateral
keel and with a large non-setigerous spur on leg coxae II; other coxae without spurs
but occasionally with small ridges. They are parasites of mammals, birds and rep-
tiles. Ornithonyssus bacoti (Hirst) and O. sylviarum (Canestrini and Fanzago) are
important parasites of birds and reptiles.

Family Hystrichonyssidae (Fig. 6.17) Second cheliceral segment normally devel-


oped, considerably shorter than greatly elongated basal cheliceral segment.
Hystrichonyssus turneri Yunker and Baker are reported as parasites of porcupines
and snakes.

6.1.4.4 Superfamily Parasitoidea

Family Parasitidae (Fig. 6.18) In case of males, chelicerae with a spermatotreme –


a sperm holding process that is coalesced distally with movable digit. Males usually
with highly developed, sexually dimorphic, spine-like setae, spurs and apophyses
on legs II. Many representatives are predaceous, free-­living or associated with
insects. Parasitellus species are phoretic on bumblebees; and Parasitus spp. are
common phoretics on carabid and silphid beetles.

6.1.4.5 Superfamily Rhodacaroidea

Family Rhodacaridae (Fig. 6.19) Chelicera usually with arthrodial brush.


Palptarsal apotele three tined. They are free-living predators. Commonly found in
surface and subsurface soils. Rhodacarus and Rhodacarellus are important genera.

Family Digmasellidae (Fig. 6.20) Palptarsal apotele two tined. They are predators
and insect associates. Dendrolaelaps fallax (Leitner) are phoretic on sciarid flies.

6.1.4.6 Superfamily Veigaioidea

Family Veigaiidae (Fig. 6.21) Palptarsal apotele three or four tined and with adja-
cent hyaline scale-like process; hypostome with internal malae typically elaborated,
often bilobed and moustache-like. They are free-living predators and are found in
soil and decaying organic matter. Veigaia and Gamasolaelaps species are reported
from upper soil region as predators.
92 6 Important Acarine Families

Fig. 6.17 Hystrichonyssidae

Fig. 6.18 Parasitidae


Fig. 6.19 Rhodacaridae:
(a) Dorsum. (b) Venter

Fig. 6.20 Digmasellidae: (a) Dorsum. (b) Venter


94 6 Important Acarine Families

Fig. 6.21 Veigaiidae

6.1.4.7 Superfamily Uropodoidea

Family Uropodidae (Fig. 6.22) Female genital shield located between coxae II–
IV. Uropoda spp. are found in highly organic, insular deposits of manure and com-
post. Uroobovella marginata is a slow-moving mite found commonly in poultry and
cattle manure.

6.1.4.8 Superfamily Diarthrophalloidea

Family Diarthrophallidae (Fig. 6.23) Deutonymphs and adults with a single dor-
sal shield that is never flanked by marginal shields; dorsal shield with several pairs
of greatly elongated marginal setae; paranal setae also markedly elongated; palpal
claw absent. Troctognathus spp. and Brachytremella spp. are found on passalid
beetles.
6.1 Superorder Parasitiformes 95

Fig. 6.22 Uropodidae

Fig. 6.23 Diarthrophallidae


96 6 Important Acarine Families

6.2 Superorder Acariformes

6.2.1 Order Trombidiformes

6.2.1.1 Suborder Prostigmata

6.2.1.1.1 Superfamily Tetranychoidea

Family Tetranychidae (Fig. 6.24) Members are commonly known as spider mites
as unlike spiders many species spin a web on plants’ leaves, feed and mate inside it.
Members of this family measure 200–900 μm. Morphology of the ambulacrum and
dorsal chaetotaxy are the main characteristics used for the classification of the
Tetranychidae. Leg chaetotaxy and its ontogeny offer additional potential and
appear to be of good phylogenetic value. The structures of the termination of the
‘peritremes’ are sometimes considered important for identification. The tarsi and
tibiae of the legs often carry some ‘tactile’ and fewer sensory ‘setae’. The shape of
the sensilla is often important for species identification. The dorsum of the tarsus I
of most tetranychids has two pairs of closely associated setae termed as ‘duplex
setae’. A single pair of duplex setae is present on the dorsum of tarsus II. Each pair
of the duplex setae consists of a long distal sensory member and short proximal
tactile member. The empodium and tenent hairs (homologous with true claws) vary
between different genera of tetranychid mites (Baker and Tuttle 1970, 1994; Bolland
et al. 1998; Helle and Sabelis 1985). Dorsal body integument pattern, leg length and
shape of the aedeagus many times prove useful in identification (ER3 and ER7).
Bryobiinae, Eurytetranychinae and Tetranychinae are the important subfamilies
under family Tetranychidae. A wide variety of dorsal body integument pattern
occurs in the Bryobiinae, whereas the dorsal body integument pattern of the

Fig. 6.24 Tetranychidae


6.2 Superorder Acariformes 97

web-­spinning Tetranychinae tends to have more or less uniform striation (ER14). In


subfamily Bryobiinae, empodium is without tenent hairs; females with two or three
pairs of anal setae; and males with five pairs of genito-anal setae. But in subfamily
Tetranychinae, empodium is absent, or if present then without tenent hairs; females
with one or two pairs of anal setae and males with four pairs of genito-anal setae.
Subfamily Bryobiinae is divided into three tribes – Bryobiini (with true claws unci-
nate and empodium pad-like), Hystrichonychini (with true claws and empodium
pad-like) and Petrobiini (claws are pad-like and empodium uncinate). Bryobia,
Bryobiella, Hystrichonychus and Porcupinychus are important genera under sub-
family Bryobiini. Subfamily Tetranychinae is divided into three tribes,
Eurytetranychini (empodium claw-like when present; tarsus I with loosely associ-
ated setae or with one pair of duplex setae) and Tenuipalpodini (empodium claw-­
like or split distally, tarsus I with two pairs of duplex setae and tarsus II with one
pair and opisthosoma with f1 in normal dorsal position or absent). Tenuipalpoides
and Tenuipalponychus (tribe Tenuipalpodini); Eurytetranychus, Eutetranychus and
Aponychus (all of tribe Eurytetranychini); and Schizotetranychus, Oligonychus,
Panonychus and Tetranychus (all of tribe Tetranychini) are important genera.
In Tetranychini, legs are never longer than the body, but long legs are found in
some Bryobiinae and in less derived Tetranychinae such as tribe Eurytetranychini.
In the other tetranychid families, a moderate leg length is the rule. In Bryobiinae,
aedeagus is of simple shape. But in Tetranychini, the aedeagus is more complicated
and is formed of a shaft and a knob.

Spinning Behaviour The use of silk and webbing is a major difference between
the Bryobiinae and the Tetranychinae. Little or no webbing is found in
Eurytetranychinae and Bryobiinae, but moderate webbing (Schizotetranychus spp.
and Oligonychus spp.) to complicated webbing (Eotetranychus spp. and Tetranychus
spp.) is found in Tetranychinae. But in many Eurytetranychinae (Panonychus spp.),
the females spin a network of guy ropes over each egg.

Habitat In case of Bryobiinae, most species live on the twigs or on the upper side
of leaves, but many Eurytetranychini mites live on the upper surface of leaves only.
In subfamily Tetranychini, members of most advanced genera (Eotetranychus and
Tetranychus) feed exclusively on the undersurface, but some members of
Oligonychus and Schizotetranychus live on both the upper and lower surfaces of the
leaves. In Panonychus species, immatures feed on lower surface, but adults live on
both the surfaces (Holtzer et al. 1988).

Host Range Many species live on only a few taxonomically close plant species; a
few are monophagous; others are extremely polyphagous. In general, members of
family Tetranychidae infest a wide range of plants belonging to different botanical
groups. Almost all species of Schizotetranychus live on monocotyledonous plants.
Tendency of polyphagy is observed in species living on herbaceous plants, being an
unstable habitat. Highest polyphagy is observed in Tetranychus species belonging
to subfamily Tetranychinae.
98 6 Important Acarine Families

Population Dynamics The intrinsic rate of natural increase, rm, found from life
table data is helpful in determining the potential of increase in a population. Intrinsic
rate of increase is large when the duration of the generation is short and fertility
high; rm of spider mites is dependent on the breeding conditions like nature of the
host plant, surface available to each individual, temperature and humidity. The high-
est rm is found in different spider mites belonging to Tetranychini, particularly
among Tetranychus species which are most prolific. An increase in rm leads to an
increase in the number of annual generations and destruction of its host plants.
Moreover, these species have much more effective means of dispersal. On the con-
trary, most of the mites belonging to Bryobiinae have low rm, and these mites live
on perennial plants having a more stable habitat.

Diversity In family Tetranychidae, about 1272 species have been described in 95


genera. More than one hundred species can be considered as pests under different
agroclimatic conditions. About ten species are considered as major pests of different
crops, fruit trees, etc. The most well-known and widespread species of spider mite
is commonly known as two-spotted spider mite, Tetranychus urticae Koch.

Websites on Spider Mites Dr. Phillip in France has maintained a special website
named as Spider Mites Web (www.montpellier.inra.fr/CBGP/spinweb/). On this
website, taxonomic data relating to history of nomenclature, the geographical distri-
butions (with maps), and the host range of 1272 valid species recorded till November
2011 have been given. In addition about 1350 references on spider mites found from
the year 1758 till date are also listed on the website for use of the acarologists and
entomologists.

Injury by Spider Mites Spider mites injure leaves by piercing cells and sucking
their cell contents. The injury produces a white or yellow spots or ‘stipplings’ that
are heaviest on the undersurface of the leaves. The leaves loose photosynthetic sur-
face as mite feeding continues. Both photosynthetic rate and leaf water status
decline with increasing levels of spider mite injury. As colonies grow and feeding
intensifies, entire leaves progress from greyish green to yellow, brown or coppery
and eventually drop off. Damage begins in the lower canopy and progresses
upwards. Heavily stippled upper leaves may exhibit deformations reminiscent of
herbicide injury. If the infestation is unchecked by disease, predators or miticides,
spider mites may kill the entire plant. Many times, progression of symptoms from
silvering, yellowing, browning, lower leaf loss and death of plants may be mistaken
from the drought symptoms. Drought triggers spider mite outbreaks in many crops
by upsetting balance of spider mite by natural enemies, weather and host quality
(ER3 and ER5).

Spider mites cause widespread changes in their host at cytological and histologi-
cal levels. Leaf surfaces are damaged including punctured and collapsed epidermal
cells, disrupted cuticles and stomata. Mesophyll tissues including vascular bundle
sheaths, palisade and spongy parenchyma are also affected.
6.2 Superorder Acariformes 99

The effective micro-environment of a small arthropod such as a spider mite


seems very different from representative nearby conditions. Temperature and
humidity also have been shown to affect the behaviour of spider mites in several
ways that may have important consequences for their population dynamics. These
include modifications of dispersal and web building; air movement may also influ-
ence these factors. Prolonged direct contact with free water and from rain or simu-
lated rains also has detrimental effects on spider mites.

Factors Promoting Mite Outbreaks Generally spider mite outbreaks were


uncommon in situations where productivity was low, since the mite populations
remained low due to natural regulation by predators, disease and poor nutrition from
low-quality host plants. Biological disruption from use of fertilizers and pesticides
and good nutrition speeds up the developmental rate and increased fecundity and
longevity of mites. Under such modern techniques of crop production, spider mite
outbreaks are quite common.
Rainfall may slow down the infestations, but it will not destroy all the popula-
tions. Spraying is the best option to control spider mites. Pyrethroid insecticides
perform poorly against mites and could even aggravate the situation by ‘flaring’
their populations. Eggs are difficult to kill with insecticides/miticides, so reinfesta-
tion is likely to occur in 7–10 days after treatment as a result of egg hatching. The
reinfestation is frequently heavy because natural enemies have been reduced or
eliminated. A second application may be necessary to kill newly hatched mites
before they mature and deposit more eggs. Details about pesticides being used
against these pests are given in a separate chapter of this text.

Family Tenuipalpidae (Fig. 6.25) Members of this family are commonly known
as false spider mites, flat mites and red tea mites. They are variously coloured; have
different shapes like ovoid, round and elongate with flat venter or flat dorsum; and
measure about 0.3–0.4 mm. Cheliceral bases adnate, fused mesally into a stylo-
phore (sometimes withdrawn into the body). Chelicerae with fixed digit reduced
and movable digit whip-like. Simple palpus which lacks a thumb–claw complex on
the penultimate segment, palpal segmentation often reduced. Adults with three or
four pairs of legs; tarsal claws usually with tenent hairs on both claws and empo-
dium. Genital papillae absent; males with an intromittent organ (Smiley and Gerson
1995).

Importance They are generally reddish looking, slow moving and normally feed
near the midrib or veins on the underside of leaves. Tenuipalpid species Raoiella
indica Hirst, Brevipalpus californicus Banks, B. obovatus Donnadieu, B. phoenicis
Geijskes and Tenuipalpus punicae Pritchard and Baker are important pests. They
are commonly encountered on different plants and are of high quarantine impor-
tance. They feed on stems, fruits, flowers and leaves (often on the lower surface).
Some species like Larvacarus transitans (Ewing) produces twig galls on jujube
trees in India, and Obdulia tamaris Pritchard and Baker on Tamarix trees in Mideast.
100 6 Important Acarine Families

Fig. 6.25 Tenuipalpidae: (a) Tenuipalpus. (b) Brevipalpus

Serious damage is caused to many crops (e.g. citrus, tea, grapes, fruit trees, orna-
mentals, orchids, pineapple and grasses). Several tenuipalpid species are confirmed
vectors of virus or virus-like diseases of plants. Brevipalpus phoenicis vectors cof-
fee ring spot and passion fruit green spot viruses in Brazil and Costa Rica. Some
Brevipalpus spp. have been identified as vectors of rhabdovirus that cause diseases
such as ‘citrus leprosis’. Mite-vectored diseases reduce the production as well as
life span of the plants.

Family Tuckerellidae (Fig. 6.26) Also known as ‘peacock mites’ because of the
elaborate ornamentations adorning the dorsal surface of their bodies. These are
brightly coloured having fan-like dorsal body setae and also possess five to seven
pairs of whip-like caudal setae. Empodium with tenent hairs; most dorsal setae
expanded leaf-like; posterior region with a row of flagelliform setae. Cheliceral
bases adnate, fused medially into a stylophore; chelicerae with fixed digit reduced
and movable digit whip-like. Tuckerella is the only genera of this family. They are
rarely found, but all reported species are phytophagous in nature and are found on
the stems of woody plants, usually in the cracks on small twigs. Some species are
associated with grasses. Tuckerella pavoniformis (Ewing) is reported commonly on
papaya.
6.2 Superorder Acariformes 101

Fig. 6.26 Tuckerellidae

Family Linotetranidae (Fig. 6.27) Members are commonly known as cryptic false
spider mites. They are rarely found in soil and on grasses in dryland habitats but not
as pests. They are usually colourless, slender and less than one-­third of a millimetre
in length. Resemble other elongate tetranychids. Linotetranidae have also been
reported from moss and rotting trees. Empodium of these mites is provided with
tenent hairs; prodorsum with four pairs of setae. Cheliceral bases adnate, fused
mesally into a stylophore (sometimes withdrawn into the body); chelicerae with
fixed digit reduced and movable digit whip-like. Chambered peritremes present at
the dorsolateral surface of the prodorsum. Eyes are absent and palps five segmented,
usually with thumb–claw process. Linotetranus is an important genus, but none of
the species are pests of plants.

6.2.1.1.2 Superfamily Bdelloidea

Family Bdellidae (Fig. 6.28) Gnathosoma is cone shaped; mouthparts are snout-
like possessing long palpi which terminate with strong setae and lack the palpal
thumb–claw complex, measuring 450–3500 μm in length. They are reddish in
colour and fast-moving mites found on plants and in soil preying on phytophagous
mites and small insects like Collembola. Bdella depressa Ewing and B. longicornis
Linnaeus are found to feed on spider mites; another species B. lapidaria Womersley
is reported as an efficient predator of lucerne flea, Sminthurus viridis. B. distincta
102 6 Important Acarine Families

Fig. 6.27 Linotetranidae

Fig. 6.28 Bdellidae


6.2 Superorder Acariformes 103

Fig. 6.29 Cunaxidae

Baker and Balock is reported feeding on eggs and crawlers of armoured scale
insects.

Family Cunaxidae (Fig. 6.29) They are small- to medium-sized (350–500 μm),
red- or brown-coloured and fairly fast-moving mites. The chelicerae are indepen-
dent and elongated; the movable digits are short and hooked, whereas the fixed
digits are reduced. The palps are raptorial, three to five segmented and often armed
with long spines on the internal margins. The stigmata are located at the bases of
chelicera without the peritremes. The prodorsum has two pairs of prominent
trichobothria. These are cosmopolitan and occur in soil, leaf litter, compost and
stored products. Members are general predators of small arthropods and nematodes.
Cunaxa capreolus Berlese and C. setirostris Hermann, Cunaxoides parvus Ewing
and C. olivieri Schruft are important predators (Meyer and Ryke 1959).

6.2.1.1.3 Superfamily Halacaroidea

Family Halacaridae (Fig. 6.30) Body rounded with enlarged palps; legs with long
setae; legs I and II anteriorly directed and legs III and IV posteriorly directed. They
are small-sized mites (220–800 μm in length). Halacaridae are the most diverse
family ecologically among all the Acari. Their biological diversity has enabled
them to invade habitats not only in salt water but also in freshwater, lakes, streams
and springs, as well as intertidal and cavernicolous habitats in fresh and saline water.
104 6 Important Acarine Families

Fig. 6.30 Halacaridae

There are phytophagous forms and carnivorous forms (predators, parasites and even
scavengers). Many halacarid species are found as both external and internal para-
sites on animals and other hosts; some species are reported from intertidal or shal-
low subtidal habitats and also reported from ocean depths of 400–1400 m.
Copidognathus, Acarothrix, and Peregrinacarus are predominantly inhabitants of
diluted brackish water and freshwater (Cook 1967; Prasad and Cook 1972; Proctor
2004).

6.2.1.1.4 Superfamily Eupodoidea

Family Eupodidae (Fig. 6.31) Anal opening terminal; intercoxal region with 15 or
more pairs of setae; femora IV normal or strongly swollen, used in jumping. They
are commonly encountered in soil, litter, low-growing vegetation, lichens or moss
in temperate, boreal and polar latitudes. They are also reported from habitats that
range in elevations from sea level to high mountains (>2200 m). Some occur in
coastal intertidal zones and others have been reported from caves and under stones.
Cocceupodes species have swollen femora IV and are capable of rapid backward
movements when threatened. Leg I of eupodids is typically elongate (e.g. Eupodes
longisetosus Strandtman), but those of Linopodes species are unusually long and
may be encountered in mushroom houses and cause economic losses.

Family Rhagidiidae (Fig. 6.32) Palptibia with three setae; tibia I with two or more
solenidia; prodorsum with one pair of bothridia and three pairs of setae. Rhagidiids
6.2 Superorder Acariformes 105

Fig. 6.31 Eupodidae

Fig. 6.32 Rhagidiidae


106 6 Important Acarine Families

Fig. 6.33 Penthalodidae

are unusual in having fields of modified recumbent dorsal solenidia on tarsi I–II that
are distinctive to be diagnostic at the generic level. These rhagidial organs are usu-
ally inserted in individual or compound grooves or troughs and typically are modi-
fied so that each solenidion takes the form of a pick hammer, with the offset stem
representing the shortened handle.
Members are soft-bodied and rapidly moving mites. They are generally opaque
white, but some are yellow, orange or red in colour. Larval and nymphal stages of
rhagidiids use spin from the podocephalic gland system to spin a web around them-
selves prior to moulting. Adults of Rhagidia longisensilla Shiba also spin silk for
catching prey for communal refuge and protection of the eggs. Reported from
alpine/high mountain regions, some are found at lower elevations. Rhagidiids com-
monly occur in the uppermost layers of soil and litter or within the soil column and
seem to prefer moist, dark cool niches. They are predaceous in nature. Rhagidia and
Coccorhagidia species are commonly reported.

Family Penthalodidae (Fig. 6.33) They are coloured and generally strongly scler-
otized; dorsum with ‘V’- or ‘Y’-shaped furrow and an anterior epirostrum project-
ing over the gnathosoma and ornamented with reticulate or punctate forms. Genus
Penthalodes is holarctic in distribution, while genus Stereotydeus is found world-
wide on moss and litter in temperate and subtropical habitats.

Family Penthaleidae (Fig. 6.34) Chelicerae separate; fixed digit regressed.


Movable digit sickle-like to prong-like – an adaptation for phytophagous behaviour.
Anal opening dorsal or dorsoterminal; femora IV similar to others. The stigmata are
6.2 Superorder Acariformes 107

Fig. 6.34 Penthaleidae

located between the bases of chelicerae. Red-legged earth mite, Halotydeus destruc-
tor (Tucker), is an important pest of many low-growing crops especially annual
broad-leaved plants and grasses. Blue oat mite, Penthaleus major (Duges), also
infests cereals, grasses, vegetables and ornamental plants in temperate and subtropi-
cal regions.

6.2.1.1.5 Superfamily Tydeoidea

Family Ereynetidae (Fig. 6.35) Presence of a peculiar sensory complex of


unknown function, the ‘Ereynetal organ’ on the tibia of leg I. Ereynetids seem to be
restricted to humid or wet habitats. They are rarely found in exposed foliar habitats
or arid regions. Some members are commonly found under bark and in dung, leaf
litter, moss and lichens, while some species are intimately associated with terrestrial
arthropods or gastropods. Riccardoella species are found on the surface and within
the mantle cavity of snails and slugs. Several Ereynetes spp. are associates of bark
beetles or phoretic associates on dung-inhabiting beetles and flies.

Family Tydeidae (Fig. 6.36) Weakly to heavily sclerotized striate or reticulate


mites. Tydeids are very small to small mites (150–500 μm) and have reduced fixed
cheliceral digits, fused cheliceral bases, and short movable cheliceral digits. Mites
are oval in shape and fast moving and can be found both singly and in congrega-
tions. The members live in a wide range of habitats and are predatory, fungivorous,
and scavenging species. Some species are considered pests or beneficial on agricul-
tural crops. Lorryia, Orthotydeus, Pronematus and Tydeus are important genera of
this family. Homeopronematus anconai (Baker) is reported as a good control of
eriophyid mite, Aculops lycopersici.
108 6 Important Acarine Families

Fig. 6.35 Ereynetidae

Fig. 6.36 Tydeidae

Family Iolinidae (Fig. 6.37) Adults with only one pair of genital papillae, or papil-
lae absent; moveable digit of chelicerae stylet-like or whip-like. Palptarsus usually
with three or four elongated setae; males typically with an aedeagus. Proctotydeus
spp. are found associated on the tympana of noctuid moths. Pronematus ubiquitous
6.2 Superorder Acariformes 109

Fig. 6.37 Iolinidae

(McGregor) feeds on honey dew, fungi and dead insects. Homeopronematus anconi
(Baker) is reported as a good control of eriophyid mite, Aculops lycopersici (Wolff.)

6.2.1.1.6 Superfamily Eriophyoidea


Eriophyoidea, also known as the Tetrapodili or four-legged mites, are worldwide in
distribution, occurring on a variety of primarily perennial hosts. All post-embryonic
instars lack third and fourth pair of legs (ER16). All the members are obligatory
phytophagous mites. They are very small to moderate in length (100–150 μm in
general, but range is 80–300 μm) (Jeppson et al. 1975).

Family Eriophyidae (Fig. 6.38) They are tiny, microscopic mites, yellowish to
pinkish white to purplish in colour. The rostrum is usually small relative to the body,
with the short form oral stylets. The distal recurved portion of the stylet is usually
much shorter than the stylet base. They are found on a wide range of plants; and
many species are major pests causing economic losses to crops, fruit trees and low-
ers quality of fruits and flowers. Some species are used as biological agents to con-
trol weeds and invasive plant species.

Family Diptilomiopidae (Fig. 6.39) Also known as big-beaked mites. Gnathosoma


is large compared to the body; chelicerae are abruptly curved and are bent down
near the base. The big-beaked plum mite, Diptacus gigantorhynchus (Nalepa), dam-
ages apple and some other Prunus species.

Family Phytoptidae (Fig. 6.40) The members are commonly known as gall mites
or rust mites. The cheliceral stylets are slightly evenly curved form of small to mod-
erate size like in Eriophyidae.
110 6 Important Acarine Families

Fig. 6.38 Eriophyidae

Fig. 6.39 Diptilomiopidae

Fig. 6.40 Phytoptidae

Body divisions, life cycle and damage by eriophyid mites


Body divisions: There are three main body divisions – the rostrum, the dorsal shield
or cephalothoracic shield and the abdomen.

Rostrum An important structure in the classification at the family level, it is of a


beak-like form and directed downwards. In the family Diptilomiopidae, the rostrum
is large in relation to the body and is apically attenuate. In the Phytoptidae and
Eriophyidae, on the other hand, the rostrum is usually small relative to the body.
6.2 Superorder Acariformes 111

Cephalothoracic Shield Also sometimes known as the propodosomal or dorsal


shield or simply shield, it is triangular or semicircular and bears a pattern of lines,
granules or broken lines which are distinctive for individual species. The pattern of
lines may vary slightly or differ in intensity in any given species, but it is an impor-
tant diagnostic feature. The cephalothoracic shield sometimes has a projection – the
anterior shield lobe, which is an important character at subfamily and genus level.
In a few instances, the lobe is thin- and flap-like. The lobes’ variation in shape and
prominence is of importance in species identification.

The Abdomen The principal part of the eriophyid body, usually tapers towards the
rear and is worm-like, is divided transversely into a number of superficial rings
which in bud and gall mites are regular dorsoventrally. Free-­living forms are less
worm-like and may have flattened abdomens with longitudinal ridges or grooves or
lateral structures. They may also have the body clearly divided dorsoventrally into
tergites and sternites. Some species have a row of a white, waxy secretion on the
dorsal surface of the abdomen or may cover themselves with flocculent wax. The
body rings are frequently studded with elongate oval or rounded structures, called
micro-tubercles, which may completely cover the body, or occur on the sternites
only. The form, number and arrangement of micro-tubercles are useful diagnostic
characters. The abdomen bears a limited number of setae. The relative length of the
setae is a valuable specific and generic character.

Genitalia The transverse genital opening is located anteriorly, just behind the hind
coxae. The male genital opening is ‘V’ shaped, but in females genital opening is
covered by a semicircular hinged flap, the cover flap, which may have longitudinal
markings or scoring. The degree and nature of the cover flap markings are of signifi-
cance in the generic and specific placement of some species.

Legs All eriophyid mites have only two pairs of legs. The forelegs and hindlegs are
essentially similar. Leg setation is an important character for identification. The
claws or ‘featherclaws’ which arise from the distal part of the tarsus are important
structures. The rock-like claw may be straight or curved and often terminates as a
knob. The length of the claws is of use for species identification. The featherclaw is
usually of simple type which has a central stem from which arise branches or rays.
The number of rays is an important species character. The second type is divided
featherclaw or having some modification. Featherclaw structure is important in
genus and species placement.

Life Cycle Two main types of the life cycle occur in eriophyid mites. The first,
sometimes called a simple life cycle, involves only one type of female. The female
lays eggs, there are two nymphal stages and finally the adults develop. The nymphs
112 6 Important Acarine Families

differ from the adults in being smaller, in lacking external genitalia and in the extent
of body micro-tuberculation. All known eriophyid mites are obligate plant parasites.
Males do not mate with females, but fertilization occurs from the contact of females
with sperm sacks laid down on the host by males.
The second type of life cycle sometimes called a complex life cycle involves two
types of females: the protogyne or primary form, which is associated with males
and structurally is very similar to them, and the secondary female or deutogyne,
which is not associated with males and is quite different in appearance to the proto-
gyne. Frequently protogynes occur on the leaves of the host plant, whereas deuto-
gynes usually appear in response to leaf hardening or the onset of winter and
hibernate in bark crevices or lateral buds, emerging in the following spring to lay
eggs on the new foliage. The eggs give rise to protogynes and males. Deutogynes
preserve the species during less favourable times of the year when food is not freely
available. Dispersal of these slow-moving mites is by wind, water, birds, insects or
humans. Commerce/trade of plants or their vegetative parts (in horticulture particu-
larly) is probably the major reason in introducing these mite species into distant
places previously un-infested areas. So any unknown plant abnormality should be
critically examined for the presence of these tiny eriophyid mites. Existence of deu-
togynes and protogynes of a species is helpful in identification at species and generic
level and is also helpful in pest management of these mites as deutogynes which
usually have different habits from the protogynes.

Types of Eriophyid Mites The eriophyid mites are often divided into several
groups.

• Russeting (Rust) Types. Feeding produces a brownish discolouration of the epi-


dermis. Heavy fine stippling introduces a blemishing or staining of leaf and fruit
surfaces. ‘Silvering and bronzing’ of mite-infested areas may also result.
• Gall Former Types. A number of certain eriophyid mites produce galls and other
plant abnormalities by injecting growth regulators into the host tissues. Modified
leaf provides shelter as well as food. There are many types of galls (bladder,
pouch, finger, nail head, etc.), but they all have an exit hole at the bottom (usually
undersurface of the leaf). Most galls are found on the leaves, but some are on
flowers, petioles, green stems and even roots.
• Erinea Forming Types. The feeding by certain eriophyid mites induces the pro-
duction of hairy patches, i.e. erineum (pl. erinea), on the surfaces of leaves and
petioles. Feeding within the thick hairs offers the mites both anchorage and pro-
tection. Some erinea on the undersurface are domed up and distort the upper leaf
surface.
6.2 Superorder Acariformes 113

Damage Caused by Eriophyid Mites (ER15)


• Leaf-Edge Rolling. In some cases, feeding by eriophyid feeding produces a roll-
ing of the leaf margin.
• Witches’ Broom. Some eriophyid mites cause the tip of branches to develop
broom-like bunches of twigs, commonly known as ‘witches’ broom’ which
result in shortening of internodal growth with resultant clustering of buds.
• Blistering and Tissue Breakdown. Eriophyids, with their small mouthparts, feed
on individual plant cells and do not usually deeply penetrate plant tissues. Mite
feeding results in loosening buds and raised blisters that dry out later in the sum-
mer leaving dead areas on the expanded leaf blades.
• Bud Destruction. Feeding by some bud-inhabiting mite species causes a blasting
of the buds. The feeding causes the interior embryonic parts of buds to swell
producing the symptom commonly known as ‘big bud’. The bud dies after the
mites leave the cavities. Partial damage results in distorted new growth that looks
like possible herbicide injury.
• Virus-Like Symptoms. The mite-injected growth regulators sometimes may cause
the entire physiology of the host plant to be disturbed (failure of fruit bud forma-
tion, loss of vigour, early death) or scattered symptoms (concentric ring blotch)
suggestive of a virus disease.
• Virus Transmission. Sometimes mites act as vectors of virus diseases and trans-
mit them by feeding. Some mosaic types of virus diseases are caused by mite-­
infected viruses.

6.2.1.1.7 Superfamily Anystoidea

Family Anystidae (Fig. 6.41) Also known as whirligig mites. They are bright red,
long legged, large (500–1500 μm) and soft bodied, possessing few dorsal setae and
no prodorsal sensilla. They possess a palpal thumb–claw complex whose tarsus is
longer than claw. Most of the members are predaceous in habit. Anystis baccarum
Linnaeus is reported quite effective against mites infesting soybeans. A. agilis
Banks is also reported effective on grapevines against spider mites and also reported
feeding on larvae and adults of citrus thrips, Scirtothrips citri.

6.2.1.1.8 Superfamily Hydryphantoidea

Family Hydryphantidae (Fig. 6.42) Adults with two-segmented chelicerae; palpi


chelate with dorsotibial portion of tibia bearing a short, thick seta extending beyond
base of tarsus. Idiosoma usually with varying degrees of plate or platelet develop-
ment dorsally. Deutonymphs and adults are sluggish and are adapted for walking or
crawling in aquatic vegetation and detritus. Hydrophantes tenuabilis Marshall is
reported to feed on dragonfly eggs.

Family Thermacaridae (Fig. 6.43) Dorsal and ventral shields present; coxal plates
I with two rows of large setae extending posteriorly from capitular bay; palp femur
with two long medial setae. Reported only from hot springs. Thermacarus is an
important genus.
114 6 Important Acarine Families

Fig. 6.41 Anystidae

Fig. 6.42 Hydryphantidae

6.2.1.1.9 Superfamily Eylaoidea

Family Eylaidae (Fig. 6.44) Prodorsal plate noticeably wider than long, con-
stricted near middle and bearing one pair of setae. Inhabits ponds and temporary
pools. Deutonymphs and adults feed on ostracods and cladoceran crustaceans and
are subelytral parasites of various aquatic Coleoptera and Hemiptera.
6.2 Superorder Acariformes 115

Fig. 6.43 Thermacaridae

Fig. 6.44 Eylaidae

Family Limnocharidae (Fig. 6.45) Prodorsal plate much longer than wide; geni-
tal acetabula scattered in ventral integument. Inhabits ponds and lakes. Limnocharis
is an important genera; adults feed on larval Chironomidae, and larvae parasitize
surface-dwelling aquatic Hemiptera (especially family Gerridae) and various
Odonata.
116 6 Important Acarine Families

Fig. 6.45 Limnocaridae

6.2.1.1.10 Superfamily Hydrachnoidea

Family Hydrachnidae (Fig. 6.46) Adult gnathosoma typically with long, curved
pointed rostrum. Palpi chelate with dorsodistal tibial claw extending well beyond
base of tissues. Legs with five movable segments, with basifemur and telofemur
fused and with leg tarsi lacking paired claws. Dorsal plate bearing eight pairs of
setae. Larval Hydrachna are parasites of various aquatic Hemiptera and Coleoptera
in pools, ponds and lakes.

6.2.1.1.11 Superfamily Hygrobatoidea

Family Limnesidae (Fig. 6.47) Two pairs of ‘urstigmata’ borne between coxal
plates I and II; bearing seven to eight setae. Limnesia spp. and some other genera of
this family are found in streams, ponds and lakes.

Family Hygrobatidae (Fig. 6.48) Coxal plates I–III on each side all fused. Found
among the dominant water mites in flowing-water habitats. Larvae are parasites of
Chironomidae. Hygrobates nigromaculatus Lebert has been reported as a limiting
factor in limiting populations of chironomid larvae in lakes (Cook 1967; Prasad and
Cook 1972).

Family Unioncolidae (Fig. 6.49) Dorsal and coxal plates reticulate, coxal plates
bearing pointed projections posteriorly. Larvae are parasites of Chironomidae or
Trichoptera. Free-living species of Unioncola are essentially planktonic; many oth-
ers have obligate relationships with molluscs.

Family Aturidae (Fig. 6.50) Coxal plate III with pointed or lobed projections
­posteriorly and bearing two pairs of setae. They are well-sclerotized and t­ypically
6.2 Superorder Acariformes 117

Fig. 6.46 Hydrachnidae:


(a) Dorsum. (b) Venter

Fig. 6.47 Limnesidae

dorsoventrally flattened mites. Adults of many genera are strongly dimorphic. This
is a species-rich family of water mites. Larvae are parasites of chironomids.

Family Pionidae (Fig. 6.51) Genital acetabula numerous (more than 10 pairs),
genus of leg IV either concave on one side and with numbers of peg-like setae or
with median surface of palptibia bearing a peg-like setae at distal end, and coxal
118 6 Important Acarine Families

Fig. 6.48 Hygrobatidae

Fig. 6.49 Unioncolidae

plates lacking well-developed projections associated with insertions of leg


IV. Members are found in springs, streams and standing water habitats. Piona con-
tains important species of water mites. Adults of many genera exhibit marked sex-
ual dimorphism.
6.2 Superorder Acariformes 119

Fig. 6.50 Aturidae

Fig. 6.51 Pionidae


120 6 Important Acarine Families

6.2.1.1.12 Superfamily Arrenurroidea

Family Neoacaridae (Fig. 6.52) Well-sclerotized dorsoventrally flattened mites;


dorsal and ventral shields present; ventral shield suture lines between coxal plates
III and IV extending posteromedially to genital field region and well separated from
each other medially. Genital acetabula in single rows on each side. Neoacarus is an
important genera; larvae are known as parasites of Chironomidae.

Family Arrenuridae (Fig. 6.53) Adults with dorsal and ventral shields present; palpi
truncate; genital acetabula numerous and lying on acetabular plates extending later-
ally from gonopore. In Arrenurus genus most importantly, its members are found in
virtually all types of freshwater habitats. Many species feed on ostracods. Larvae of
Arrenurus are parasites of various nematocerous Diptera or Odonata (Cook 1967).

6.2.1.1.13 Superfamily Erythraeoidea

Family Erythraeidae (Fig. 6.54) Members are large, reddish mites with two pairs
of prodorsal sensilla; many dorsal setae and a thumb–claw process with long straight
chelae. Larvae usually parasitize other arthropods, whereas the nymphs and adults
are predators of bugs and other small insects. Blaustium putmani Smiley is reported
to feed on eggs and mobile stages of European red mite, Panonychus ulmi (Koch).

Family Smaridiidae (Fig. 6.55) Gnathosoma is entirely retractable into idiosoma;


idiosomal setae are usually setiform. Deutonymphs and adults are m­ oderately large
(1000–2000 μm). Active predators of small arthropods are primarily found in grass-
lands and litter habitats. Larvae of Smaris prominens (Banks) are reported as para-

Fig. 6.52 Neoacaridae


6.2 Superorder Acariformes 121

Fig. 6.53 Arrenuridae

Fig. 6.54 Erythraeidae


122 6 Important Acarine Families

Fig. 6.55 Smaridiidae

sites of psocids; larvae of some other species are found on many other insects
especially scale insects.

6.2.1.1.14 Superfamily Trombiculoidea

Family Trombiculidae (Fig. 6.56) Commonly known as harvest mites or ‘chigger


mites’; larvae (170 μm) are commonly called ‘chiggers’, which are normally light
red and covered in hairs. There is a marked constriction in the front part of the body
in case of nymphal and adult stages. Chiggers attach to the host, pierce the skin and
inject enzymes into the bite wound that digest cellular contents. They do not burrow
into the skin or suck blood, but feed on skin causing itching.
They are commonly found on grasses and weeds and get transferred to their
hosts when they approach for grazing. They prefer damp areas in woodlands, grass-
lands, golf courses, parks and bushes along lakes and streams. Their feeding causes
intense ‘irritation’ or a wheal, usually with severe itching and dermatitis.
Leptotrombidium deliense Domrow is considered a dangerous pest and often may
carry tiny bacterium, Orientia tsutsugamushi, that causes scrub typhus or tsutsuga-
mushi. Trombicula alfreddugesi (Oudemans) and T. autumnalis (Shaw) are the most
prevalent species.

6.2.1.1.15 Superfamily Trombidioidea

Family Trombidiidae (Fig. 6.57) Idionotum with unpaired prodorsal plate;


palptibia usually lacking large spinose setae adjacent to claw and either lacking any
spinose setae that are not arranged in rows, when bearing large spinose setae
6.2 Superorder Acariformes 123

Fig. 6.56 Trombiculidae

Fig. 6.57 Trombidiidae

arranged in rows and then anterior edge of prodorsal plate concave. Measure 500–
4000 μm. Anal gnathosoma not retractable into idiosoma. They are active predators
of small invertebrates in habitats that range from s­ ub-­aquatic to deserts. Trombidium
spp. parasitizes many species of insects and arachnids. Adults are usually found in
moist litter, or in moss in and around bark of tree trunk.
124 6 Important Acarine Families

Fig. 6.58 Eutrombidiidae

Family Eutrombidiidae (Fig. 6.58) Idionotum bearing large plate posteriorly.


Deutonymphs and adults are large mites (500–4000 μm) and found in wide variety
of edaphic habitats worldwide. Postlarval instars of Eutrombidium locustarum
(Walsh) are predators of grasshopper eggs. Larvae of various Eutrombidium spp. are
parasites of various Orthoptera.

6.2.1.1.16 Superfamily Myobioidea

Family Myobiidae (Fig. 6.59) Chelicerae and subcapitulum fused into a gnathoso-
matic capsule that is retractable within the idiosoma; leg I modified for clasping
mammalian hosts. Ectoparasites in the fur of bats, rodents and various insectivores.
Myobia musculi (Schrank) attacks laboratory mice, and Radfordia ensifera (Poppe)
attacks laboratory rats and causes dermatitis in rodents.

6.2.1.1.17 Superfamily Pterygosomatoidea

Family Pterygosomatidae (Fig. 6.60) Chelicerae independently moveable, elon-


gate and with swollen bases and hook-like movable digit with one or more teeth;
empodia absent; peritremes emergent. Most species are reported from lizards and
tortoises. They are bright red mites found attached under scales or between toes,
feed on body fluids of their hosts and result in benign to severe pathological disor-
ders such as anaemia and intense skin irritation. Pimeliaphilus spp. are reported
parasitic on lizards and triatomine bugs.
6.2 Superorder Acariformes 125

Fig. 6.59 Myobiidae

Fig. 6.60 Pterygosomatidae


126 6 Important Acarine Families

Fig. 6.61 Caligonellidae

6.2.1.1.18 Superfamily Raphignathoidea

Family Caligonellidae (Fig. 6.61) Measures about 250–550 μm. Basal segments
of stylate chelicerae fused with each other in midline to form a conical stylophore,
which bears on its dorsal surface a pair of sinuous peritremes. Palp five segmented;
palptibial claw single but well developed. Body without striations and shields,
found under the bark, mostly associated with bark beetles and from grain storages.
They are free-living predatory mites.

Family Stigmaeidae (Fig. 6.62) Body short (200–500 μm in length), orange or red
and ovoid or round in shape. Thumb–claw process present; chelicerae with stylet-
like movable digit. Dorsal shields frequently present; ventral shields absent. Coxae
I and II distinctly separated from coxae III. Genital and anal openings contiguous.
Stigmaeid mites are commonly found on leaves of plants and feed on different
stages of pest mites. Agistemus and Zetzellia are important genera, whose members
have been reported feeding on eriophyid mites, spider mites, false spider mites and
even on some soft-bodied insects (Meyer 1969).

Family Camerobiidae (Fig. 6.63) Body length ranges from 250 to 300 μm. Leg
tibia about three times longer than respective tarsi; oval to subcircular mites with
very long legs. Dorsal idiosomal setae in marginal area not whip-­like. Idiosoma
nearly round in dorsoventral view. Gnathosoma often covered by prodorsum; che-
licerae fused to form a stumpy stylophore. Camerobia and Neophyllobius are
important genera of this family.
6.2 Superorder Acariformes 127

Fig. 6.62 Stigmaeidae

Fig. 6.63 Camerobiidae

Family Cryptognathidae (Fig. 6.64) Measures 300–400 μm in length. Fully


armoured mites with holodorsal shield produced anteriorly into a hyaline or orna-
mented collar into which the narrow elongate gnathosoma can be retracted, only the
bottom of chelicerae fused; dorsal and ventral idiosoma covered by a single shield.
Idiosoma oval in dorsoventral view, strongly sclerotized. Found in leaf litter, tree
bark, moss and lichens. Cryptognathous and Favognathous are important genera.
128 6 Important Acarine Families

Fig. 6.64 Cryptognathidae

Fig. 6.65 Raphignathidae

Family Raphignathidae (Fig. 6.65) They are small red mites (300–500 μm);
round in shape without any suture between propodosoma and hysterosoma.
Propodosoma without any sensory setae; peritremes situated between chelicerae
and prodorsum. Tarsus with two claws and a rayed pulvillus. Tarsi I and II with a
single sensory organ. No genital suckers. May be found on low-­growing herbs and
shrubs beneath tree bark, house-dust or in stored grains.
6.2 Superorder Acariformes 129

Family Eupalopsellidae (Fig. 6.66) Length varies from 250 to 400 μm. They have
four pairs of prodorsal setae, the empodia have one (rarely two) pair of tenent hairs
and palp tarsi are elongate. They are common in very dry regions. Saniosulus spp.,
Eupalopsis spp. and Eupalopsellus spp. are reported feeding on scale insects.

6.2.1.1.19 Superfamily Cheyletoidea

Family Cheyletidae (Fig. 6.67) Body oval to round (200–1600 μm in length);


palptibial claw present and well developed. Free-living predators, associated with
insects. In some studies, Cheyletus eruditus (Schrank) is considered effective in
controlling stored grain mites, Acarus siro Linnaeus. Some cheyletid mites are
found on plants feeding on crop pests, but some others are found in fur of
mammals.

Family Demodicidae (Fig. 6.68) Body elongated worm-like with numerous


annuli; legs strongly telescoped, highly reduced or modified into hooks. Measure
150–400 μm in length and are semi-transparent. Body is covered with scales for
anchoring itself to the hair follicles; possess pin-like mouthparts for eating skin
cells. Demodex folliculorum Simon is found commonly feeding in the eyelids,
cheeks and eye lashes of human beings. D. canis Leydig is found on domestic dogs
and cats and can occasionally infest humans.

Fig. 6.66 Eupalopsellidae


130 6 Important Acarine Families

Fig. 6.67 Cheyletidae

Fig. 6.68 Demodicidae:


(a) Dorsum. (b) Venter
6.2 Superorder Acariformes 131

Family Cloacaridae (Fig. 6.69) Well-sclerotized mites without opisthosomatic


striae; gnathosoma reduced to a single segmented palpi, each with a fang-like stylet
apically. Legs without distinct claws. They are found as internal parasites in the tis-
sues of reptiles and birds. Pneumocephalus bubonic Fain and Smiley is reported
infesting connective tissues of lungs and bronchi of owls.

Family Psorergatidae (Fig. 6.70) All legs ending in paired lateral claws and a pad-
like or bilobed empodia; femora I–IV each with a strong ventral spur. Members are
parasites on the skin of mammals. Psorergates, Psorergatoides and Psorobia are
important genera and their infestations cause dermatitis, mange and follicular
infections.

Family Harpirhynchidae (Fig. 6.71) Legs I–II with empodial tenent hairs and
with or without lateral claws, legs III–IV without lateral claws or leg IV absent;
femora I–IV without strong ventral spurs. Infest the skin/feathers of birds found
under scales of reptiles. Ophioptera is an important genera.

6.2.1.1.20 Superfamily Pygmephoroidea

Family Siteroptidae (Fig. 6.72) Small mites (200–300 μm); chelicerae and subca-
pitulum are fused to the gnathosomal capsule, which has a pair of small cheliceral
stylets and reduced palps. The prodorsal sclerite of the female does not cover the

Fig. 6.69 Cloacaridae


132 6 Important Acarine Families

Fig. 6.70 Psorergatidae: (a) Dorsum. (b) Venter

Fig. 6.71 Harpirhynchidae


6.2 Superorder Acariformes 133

Fig. 6.72 Siteroptidae: (a)


Dorsum. (b) Venter

entire propodosoma. The bases of legs II are twice as widely apart as the bases of
legs III–IV. Siteroptes spp. are pests of grasses and cereals and are responsible for
transmissions of pathogenic fungi. Siteroptes avenae is of economic importance
under greenhouse conditions. Pediculaster flechtmanni infests button mushrooms
and results in qualitative losses.

Family Pygmephoridae (Fig. 6.73) Gnathosoma of male longer than wide; tro-
chanter IV of female quadrangular. Prodorsum of females with stigmata round or
oval. Found associated with fungi, insects and animal nests. Pygmephorus sellnicki
Krczal has commonly been reported on cultivated mushrooms.

6.2.1.1.21 Superfamily Scutacaroidea

Family Scutacaridae (Fig. 6.74) Broad, tortoise-shaped bodies (120–370 μm in


length). Female with hood-like podonotal shield covering propodosoma and gna-
thosoma; margin of podonotal shield is striated. Scutacarus, Nasutiscutacarus and
Parascutacarus are important genera found commonly associated on halictid and
bumblebees, carabids and ants (Mahunka 1965).

Family Microdispidae (Fig. 6.75) They are small (80–315 μm) and possess greatly
enlarged and/or ornate dorsal setae. Idiosoma oval; distance between insertions of
legs II and III at least twice that between legs III and IV. Mostly free-living or asso-
ciated with insects and animal nests. Microdispus lambi (Krczal) is an important
134 6 Important Acarine Families

Fig. 6.73 Pygmephoridae

Fig. 6.74 Scutacaridae


6.2 Superorder Acariformes 135

Fig. 6.75 Microdispidae: (a) Dorsum. (b) Venter

species found in soil, forest litter, decaying vegetation, rotting wood, moss beds and
other similar habitats.

6.2.1.1.22 Superfamily Pyemotoidea

Family Pyemotidae (Fig. 6.76) Female coxisternal plates II–IV separated from
each other medially by soft cuticle that bears a separate triangular sternal plate.
Male legs IV differentiated from legs II and III in being somewhat stouter and end-
ing with a single sessile claw. The chelicerae are minute and needle-like; the perit-
remes are barely absent and are dorsal in position directed anteriorly on the shoulders
of the propodosoma. Pyemotids reported as ectoparasites of insects and other
arthropods. Pyemotes anobii Krczal paralyses larvae and pupae of bees. P. herfsi
(Oudemans) is reported feeding on gall-­making midges on oak leaves. P. tritici (La
Greze-Fossat and Montagne), straw itch mite, is associated with dermatitis in horses
and man and P. scolyti (Oudemans) with beetles.

Family Acarophenacidae (Fig. 6.77) Whitish-coloured adults measure about 140


μm in length; gnathosoma capsule, palps not distinct, chelicerae stylet-like. Body
covered by a series of sclerotized plates; prodorsum with anterior stigmatal
136 6 Important Acarine Families

Fig. 6.76 Pyemotidae: (a)


Dorsum. (b) Venter

Fig. 6.77 Acarophenacidae


6.2 Superorder Acariformes 137

o­ penings; sensillum absent. Legs with strong claw. Acarophenax lacunatus Cross
and Krantz is an egg parasite of stored grain pest, Rhyzopertha dominica, on wheat.

Caraboacaridae (Fig. 6.78) Female with gnathosomatic capsule large, clearly


wider than long; female leg I lacking claws, smaller than legs II–III. Leg IV with
tibia and tarsus at least partially fused and with reduced claw. Mites are associated
with carabid beetles. Caraboacarus is an important genera.

6.2.1.1.23 Superfamily Tarsonemoidea

Family Tarsonemidae (Fig. 6.79) Females with four pairs of legs; legs IV three
segmented, usually much more slender than legs II–III. Males always with four
pairs of legs; legs IV inserted ventrally; male genital capsule caudal in position.
Members of Steneotarsonemus, Polyphagotarsonemus, Phytonemus, Florida and
Tarsonemus (Lindquist 1969) are known to feed on higher plants, while most tarso-
nemid species feed on the mycelia of fungi and alga. Polyphagotarsonemus latus
(Koch) and S. pallidus Banks are pests of agriculture. Acarapis woodi (Rennie) is an
endoparasite of bees (ER17).

Family Podapolipidae (Fig. 6.80) Females typically with one to three pairs of
legs; leg IV if present is five segmented and more slender than legs II and III. Male
genital capsule caudal or dorsal in position. Several species belonging to genera
Locustacarus and Podapolipoides are parasitic on insects and sometimes prove
detrimental.

Fig. 6.78 Caraboacaridae


138 6 Important Acarine Families

Fig. 6.79 Tarsonemidae: (a) Dorsum. (b) Venter

Fig. 6.80 Podapolipidae


6.2 Superorder Acariformes 139

6.2.1.2 S
 uborder Oribatida (Including Cohort Astigmatina)
(Balogh and Balogh 1992)

6.2.1.2.1 Superfamily Histiostomatoidea

Family Histiostomatidae (=Anoetidae) (Fig. 6.81) Chelicerae laterally flattened;


fixed digit typically comb-like with numerous finely spaced teeth; movable digit short.
Palpi short, simple and often with a membranous complex. Males are more heavily
sclerotized than females. Genital papilla ring-like, usually large, not using from a
common furrow and not directly associated with the genital opening; body relatively
larger and not dorsoventrally flattened. Histiostoma spp. are typical inhabitants of
dungs of vertebrates, and deutonymphs often specialize for phoresy on staphylinid
beetles. Histiostoma murchiei Hughes and Jackson is reported as a predator of earth-
worm egg cocoons. Anoetus spp. live in mutualistic association with halictid bees.

6.2.1.2.2 Superfamily Canestrinioidea

Family Canestriniidae (Fig. 6.82) Idiosoma with cuticle smooth, striated, reticu-
late, scaly or sometimes well sclerotized. Gnathosoma normally developed; palp
and chelicerae occasionally elongated. Females with genital valves fused to body
anteriorly, forming V-shaped oviparous. Members are moderate to large in size and
are very weakly sclerotized. They are external associates or parasites commonly
associated with many beetle families Carabidae, Scarabaeidae, Lucanidae,
Passalidae, Cerambycidae and Chrysomalidae. Passalophagus georglei is reported
associated with passalid beetles.

Fig. 6.81 Histiostomatidae


140 6 Important Acarine Families

Fig. 6.82 Canestriniidae

6.2.1.2.3 Superfamily Hemisarcoptoidea

Family Hemisarcoptidae (Fig. 6.83) Adults with gnathosoma normally formed;


chelicerae chelate-dentate. Idiosoma with cuticle smooth, mammilate, microtri-
chous or occasionally strongly sclerotized female with oviparous in form of inverted
‘V’. Nanacarus minutus (Oudemans) is a common inhabitant of house-dust and
stored food products. Hemisarcoptid mites are common inhabitants of various arbo-
real and aquatic habitats. Hemisarcoptes spp. are obligate parasites of diaspidid
scale insects, and some species have been employed in biological control of scale
insects.

Family Chaetodactylidae (Fig. 6.84) Members have soft globose bodies with a
distinctly mammilate cuticle, lack sejugal furrow and have greatly enlarged empo-
dial claws that allow for movement on host pollen/nectar stores. Supracoxal gland
opening on an elongate, oval sclerotized region restricted to dorsal above leg I;
female pretarsus with paired elongate condylophores. Males with condylophores
fused along their length and emerging ventrobasally as a hook-like structure in the
base of the pretarsus. They are cosmopolitan in the distribution and are commonly
found in bee nests. Sennertia cerambycina Oudemans is found in nests of Xylocopa
and Ceratina bees. It is a kleptoparasite, destroying host eggs or larvae and feeding
on pollen and nectar stores.

Family Winterschmidtidae (=Saproglyphidae) (Fig. 6.85). Empodial claws pres-


ent; condylophorus usually fused into a V-shaped sclerite or a more elongate sclerite
6.2 Superorder Acariformes 141

Fig. 6.83 Hemisarcoptidae

Fig. 6.84 Chaetodactylidae


142 6 Important Acarine Families

Fig. 6.85 Winterschmidtiidae

in base of ambulacral stalk. They are found in a wide variety of habitats. Saproglyphus
spp. are fungivorous in decaying materials, bark crevices and woody fungi.
Winterschmidtia spp. are found as predators of the eggs of bark beetles. It is
Procalvolia zacheri and Acalvolia squamata (Oudemans) sometimes infest stored
food products.

Family Carpoglyphidae (Fig. 6.86) Prodorsal sclerite absent; coxal apodeme I


fused medially with coxal apodeme II closing coxal fields I in both sexes.
Condylophores elongate and separate in female, asymmetrical in male. Carpoglyphus
lactis (Linnaeus) is a cosmopolitan pest of stored materials with a high sugar con-
tent such as dried fruit, powdered milk, honey and many other stored food material.
Their deutonymphs have also been found as phoretic on insects of order Lepidoptera
and Coleoptera.

6.2.1.2.4 Superfamily Glycyphagoidea

Family Glycyphagidae (Fig. 6.87) Ventral subcapitulum with a prominent pattern


of external transverse and oblique ridges; empodial claws small or absent; condylo-
phores very thin or absent. Female with an epigyneal apodeme and often with an
external copulatory tube; males without paranal suckers or modified setae on tarsus
IV. Commonly found in nests of vertebrates, stored products, house-dust or occa-
sionally on plant foliage. Aeroglyphus robustus (Banks) is commonly found in
stored food products.
6.2 Superorder Acariformes 143

Fig. 6.86 Carpoglyphidae: (a) Dorsum. (b) Venter

Family Chortoglyphidae (Fig. 6.88) Body without microtrichia; male with


paranal suckers and often with sucker-like setae on tarsus IV. Chortoglyphids have
highly reduced coxal apodemes III–IV and the males are only glycyphagids to pos-
sess paranal suckers. Dorsal setae are mostly spine-like, b­ ifurcate apically and ante-
riorly directed. Commonly found in nests of rodents, stored products and house-dust.
Chortoglyphus arcuatus (Troupeau) is many times reported on stored foods.

6.2.1.2.5 Superfamily Acaroidea

Family Acaridae (Fig. 6.89) Prodorsal sclerotization in the form of a rectangular


field that may be incised posteriorly, or prodorsal sclerotization absent; dorsal setae
variable but not usually both elongate and heavily barbed. Cuticle smooth or rarely
with small rounded protuberances. Acarus siro and Tyrophagus putrescentiae
(Schrank) are pests of stored grains, cheese and other stored food products.
Rhizoglyphus echinopus (Fumouze and Robin) is reported to feed on bulbs, corms
and tubers.
144 6 Important Acarine Families

Fig. 6.87 Glycyphagidae

Fig. 6.88 Chortoglyphidae


6.2 Superorder Acariformes 145

Fig. 6.89 Acaridae

Family Lardoglyphidae (Fig. 6.90) Empodial claw bifurcate in females; males


with leg III enlarged, terminating in a large, straight empodial claw. Males with
paranal suckers and sucker-like setae on tarsus IV. Lardoglyphus konoi Sasa and
Asanuma and L. zacheri Oudemans are known as pests of dried meat and fish
products.

Family Suidasiidae (Fig. 6.91) Body distinctly longer than wide; dorsal setae fili-
form and unbarbed; tarsi with proral setae enlarged and claw-like. Deutonymphs
lack a pretarsus IV, but feeding stages possess a mammillate cuticle; sejugal furrow
is found. Suidasia and Neosuidasia species are known from stored food products
and in nests of bees, wasps and birds.

6.2.1.2.6 Superfamily Pterolichoidea

Family Freyanidae (Fig. 6.92) Body is generally broad; prodorsum with two ros-
tral setae or none; posterior legs ventrally inserted. Parasites of many orders of
aquatic birds. Freyana spp. are found associated with the wing and occasionally tail
feathers of aquatic birds.

6.2.1.2.7 Superfamily Analgoidea

Family Analgidae (Fig. 6.93) Ambulacral stalk inverted on the ventral tarsal apex
to form a distinctly convex surface; the ambulacral discs are relatively small and the
anterior legs bear apophyses and spines. Males of many genera exhibit polymor-
146 6 Important Acarine Families

Fig. 6.90 Lardoglyphidae

Fig. 6.91 Suidasiidae


6.2 Superorder Acariformes 147

Fig. 6.92 Freyanidae

Fig. 6.93 Analgidae


148 6 Important Acarine Families

phism, with the enlargement and the elongation of leg III. Large populations of
Megninia ginglymura (Megnin) mites result in weight loss and decline in egg pro-
duction of poultry birds.

Family Pteronyssidae (Fig. 6.94) Proximal enlargement of the ambulacral stalk as


a membrane along the ventral side of the tarsus; lateral sclerites of ambulacral disc
with distinct clear areas (lacunae). They are parasites on many bird species.
Pteroherpus is an important genus of feather mites.

Family Proctophyllodidae (Fig. 6.95) Most species are elongate and narrow, and
all lack propodosomatic setae and the solenidion on genu II. Females and many
males are bilobed posteriorly, with females possessing elongate projections from
the posterior lobes. Parasites of many hummingbird species. Allodectes and
Proctophyllodes are important genera containing important bird parasites.

Family Dermoglyphidae (Fig. 6.96) Elongated; both sexes with legs III and IV
ventrally inserted; anus terminal. They are parasitic in quills region of many bird
species. Female epigynial shield and male paranal suckers absent; and ambulacral
stalk convex ventrally.

Fig. 6.94 Pteronyssidae


6.2 Superorder Acariformes 149

Fig. 6.95 Proctophyllodidae

Fig. 6.96 Dermoglyphidae


150 6 Important Acarine Families

Family Epidermoptidae (Fig. 6.97) Earlier known as Knemidocoptidae . Body


sac-like or elongate; prodorsal sclerite with a strongly developed internal apodeme
on each side. All legs very short. Infests feather follicles or skin lesions of numerous
bird species. Epidermoptes and Myialges are containing representatives ectopara-
sitic on birds and mammals. Knemidocoptes spp. are burrowing mites found on
poultry and domestic birds.

Family Syringobiidae (Fig. 6.98) Dorsal hysterosoma with a single, undivided


shield; adults are comparatively well sclerotized. They are inhabitant of quills of
birds. Plutarchusia spp. cause considerable damage and weakening of the feathers
(Proctor 2003).

Family Pyroglyphidae (Fig. 6.99) Tarsus II with solenidion ω (omega) inserted in


basal half of the segment. Largely unsclerotized dorsally except for a propodoso-
matic sclerites (e.g. Dermatophagoides) to being virtually completely sclerotized
(e.g. Pyroglyphus). Coxal fields are well sclerotized. The female epigyneal apodeme
ranges from a large inverted ‘U’ to a smaller almost transverse form. Pretarsal
ambulacral disc may be relatively large and sclerotized internally. Infests many bird
species or mammal nests, house-dust or stored food products.

Family Cytoditidae (Fig. 6.100) Chelicerae with fixed digit absent or entire che-
licerae vestigial; leg and body setae highly reduced. Body rounded; strongly
regressed chelicerae; strong reduction of sclerotizations. Loss of all paraproctal
setae and strong reduction in size of the remaining setae. Members inhabit the respi-

Fig. 6.97 Epidermoptidae


6.2 Superorder Acariformes 151

Fig. 6.98 Syringobiidae

Fig. 6.99 Pyroglyphidae: (a) Dorsum. (b) Venter


152 6 Important Acarine Families

Fig. 6.100 Cytoditidae

ratory tract of birds. Cytodites nudus (Vizioli) inhabit the lungs, bronchi and air sacs
of poultry birds.

6.2.1.2.8 Superfamily Sarcoptoidea

Family Listrophoridae (Fig. 6.101) Commonly known as fur mite family.


Presence of a projecting tegmen that extends over the gnathosoma to contact the
host air. Propodosoma is sclerotized, and hysterosoma may also bear sclerites.
Parasites of many mammals. Listrophoroides cucullatus (Trouessart) is a wide-
spread fur mite of rats in tropical regions.

Family Chirodiscidae (Fig. 6.102) Strongly modified legs I–II terminate in flat,
plate-like attachment organs clasp host hairs; pretarsal elements absent from legs
I–II. They are parasites of mammals and rodents. Chirodiscoides caviae Hirst is a
common fur mite of laboratory guinea pig.

Family Psoroptidae (Fig. 6.103) These mites possess long legs with all the legs
projecting beyond the body margin. Anterior apodemes of coxal field not fused to
form a sternum; body rounded; pointed mouthparts; female opisthosoma rarely
bilobed; males with or without bilobed opisthosoma. They are mostly skin parasites
and sometimes also infest the inside of ears of many mammals. Psoroptes spp. and
Otodectes cynotis (Hering) feed on the skin or ears of sheep, livestock, horses and
domestic rabbits.
6.2 Superorder Acariformes 153

Fig. 6.101 Listrophoridae

Fig. 6.102 Chirodiscidae


154 6 Important Acarine Families

Fig. 6.103 Psoroptidae

Family Sarcoptidae (Fig. 6.104) Also known as burrowing mites; short legs which
rarely project the body region. Females are typically globose, but some are elon-
gated dorsoventrally. Parasites of many mammalian hosts. Sarcoptes scabiei
(Linnaeus) is a parasitic mite of humans that form subdermal burrows parallel to the
skin surface in susceptible individuals.

Family Pneumocoptidae (Fig. 6.105) Rounded body with strongly abbreviated


opisthosoma; leg IV situated near the posterior end of the body and a well-devel-
oped sclerite that covers most of the dorsal hysterosoma. Pneumocoptid mites infest
the lungs of rodents. Pneumocoptes penrosei (Weidman) and P. banksi (Baker)
parasitize ground squirrels and dogs.

Family Gastronyssidae (Fig. 6.106) Dorsal hysterosoma without sclerites; tarsus


is very short and is equal in length or shorter than tibia. They are endoparasites of
nasal passages of bats.

Family Myocoptidae (Fig. 6.107) Legs III–IV of female and III of male strongly
modified for clasping hairs; rarely leg IV of female reduced. They have well-devel-
oped apophyses in leg segments. Body usually dorsoventrally flattened and grasp
one hair in legs III–IV on each side. Myocoptes musculinus Koch sometimes appears
in serious proportions in laboratory mouse colonies.
6.2 Superorder Acariformes 155

Fig. 6.104 Sarcoptidae

Fig. 6.105 Pneumocoptidae


Fig. 6.106 Gastronyssidae

Fig. 6.107 Myocoptidae


Further Reading 157

6.3 Conclusions

The Acari are quite small in size (80 μm to 13 mm) having varied modes of life.
More than 55,000 species of different acarine species are known (5 % of king-
dom Animalia) distributed in 540 families among 124 superfamilies. Subclass
Acari is divided into two superorders, namely, Parasitiformes and Acariformes.
There are over 40,000 species found under superorder Acariformes, while more
than 15,000 species have been reported under superorder Parasitiformes. In
superorder Parasitiformes, representatives belonging to orders Holothyrida and
Opilioacarida are comparatively less diversified and have lesser economic
­importance. However, representatives belonging to order Ixodida to which ticks
belong are of great economic importance and have been studied in greater details
because of their more concern for mankind and veterinary animals. More than
896 species of ticks (including both hard and soft ticks) are described. Order
Mesostigmata includes many species of great economic importance under fami-
lies Phytoseiidae, Dermanyssidae, Varroidae, Parasitidae, Digmasellidae,
Eupodidae and Diarthrophallidae.
Superorder Acariformes is more diverse of the two superorders and includes over
40,000 described species in 351 families. Acariformes is divided into two orders,
namely, Trombidiformes and Sarcoptiformes. Order Trombidiformes is further
divided into two suborders, namely, Prostigmata (including almost all phytopha-
gous mites) and Sphaerolichida. Order Sarcoptiformes earlier known as Acaridida
and Oribatida is further divided into suborders Endeostigmata and Oribatida.
Endeostigmatans are soil inhabiting, but their importance is not well known.
However, suborder Oribatida has too much diversity of mites and includes impor-
tant representatives under superfamilies Histiostomatoidea, Hemisarcoptoidea,
Glycyphagoidea, Acaroidea and Sarcoptoidea.

Further Reading
Baker EW, Tuttle DM (1970) The false spider mites of Mexico (Acari Tenuipalpidae). USDA Tech.
Bull No. 1706. USDA, Washington, DC
Baker EW, Tuttle DM (1994) A guide to the spider mites (Tetranychidae) of the United States.
Indira Publishing House, West Bloomfield
Balogh J, Balogh P (1992) The oribatid mites genera of the world, vol 1. National Museum Press,
Budapest
Bolland HR, Gutierrez J, Flechtmann CHW (1998) World catalogue of spider mite family (Acari :
Tetranychidae). Koninklijke Brill, Leiden
Cook DR (1967) Water mites from India. Mem Amer Entomol Inst 9. Gainesvilla, FL
Helle W, Sabelis MW (eds) (1985) Spider mites: their biology, natural enemies and control. World
Crop Pests, vol 1B. Elsevier, Amsterdam
Holtzer TD, Norman JM, Perring TM, Berry JS, Heintz JC (1988) Effects of microenvironment on
the dynamics of spider mite populations. Exp Appl Acarol 4:247–64
Jeppson LR, Keiffer HH, Baker EW (1975) Mites injurious to economic plants. University of
California Press, 529 pp
158 6 Important Acarine Families

Krantz GW, Walter DE (2009) A manual of acarology 3rd edn, pp 97–103. Texas Tech University
Press, 807pp
Lindquist EE (1969) Review of holarctic tarsonemid mites (Acarina: Prostigmata) parasitizing
eggs of pine bark beetles. Mem Entomol Soc Can 60. Ontario Entomol Soc Canada
Mahunka S (1965) Identification key for the species of the family Scutacaridae (Acari:
Tarsonemini). Acta Zool Acad Sci Hung 11:353–401
Meyer MKP (1969) Some stigmaeid mites from South Africa (Acari: Trombidiformes). Acarologia
11:227–271
Meyer MKP, Ryke PK (1959) Cunaxoidea (Acarina: Prostigmata) occurring on plants in South
Africa. Ann Mag Nat Hist Ser 132:369–384
Prasad V, Cook DR (1972) The taxonomy of water mite larvae. American Entomological Institute,
Gainesville
Proctor HC (2003) Feather mites (Acari: Astigmata): ecology, behaviour and evolution. Annu Rev
Entomol 48:185–209
Proctor HC (2004) Aquatic mites: from genes to communities. Kluwer Academic Publishers,
Dordrecht/London
Smiley RL, Gerson U (1995) A review of the Tenuipalpidae (Acari: Prostigmata) of Australia with
description of two new genera and four new species. Int J Acarol 21:33–45
Sonenshine DE (1993) Biology of ticks, vol 2. Oxford University Press, New York
Sonenshine DE, Roe BM (2014) Biology of ticks, 2nd edn. Oxford University Press, London
Sonenshine DE, Lane RS, Nicholson L (2002) Ticks (Ixodida). In: Mullen GR, Durden LA (eds)
Medical and veterinary entomology. Academic, San Diego, pp 517–558
Walter DE, Proctor HC (2013) Mites: ecology, evolution and behaviour. Springer, Dordrecht/
Heidelberg/New York

Electronic References (ER)

1. https://www.youtube.com/watch?v=NOwnSW4v-Qs. Introduction to Acariform Morphology


2. http://itp.lucidcentral.org/id/mites/invasive_mite/Invasive_Mite_Identification/key/Whole_
site/Acari_Mites.htm. Mites or Acari
3. https://www.youtube.com/watch?v=-yolQljY_6I. Destroy Spider Mites On Marijuana
Cannabis
4. https://www.youtube.com/watch?v=3JNprSWv96w. Introduction to the Acari
5. https://www.youtube.com/watch?v=2rz36ufAfEQ. Life cycle of spider mites
6. https://www.youtube.com/watch?v=kAQNoEp8P0o. Ticks up close
7. https://www.youtube.com/watch?v=zZigF4_t1Zk. Family Tetranychidae
8. http://idtools.org/id/mites/flatmites/index.php. Flat Mites of the World
9. http://eol.org/pages/2586535/names?all=1. Acari: About Higher Groups & Families
10. http://www.cdc.gov/nceh/ehs/Docs/Pictorial_Keys/Acarina.pdf. Acarina: Illustrated Key to
some common adult female mites and adult ticks
11. http://www.cdc.gov/nceh/ehs/Docs/Pictorial_Keys/Ticks.pdf. Ticks: Key to Genera of Ticks in
United States
12. https://www.youtube.com/watch?v=eATpttq6DfA&index=198&list=PLlRvaTnNBxia9NlhGf
dI5-zgytIv75qbG. The Tick Key
13. https://www.youtube.com/watch?v=QM0kBwSU3hc&index=145&list=PLlRvaTnNBxia9Nl
hGfdI5-­zgytIv75qbG. Tick borne diseases of cattle
14. http://www.nappo.org/files/3714/3782/0943/DP_03_Tetranychidae-e.pdf. DP 03:
Morphological Identification of Spider Mites (Tetranychidae) Affecting Imported Fruits
15. https://www.unce.unr.edu/publications/files/nr/2004/FS0447.pdf. Ornamental Plant Damage
By Eriophyid Mites
Further Reading 159

16. https://www.youtube.com/watch?v=ecTauvwdeZg#t=1116.549386. Eriophyoidea


17. https://www.youtube.com/watch?v=UjtdMivX7x0&ebc=ANyPxKqTbw8oA1uKdtQf56PjXe
07YSkF2kai9Rz_ECNi0nrgQpn78-­B82EiTNeDOWYVANk3QbO9MIi8FmfcFIh5XiDMMn
0dUfA. Family Tarsonemidae
18. http://itp.lucidcentral.org/id/mites/invasive_mite/Invasive_Mite_Identification/key/0_
Glossary/Mite_Glossary.htm#Vertical-scapular system.

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