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BIOLOGICAL CONTROL OF AND BY ACARINA
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Overview Phytophagous
mites appear as pests in an array of agroecosystems, but have not been
extensively discussed as a separate group for biological control. In most
cases predatory mites are the key natural enemies of phytophagous mites.
Gerson et al. (1990) were perhaps the first to elaborate on the Acari as a
separate biological control category for armored scale insects. The following
discussion relies extensively on their report: Acarina
for Biological Control of Phytophagous Mites Dermacentor variabilis
(Say), American dog tick.--
This species is widely distributed in the U.S. east of the Rocky Mountains,
but is also found in California, Mexico and Canada (McMurtry 1977b). It
causes irritation to dogs and sometimes to livestock. Its greatest importance
is as a vector of Rocky Mountain spotted fever in the Central and Eastern
U.S., and is occasionally known to vector tularemia. The life cycle may vary
from 1-3 yrs. There is little activity during winter or in the warmest part
of summer. Adults are most active in the spring and may live more than 2 yrs
without food. This is the only stage known to infest humans, dogs and
domestic animals. Small mammals, especially mice and rabbits, are considered
to be the main hosts. Mating occurs on the host. After becoming engorged,
they drop from the host, and the females deposit their eggs in protected
places in masses of 4,00- 6,500 eggs after which the females die. Eggs hatch into six-legged larvae, which
attach to a passing host. After feeding for several days, they become
engorged, drop to the ground and molt to the nymphal stage. When the nymph is
ready to feed, it similarly seeks a host on which to attach. When the nymph
has become engorged, it also drops to the ground where it molts to the adult
stage. Both larvae and nymphs were observed to live over a year if food was
not available (Smith, Cole & Gouck 1946). Natural Enemies Sought.--In the U.S. a culture of the encyrtid parasitoid Hunterellus
hookeri Howard (formerly
Ixodiphagus caucutei du Buysson) was
introduced from France where it was propagated and released on Naushon
Island, Mass (Larrouse, King & Wolbach 1928). Small numbers of nymphs of D. variabilis parasitized by the French strain of H. hookeri were released on Capers Island, SC. in 1931 (Bishopp
1934). A larger effort was made on Martha's Vineyard Island, Mass, where an
estimated 90,000 females of H.
hookeri were released in two
locations on the island during 1937-39. The strain of parasitoid used
originated in Texas (Smith & Cole 1943). In the season following the releases of H. hookeri on Naushon Island, immature parasitoids were found
in a single nymph of the American dog tick and a single nymph of another tick
species (Larrouse, King & Wolbach 1928). Subsequent surveys were made in
1940 by Cobb (1942) and in 1941 by Smith & Cole (1943). In both a few H. hookeri were found, but none was recovered from the
American dog tick. Both this species and Ixodes
scapularis Say were still
observed in abundance; therefore, there was no evidence that any success was
achieved on the island (McMurtry 1977b). Bishopp (1934) reported recovery of the parasitoid from a single
nymph of D. variabilis on Capers Island two
yrs after release were made. In an assessment of results of releases of H. hookeri in Martha's Vineyard in 1937-39, Smith & Cole
(1943) recovered no parasitoids from ticks in the release areas and observed
no reduction in tick abundance that could be attributed to the parasitoid. A
later report by Smith, Cole & Gouck (1946) also indicates that the attempt
was unsuccessful. Natural Enemy Biology.--Hunterellus
hookeri is an internal
parasitoid of wide distribution, having been recorded not only from North
America but from Europe, Africa and South America (McMurtry 1977b). It was
reared from several species of Dermacentor,
Ixodes, Haemaphysalis, Thripecephalus
and Hyalomma. The biology of
this parasitoid was studied by Wood (1911), Cooley (1928), Cooley & Kohls
(1933) and Smith & Cole (1943) summarized by Cole (1965). The parasitoid oviposits in the body cavity
of fed larvae and fed or unfed nymphs of the ticks (McMurtry 1977b).
Oviposition may occur when the ticks are attached to the host animals.
Apparently the parasitoids do not develop in the larvae or the unfed nymph,
development proceeding after the nymph has become engorged. Overwintering may
thus occur in the unfed nymph, with the parasitoids emerging the following
spring after the nymph ticks have engorged with blood. The nymphs show no
signs of parasitism until sometime after feeding on the host animals is
completed. The period of development appears to be rather long. Cooley (1928)
found that at 22°C. the average time from dropping of engorged nymphs from
the host animal to emergence of adult parasitoids was 45 days. A number of eggs is laid in a single host,
and it was observed that more than one parasitoid may lay eggs in the same
host. The parasitoid larvae seem to consume all of the contents of the body
cavity of the host for successful transformation to the adult stage. Therefore,
the size of the adult is inversely proportional to the number in a single
host. An average of ca. 20 parasitoids emerges from a single nymph of Dermacentor andersoni Stiles or D. variabilis, and the highest number observed by Cooley
(1928) was 73. Dermacentor andersoni Stiles, Rocky Mountain Wood Tick.--
This tick is a vector of Rocky Mountain spotted fever, a rickettsial disease
that can be fatal to humans, but is primarily a disease of wild animals. It
can also harbor tularemia, another disease primarily of wild animals but also
infectious to humans. This tick is also responsible for tick paralysis, which
affects the motor nerves starting in the legs and gradually spreading to the
rest of the body (McMurtry 1977b). It results usually if the tick feeds at
the back of the neck or the base of the skull, and removal of the tick
usually results in recovery. The species occurs in the western U.S.,
primarily in the Rocky Mountains and also in Canada. Spotted fever occurs in
other areas also, but its chief vector there is the American dog tick, D. variabilis. Eggs of D.
andersoni are deposited on
the ground. They hatch in springtime or early summer into six-legged larvae
and climb onto grass or other vegetation where they wait attachment to
passing animals, usually small rodents (McMurtry 1977b). When fully fed in a
few days, the larvae drop to the ground to molt to the nymphal stage, which
usually does not feed until the following spring, when they attach to small
animals, become engorged and drop to the ground to transform to the adult
stage. Although some adults may attach to hosts the same season, they
seemingly pass the rest of the summer and winter in hiding and find a host
the following spring. Mating takes place on the host, and when fully fed the
female drops to the ground to deposit her eggs. Only the adult stage is known
to attack humans and large animals (Cooley 1932). Natural Enemies Sought.--In the U.S. a culture of the encyrtid parasitoid H. hookeri Howard, originating in France was started in Montana
for colonization against the Rocky Mountain wood tick (Cooley 1928, Cooley
& Kohls 1933). More than 4 million parasitoids were liberated during
1927-32, mostly in Montana but also in Colorado, Idaho and Oregon. Various
methods were used, including release of adult parasitoids, scattering
parasitized nymphs in grass and low vegetation, and liberating squirrels
which had been infested with parasitized nymphs. The method of mass rearing
the parasitoid on D. andersoni was described by
Morton (1928). Only one instance of recovery occurred in
1929, when a few parasitoids emerged from D.
andersoni nymphs taken from
squirrels captured in the Bitter Root Valley of Montana (Cooley & Kohls
1933). Cole (1965) cited from a personal communication from G. M. Kohls in 1963
that no reduction in the tick population was observed and no evidence had
been obtained that the parasitoids were established in nature (McMurtry
1977b). Other Ixodidae.--Alfeev
(1940) reported on an experiment in which it was attempted to control Isodes ricinus (L.) and I.
persulcatus Schulze in a
250-acre pasture in the province of Leningrad, USSR. H. hookeri,
obtained from Montana in 1935 was propagated and 2,600 adult parasitoids and 38,000
parasitized ticks were liberated. No recoveries were noted (McMurtry 1977b). Tetranychus urticae Koch, Two-Spotted Spider Mite.--McMurtry (1977) noted that there is a long list of synonyms
for this mite, the more common in the early literature being Tetranychus bimaculatus Harvey, T. telarius (L.) and T.
multisetus McG. The mite is
worldwide in distribution and has an extremely wide range of host plants
including fruit trees, ornamentals, vegetables and forage crops. The mites
increase to high populations causing stunting, drying of leaves and even
defoliation. This mite usually overwinters as orange-red diapausing adult
females, which do not feed or lay eggs until the following spring.
Fertilization apparently takes place in the fall and at lower latitudes
reproduction may occur throughout the winter (McMurtry 1977b). The species is typical of most Tetranychidae
in having egg, larva, protonymph, and deutonymph stages before becoming
adult. it is arrhenotokous (virgin females produce male progeny, while mated
females produce both sexes). A generation from egg to egg may be completed in
9 days. The rate of egg production at warm temperatures has been observed to
be five or more per female per day, and the total number of eggs may exceed 100.
This mite has not been observed to suspend itself on silken threads by which
it can be transported by air currents, such as occurs with some other
tetranychids. But, they can be dispersed by wind, although a higher velocity
may be required (Baker & Pritchard 1953, Boudreaux 1963, Boyle 1957,
Cagle 1949, Fleschner et al. 1956, Watson 1964). Natural Enemies Sought.--The phytoseiid predator Phytoseiulus persimilis Athias-Henriot
was introduced into Germany from Chile by Dosse (1958), who noted its potential
for controlling T. urticae in glasshouses (Dosse
1959). From Germany it was dent to other countries of Europe (Bravenboer
& Dosse 1962, Hussey & Parr 1965) and to Canada (Chant 1961) from
where it was sent to the U.S. (Smith, Henneberry & Boswell 1963, Oatman
1965). Due to most work with P. persimilis
being in glasshouses or on annual crops outdoors, almost all results have
been based on periodic releases rather than permanent establishment. In
Germany Dosse (1959) showed that P.
persimilis could increase
rapidly and decimate populations of T.
urticae in the glasshouse.
The possibilities of using the predator were further studied by Langenscheidt
(1966). In the Netherlands Bravenboer & Dosse (1962) reported that
releases of P. persimilis on cucumbers in the
glasshouse gave control of T.
urticae that was comparable
to 3-5 applications of insecticides or acaricides, but after the prey was
eliminated the predator also died out. Bravenboer (1969) indicated that there
are possibilities of practical application of this method for cucumbers, but
that much information is still required before the practice can be
recommended. But the use of Phytoseiulus
appeared to have little possibility in flower growing because of the low
damage tolerance to plants. In Great Britain considerable progress was made
at the Glasshouse Crops Research Institute in Littlehampton, Sussex on the
use of P. persimilis for commercial
control of T. urticae on cucumbers in
glasshouses. Experiments showed that when the predator was introduced at low
population densities of the prey, the latter could be eliminated before leaf
injury became serious (Hussey & Parr 1965). Larger tests showed that
releases of P. persimilis over ca. 3 acres of
glasshouse space resulted in successful control (Gould, Hussey & Parr
1968). One advantage was the generally severe leaf injury that cucumbers can
tolerate without yield loss. Several methods of establishing a uniform
pattern of control were tried, such as general inoculation with spider mites
as well as predators and a banker
method where infestations of the pest mites would be established on
one plant in every glasshouse so that sufficient mites could develop to
produce adequate numbers of predators for maintaining control over the entire
glasshouse. In the U.S. Smith, Henneberry & Boswell (1963) observed that P. persimilis showed promising possibilities in the control
of T. urticae on glasshouse ornamentals. However, the use of
predators alone may be deterred by a demand for flowers free from pests or
imperfections, and the possibility may be greater for an integrated control
program (McMurtry 1977b). Commonly used pesticides were very toxic to the
predators, but some were only slightly or nontoxic. Studies on annually
planted strawberries grown outdoors in southern California showed promising
results with releases of predators at the rate of ca. 300,000 per acre before
the spider mite population exceeded one per leaf (Oatman 1965, Oatman &
McMurtry 1966, Oatman et al. 1967). In central California this predator has
been observed to survive the winter but permanent establishment was uncertain
(McMurtry 1977b). The life history of P. persimilis
typically has an egg stage, a larval stage with three pairs of legs followed
by the protonymph and deutonymph, each having four pairs of legs, and the
adult. The period of development from egg laying to adult can be as quick as
4-5 days, which is more rapid than the rate of development of the prey, T. urticae, and also more rapid than most other species of
Phytoseiidae (Bravenboer & Dosse 1962, Dosse 1958, 1959). The rate of
oviposition may average as high as four eggs per female per day (Bravenboer
& Dosse 1962, Dosse 1958, 1958; Laing 1968, McClanahan 1968, McMurtry
1977b). Although any stage of the prey may re readily consumed, Chant (1961,
1963) found that the adult predators prefer adult or nearly mature spider
mites and that since the predator feeds directly on the reproductive units of
the prey population, it should be better able to suppress the prey than one
which feeds primarily on eggs and early immature stages. The functional response by P. persimilis to increasing prey density studied by Mori
& Chant (1966a) showed a domed curve of prey consumption with increasing
prey density in a relatively simple experimental arrangement. It seemed that
when prey were numerous there was a disturbance effect which reduced the
predator's rate of consumption. The high mobility of P. persimilis
appears to be an important factor in its effectiveness (Chant 1961). It
quickly moves down the rows in strawberry plots, and is also able to migrate
around barriers and over bare ground to invade control plots (Oatman 1965,
Oatman & McMurtry 1966, Oatman et al. 1967). Combined with its high
mobility is a marked ability to remain on and lay eggs only on infested
leaves (Chant 1961, Oatman & McMurtry 1966). This predator is seemingly
very dependent on spider mites for food, and therefore there is usually a
marked response to a change in density of the host. Nutritive substances such
as sucrose, honey, pollen and fish meal had no effect on longevity or
reproduction (Laing 1968, Mori & Chant 1966b). The optimum temperature
for reproduction was ca. 25-30°C (Bravenboer & Dosse 1962, Dosse 1958),
but reproduction can occur at much lower temperatures (Böhm 1966, McClanahan
1968). Mori & Chant (1966a, 1966b) studied behavior in relation to
humidity and found that activity of both predator and prey increased at low
RH, whereas the prey avoided high RH but the predator did not. Prey consumption
was highest at low RH. Panonychus citri (McGregor), Citrus red mite.--This mite has also been noted as Tetranychus mytilaspidis
Banks, T. citri McGregor, Paratetranychus citri, and Metatetranychus citri
(McMurtry 1977b). Reports occur from North and South America, China, India,
Japan, South Africa and Russia, but presumably are native to the Orient where
citrus originated. It is considered the most important pest of citrus in
California (McMurtry 1977b), but attacks other plants as well. P. citri feeds on leaves and fruit, causing a bronzing or
silvering of the surface. High infestations can cause defoliation, which is
enhanced under hot, dry conditions (McMurtry 1977b). One generation may be
completed in 3 weeks during warm weather, and 12-15 generations may occur per
year. An average of 96 eggs per female and an average longevity of 23 days at
24°C was reported in southern California, but other researchers reported a
maximum of only 50 eggs. The life span and period of oviposition are
considerably longer during cool months. There are commonly two peaks of
abundance in spring or early summer and again in the autumn or early winter.
During these periods the temperature may be most favorable, but the age of the
foliage may also be an important factor (McMurtry 1977b). High populations
can occur in some places at virtually any time of the year, however. All
developmental stages can be found in midwinter in California and Florida,
although in the colder areas of Japan it was reported that the winter is
passed in an egg diapause. An important means of dispersal is air drift;
adult females spin silken threads and are carried by air currents. This
action seems induced when the foliage becomes unfavorable through excess
feeding or other causes (Boyce 1936, Ebeling 1959, Ehara 1964, English &
Turnipseed 1941, Fleschner 1953, Fleschner et al. 1956, Fukuda & Shinkaji
1954, Henderson & Holloway 1942, Jeppson et al. 1957, Muma 1961a, Munger
1963, Quayle 1912). Fleschner (158) reported other factors affecting the
abundance of citrus red mite, such as predation, pesticides, water, soil,
direct and indirect effects of climate and host plant genetics. Natural Enemies Sought.--In the U.S. importation of predatory mites into southern
California began in 1953 which resulted in the importation of several species
of Stethorus from the Middle
and Far East and Central America, and releases made in orchards infested with
citrus red mite (McMurtry 1977b). Although one species, S. gilvifrons
(Muls.) was recovered in large numbers several months after release,
establishment did not occur (McMurtry 1977b). One species of the
Phytoseiidae, Typhlodromus floridanus, was imported and
colonized in 1955. This was followed by importation of 10 different species
between 1961 and 1968. T. rickeri Chant was released in
the largest numbers (ca. 1/2 million), and became established on lemon trees
(McMurtry 1977b). But by 1958 this species disappeared from release orchards.
Several other species of Phytoseiidae, especially Iphiseius degenerans,
were recovered in large numbers during the season of release, but did not
become established. The life cycle of T. rickeri
Chant is typical of the Phytoseiidae, having egg, larva, protonymph and
deutonymph stages. No feeding occurs in the larval stages. At 22°C a
generation is completed in 9.4 days, and an initial mating is insufficient
for continued oviposition (McMurtry 1977b). The average rate of oviposition
ranges from 0.7/female/day at 15°C to almost two per day at 24-27°C.
Ovipositing females consume an average of 4.3 adult female hosts or 13.4
protonymphs of Tetranychus pacificus McG. per day at 24°C.
Feeding and reproduction occur readily on a variety of tetranychid mites,
including those which produce large amounts of webbing, such as T. pacificus, and those producing only a small amount such as
Panynychus citri (McMurtry 1977b). The
citrus rust mite P. oleivora is also a favorable
prey species. However, the common native species of southern California, Amblyseius hibisci (Chant) and A.
limonicus Garman &
McGregaor, fed bud did not reproduce on this prey. In contrast to the latter
predators, T. rickeri was found to be more
dependent on mite prey for reproduction, although pollen, honeydew, and scale
crawlers are fed on to some extent (McMurtry 1977b). Due to these biological
differences, it seemed that T.
rickeri would a significant addition to the predator complex on citrus in
California if establishment were possible (McMurtry & Scriven 1964b). Avocado Brown Mite, Oligonychus punicae (Hirst) [= coiti McGregor].--Presumably native to Central America and Mexico, this
tetranychid is the most injurious pest of avocado in southern California
(McMurtry 1977b). It feeds on foliage and causes a brownish discoloration and
some leaf drop when at high densities (Ebeling 1959). A classical biological
control program in southern california was initiated (Fleschner 1955,
McMurtry 1961), with emphasis since 1961 on predacious mites of the family
Phytoseiidae. The common native species seem to have certain limitations in
their ability to attain control (McMurtry & Johnson 1966). Field releases
of the imported predators were summarized by McMurtry (1977). No establishment
of any predatory species was reported, however. Other Pestiferous Acarina.--McMurtry (1977) reported that in the U.S. a stock of the
phytoseiid T. rickeri was sent from
California to Florida in 1962 and released against Texas citrus mite Eutetranychus banksi (McG.), six-spotted mite
Eotetranychus sexmaculatus (Riley) and citrus
rust mite P. oleivora, as well as the citrus
red mite. Short term recoveries were made but there were no reports of
establishment (Muma 1964). T.
rickeri was also shipped to
Texas from California where direct releases of several hundred did not result
in establishment (McMurtry 1977b). In Israel several phytoseiid mites
associated with citrus rust mite and tetranychids were introduced from Hong
Kong in 1960 (Swirski & Schlechter 1961), and it was reported that one, Amblyseius largoensis (Muma) was recovered the following season on Convolvulus sp. ca. one mile
from the release point (Swirski & Amitai 1961). Over 1/2 million A. largoensis were released, and establishment was thought to
occur. Several species of mite predators were sent to Israel from California
during 1960-65 (three indigenous species and two introductions from India),
and Phytoseiulus persimilis of South American
origin, was imported from Germany. Recoveries of P. persimilis
and T. rickeri were reported (Rosen 1967). Acarina
for Biological Control of Armored Scale Insects Mites and ticks (Acari) include a vast
assemblage of small arthropods which rivals the Insecta in diversity of
living habitats. They can be readily distinguished from insects by a
reduction in segmentation, presence of four pairs of legs in adults, and the
absence of compound eyes, antennae and wings. The Acari are separated into
several subgroups, generally recognized at ordinal or subordinal rank. Three
of these, the Astigmata, Mesostigmata and Prostigmata, include species that
prey on or parasitize armored scale insects. These species included within 10
families may be divided into two functional groups: those for which
biological data or claims for control are available and those which seem to
be of lesser importance. These taxa are discussed separately, with families
containing obligate or potentially important diaspidid parasites or predators
considered first. Secondly, taxa occasionally associated with diaspidids and
polyphagous predators will be mentioned. Finally, some mites which are often
found in association with scale insects, but which do not appear to have any
potential for control, will be noted. Hemisarcoptidae.--The Hemisarcoptidae (Astigmata) is a group of small,
soft-bodied mites associated with arboreal habitats such as polypore fungi,
vertebrate nests, and subcortical habitats. The family may be recognized in
the female by the position of the ovipore between or behind coxal fields IV,
in the male by the presence of a median sucker anterior to the genital region
and in all feeding stages by the sucker-like pretarsi which lack empodial
claws (Gerson et al. 1990). Deutonymphs are characterized by the loss of pretarsi
from legs IV, the reduction to a maximum of four setae of tarsi III-IV, and
the presence of a single large pigment spot under the propodosomal ocelli.
The genus Hemisarcoptes
Lignières is the only genus in this family associated with armored scale insects,
but all known species of this genus are obligate parasites or predators of
diaspidid scales. Species of Hemisarcoptes have been
known as important generalized predators of diaspidids for >100 yrs and
are found on many genera of host scale insects. Hemisarcoptes malus
(Shimer) was not only one of the first mites described from North America,
but was also the first mite utilized in a biological control program for
insect pests (Shimer 1868, Riley 1973). Ewing & Webster (1912) stated
that "it is quite evident that the oyster-shell scale [Lepidosaphes ulmi (L.)] is in many places
kept in check by mites... Of these mites, the most efficient was Hemisarcoptes malus." Similar claims
regarding the same pest in Canada were made by Lord (1947), and by Samarasinghe
& LeRoux (1966). Kaufmann (1977) reported that Hemisarcoptes were the most efficient predators of the
date palm scale, Parlatoria blanchardi (Targioni Tozzetti)
in the Sahel region of Niger, West Africa. Claims of relatively high rates of
predation affecting other economically important diaspidids were summarized
by Gerson & Schneider (1981). A recent literature survey on the worldwide
distribution of these mites shows non-specificity of diaspidid host
preference. A surprising feature is that no records appear for one of the
five major divisions of the Diaspididae, namely, the Odonaspidini (Gerson et
al. 1990). Regardless of the enthusiastic reports concerning Hemisarcoptes, very little data
is available on their biology and potential for biological control. Problems
include taxonomic uncertainties, scattered information on distribution and
bionomics, apparent uneven predation performance in the field, and lack of
publications on mass production techniques. Taxonomic Ambiguities.--Problems of misidentification and incomplete description
are found in the literature on Hemisarcoptes.
Shimer (1868) described the adults of the first species which he named "Acarus" malus, from Illinois. This
species was apparently first noted by Riley (1873), but Riley mistook another
mite for malus, and acrid
mite of the genus Thyreophagus.
This confusion most likely arose because these mites occur in association
with many species of diaspidid scale insects, both are very small, and the
general body forms are similar enough to be confused considering the optics
of the era. This misrepresentation of malus
led Lignières (1893a,b) to propose a new genus, Hemisarcoptes, for a species he described as H. coccisugus from France, while he regarded a species of
what is now recognized as Thyreophagusas
being identical with malus.
The confusion of the genera Hemisarcoptes
and Thyreophagus was
recognized by Michael (1903) who correctly aligned the European species of
Lignières (H. coccisugus) with its American
cogener (H. malus). All researchers after
Michael have regarded the European H.
coccisugus as synonymous
with the American H. malus despite the lack of
detailed study. Contemporary workers have also had to rely on erroneous
illustrations to distinguish species of Hemisarcoptes.
The species H. coccophagus Meyer, described
from South Africa, and H. dzhashii Dzhibladze, described
from Soviet Georgia, were distinguished from H. malus
only on the basis of very schematic figures of H. malus.
None of these species is recognizable on the basis of the original
descriptions. More confusion regarding Hemisarcoptes concerns the
dimorphic life cycle of these and other free-living astigmatid mites. The
deutonmyph (second nymphal instar, or hypopus) of these species is highly
modified morphologically and disperses by phoretic association with other
animals. These deutonymphs are so morphologically divergent from the other
life-cycle stages that association between stages is only possible through
rearing or collection of moulting deutonymphs. Deutonymphs of Hemisarcoptes were first
positively identified by Bartlett & DeBach (1952) in phoretic association
with the coccinellid beetle, Chilocorus
stigma (Say), in laboratory
cultures in California. The specific identity of these mites in uncertain.
Gerson (1967b) first described deutonymphs of H. coccophagus
from laboratory cultures and natural populations in Israel. These deutonymphs
were associated with the coccinellid, Chilocorus
bipustulatus (L.). Thomas (1961)
described a deutonymph collected from Chilocorus
cacti (L.) in Texas, as Vidia cooremani. Gerson (1967b) placed this species in the genus
Hemisarcoptes. The adults of
H. cooremani (Thomas) remain undescribed. The species-level
systematics of Hemisarcoptes
on a worldwide basis is currently under study by O'Connor & Houck (Gerson
et al. 1990). Bionomics.--Hemisarcoptes
coccophagus is most abundant
in the field in Israel during summer, although winter activity also occurs
(Gerson & Schneider 1981). Worldwide, Hemisarcoptes
species seem to be quite resistant to extreme climatic conditions. In Canada H. malus is the major natural control agent of the
oyster-shell scale during cold periods, as the mites may survive even when
temperatures decrease to -34°C (Lord & MacPhee 1953). The other major
natural enemy in these areas, the aphelinid wasp, Aphytis mytilaspidis
(LeBaron) is killed at -25°C. Observations on H. malus
in New York by Houck & O'Connor indicate that egg production continues
throughout the winter (Gerson et al. 1990). In the other extreme, Hemisarcoptes coccophagus acted as "a
most efficient predator" of date palm scale in the hot, dry climate of
the Sahel region of Niger, while Chilocorus
bipustulatus, which was
introduced to control the pest, was rendered ineffective by the unusually
harsh environment (Kaufmann 1977, Gerson et al. 1990). Freshly laid H. coccophagus
eggs hatch within 4-7 days at 21°C in the laboratory, and within 2-5 days at
28°C. Emerging larvae wander around the host scale avoiding strongly lighted
sites, and settle down to feed. These usually progress through thee moults
(to protonuymph, tritonymph and adult), feeding during each active stage. The
adults mate and females produce an average of 16 eggs. A complete life cycle
uninterrupted by a deutonymphal stage, requires ca. 26-28 days at 21°C, and
15-17 days at 28°C. The sex ratio is ca. 2 females/male (Gerson &
Schneider 1981). Individual H.
coccophagus which subsisted
on insufficient food (i.e., moribund scales) as larvae or protonymphs went
through a deutonymphal (hypopodial) stage in their development which was
consequently quite prolonged (Gerson et al. 1990). The deutonymphs, which
also serve to disperse the species, survived for 2-3 weeks in the laboratory
at 22°C under saturation conditions (Gerson & Schneider 1982). In
cultures of H. malus grown by Houck &
O'Connor, deutonymphs have never been produced in one year and 6 months of
continuous culture even though the scale hosts were allowed to completely
desiccate. Since deutonymphs of H.
malus do occur in field
populations, they may be rare, or their appearance may require chemical or
mechanical stimulation by the scale-piercing behavior of the Chilocorus beetles upon which
the deutonymphs are phoretic (Gerson et al. 1990). The deutonymph of H. coccophagus
may be seen wandering among scale insect colonies, but it is most commonly
encountered on Chilocorus bipustulatus in israel.
Occurrence on the beetles followed a seasonal trend, peaking in late summer.
By that time most beetles examined carried some deutonymphs, with an average
of over 30 per beetle (max. 202) (Gerson 1967b). The deutonymphs lack
mouthparts and do not harm the beetles, although heavily-laden Chilocorus appeared somewhat
sluggish. Deutonymphs were evenly distributed on male and female beetles,
indicating a similar attraction. This was later confirmed by choice-chamber
experiments, which also demonstrated strong vector attraction for the
deutonymphs, as 84.7% of all mites moved towards the Chilocorus-containing cells (Gerson & Schneider 1982).
Species of Chilocorus are
also predators of diaspidids, with the various beetle species attacking a
wide range of scale insect taxa. The potential for defining the full
geographic range for Hemisarcoptes
can be evaluated in terms of the known ranges of the phoretic partner as
indicated above. The affinity of Hemisarcoptes
deutonymphs for Chilocorus
beetles has been demonstrated by examination of museum collections of these
and related beetle species, as first suggested by Gerson (1967b). O'Connor
& Houck have examined specimens of 29 of the known species of Chilocorus in American museums,
with 12 of these species yielding collections of Hemisarcoptes (Gerson et al. 1990). Examination of their
scale-feeding beetles has yielded only one non-Chilocorus host for these mites, the related chilocorine
species Axion tripustulatum (DeGeer). The
only other reported host for these deutonymphs is the coccinellid Zagloba ornata Casey, and this record is from laboratory cultures
(Sellers & Robinson 1950). Distribution.--Hemisarcoptes
species distribution may be estimated from the literature and records of mite
deutonymphs obtained from museum collections. Knowledge of actual species
distributions is encumbered by problems of identification. On the basis of
specimens examined by Gerson et al. (1990), Hemisarcoptes malus
is regarded as widely distributed in North America, probably corresponding to
the range of its phoretic host, Chilocorus
stigma. Hemisarcoptes cooremani
is probably parapatric with H.
malus, with a known range
extending from southern Texas and California south through Honduras in
association with Chilocorus cacti. In the Old World, the
only recognizable species is H.
coccophagus. This species
has been verified from southern Europe (Spain), North Africa and the Middle
East in association with Chilocorus
bipustulatus and from
eastern and southern Africa associated with C. distigma
(Klug). Collections of Hemisarcoptes
deutonymphs from other areas in western North America, Africa, India,
Indonesia and the Philippines represent undescribed species. The specific
identity of central European Hemisarcoptes
remains questionable pending the examination of specimens. No deutonymphs
have as yet been recovered from Chilocorus
bipustulatus nor C. renipustulatus (Scriba) from this area. Also, the identity
of Hemisarcoptes reported
from South America (Flechtmann 1968, Fernandez 1973) must be reexamined.
There are no species of Chilocorus
native to this region, although C.
bipustulatus has been
introduced, probably from Europe, and is now widespread. There is a
possibility that South American and European populations may be conspecific. Hemisarcoptes species are not
yet reported from Japan or China, despite the diversity of species of Chilocorus in these areas. A
large series of Japanese Chilocorus
were examined by Gerson et al. (1990) without obtaining any Hemisarcoptes, although future
collecting in these areas may reveal their presence. However, the absence of Hemisarcoptes from the Australian
region may be predicted on the basis of the absence of Chilocorus species from that area (Gerson et al. 1990). Field Investigations.--There have been no controlled experimental studies
published concerning the field potential for biological control of scale
insects using Hemisarcoptes
(Gerson et al. 1990). The uneven field performance of these mites has been
noted by several authors. Simmonds (1958) reported them to attack from 1-100%
of the white peach scale, Pseudaulacaspis
pentagona (Targioni Tozzetti),
in Bermuda. Gerson (1967b) found that >70% of one population of the
California red scale, Aonidiella
aurantii (Maskell) were
attacked by H. coccophagus in Israel, but that
this rate dropped to ca. 20% later. Gulmahamad & DeBach (1978) recorded
mite parasitism rates of 42-66% on the San Jose scale, Quadraspidiotus perniciosus
(Comstock), in California during certain months, but scarcity or absence
during others. Some of this variance in predation might be due to variable
occurrence of mite predators (e.g., Cheletomimus
berlesei Oudemans), slow
dispersal rate of mobile stages, undetermined responses to chemical sprays,
and seasonal shifts in temperature and moisture conditions. When living under
optimal physical conditions and without chemical assault, as in laboratory
populations of diaspidids, Hemisarcoptes
mites may reduce population growth and actually endanger these scale cultures
(Sellers & Robinson 1950. Hemisarcoptes species usually occur in the field on or under ovipositing
scale insects which may still continue to produce progeny (Gulmahamad &
DeBach 1978, Gerson & Schneider 1981). Both female scale insects and
their eggs are fed upon (Ewing & Webster 1912), although crawlers, second
instar nymphs and prepupal male scale insects may also be less frequently
parasitized (Gulmahamad & DeBach 1978). Feeding mites (usually more than
one per scale) tend to take up the body color of their hosts (André 1942,
Gerson 1967b, Kaufmann 1977). For example, Hemisarcoptes malus
is bright purple on Lepidosaphes
beckii (Newmann), red on Epidiaspis leperii (Signoret), and yellow on Quadraspidiotus juglansregiae
(Comstock). This coloration often makes them difficult to locate (Gerson et
al. 1990). Regarding control potential, the effect of Hemisarcoptes species on their
host scale insects appears to be cumulative; i.e., parasitized scale insects
continue to deposit at least some eggs (Gulmahamad & DeBach 1978). Gerson
& Schneider (1981) applied the following general rule to female scale
insects parasitized by H. coccophagus: when fewer than
five mites developed on a single host, its fecundity would be reduced. A
scale insect attacked by five to 10 mites would fail to produce any progeny,
while the feeding of more than 10 mites usually causes the death of the host.
Scale insect species, size, age and sex, as well as mite species, may modify
this generalization (Gerson et al. 1990). The efficacy of Hemisarcoptes as biological control agents of scales was
verified by two introduction projects. The apparent absence of these mites
from western Canada suggested that they could be used there to control the
oystershell scale. Introductions of H.
malus from eastern Canada
began in 1917, and 23 years later the mite was widely distributed and at
times important in British Columbia. Turnbull & Chant (1961) rated this a
successful biological control attempt. The other project took place in
Bermuda, following an outbreak of Lepidosaphes
newsteadi Šulc on cedar
trees. Several natural enemies were introduced against this pest, including Hemisarcoptes malus. The mites were
introduced as deutonymphs on the bodies of 235 coccinellid beetles, mostly Chilocorus spp. (Bedford 1949),
and were subsequently found to attack the purple scale, Lepidosaphes beckii
on citrus. Hemisarcoptes mites are susceptible to many common pesticides. Sulfur and
winter oil were quite detrimental to the mite, but DDT, lead arsenate,
nicotine sulfate or summer oils had little effect under field conditions in
Canada (Lord 1947). Sellers & Robinson (1950) who had to eliminate Hemisarcoptes from their
laboratory cultures of diaspidids, used the acaricide Neotran with success. Mass Production.--Mass and individual mite rearing methods were described by
Gerson (1967b) and by Gerson & Schneider (1981), respectively. Large
numbers of H. coccophagus were produced by
growing diaspidids on potato tubers at 80% RH and colonizing them with
deutonymphs obtained from elytra of chilocorus
bipustulatus. Observations
on individual mites were made possible by substituting the scale insects'
shields with artificial covers. These consisted of a small amount of
collodion dissolved in iso-amyl-acetate. A few drops of the resultant
solution were placed on a smooth surface, and upon drying were used to cover
young female scale insects whose original shields had been removed. Only a
small aperture was left open, through which mites or their eggs were
introduced. Use of the artificial shield made direct observations on these
mites possible (Gerson et al. 1990). Gerson et al. (1990) concluded that under
certain conditions, especially when they are the only active natural enemies,
Hemisarcoptes species may be
important control factors of armored scale insects. However, this implies
that they are not very efficient in the presence of other predators and
parasites. The diversity of species of Hemisarcoptes,
their close association with Chilocorus
beetles, and their restriction to diaspidid hosts imply a relatively long
evolutionary association among members of this community. Therefore, it is
not surprising that the mites appear to be better adapted for coexisting with
their diaspidid hosts than for killing them directly, since such long
associations often tend toward reduced pathogenicity of the parasite. This
evolutionary trend might also explain why scales parasitized by Hemisarcoptes normally produce
at least some progeny, ensuring hosts for the progeny of the mites. However,
deductions based on the natural biology of the mite- scale insect- Chilocorus community may not be
valid in managed agroecosystems. Unpredictable performance, as has been
reported for Hemisarcoptes,
upsets control schedules and introduces unknown factors, detracting from the
mites' potential for biological control. Future studies should strive to
better understand Hemisarcoptes
control performance in such managed systems. A sound systematic base is an
obvious prerequisite; some of the unpredictability in prior studies may have
resulted from the interaction or succession of more than one species (Gerson
et al. 1990). Camerobiidae.--The Camerobiidae (Prostigmata) are a small family of mites
with long, "stilted" legs, a ventrally directed gnathostoma, weak
palpi and looped peritremes. Species in one genus, Neophyllobius, have been reported to feed on diaspidid
crawlers. McGregor (1950) quoted unpublished observations made by Pence, who
noted that when attacking crawlers, the mites inject their prey with some
opiate. The crawlers subsequently relax and allow their body juices to be
sucked out. Meyer (1962) added to these observations, reporting that nymphs
and adults of N. ambulans Meyer fed on crawlers
of the California red scale, Aonidiella
aurantii, but not on settled
scale insects. The predator appeared to be rather scarce on South African
citrus trees, and therefore Meyer noted that it was probably of no economic
importance in natural control of red scale. A different opinion was by
Richards (1962) who believed that a species of Neophyllobius was the principal predator of Quadraspidiotus ostreaeformis (Curtis) in New
Zealand. The mites were very common wherever the scale insect was abundant,
but no crawlers were actually observed consumed. In the laboratory the
predatory mites were seen with their mouthparts inserted in adult scales,
sucking them dry. Richards (1962) also thought the mites appeared to be
injecting some relaxing chemical into prey, as the latter did not struggle. Cheyletidae.--The majority of the prostigmatid family Cheyletidae are
free-living predators, while others are ectoparasites of birds, mammals or
rarely insects. Free-living cheyletids are slow-moving, yellow or orange and
usually ambush prey. The morphological characteristic best defining the
Cheyletidae is the prominent palpal thumb-claw complex, with the palptarsus
bearing strong sickle and/or comb-like setae (Gerson et al 1990). These mites
often occur on plants, and several species have been observed to feed on
diaspidid crawlers. Cheletogenes
ornatus (Canestrini &
Fanzago) was observed feeding on crawlers in many parts of the world (Avidov
et al. 1968, Gerson et al. 1990). The role of this predator in citrus groves
in Israel was studied by Avidov et al. (1968). The mite was reared in
plaster-of-Paris cells and fed crawlers of the chaff scale, Parlatoria pergandii Comstock. Females deposited <a dozen eggs
throughout their lives under these conditions. Egg development took ca. 10
days, the larva and two nymphal instars another 47 days, and each molt
required 2.5 days, total immature development taking 64 days. Oviposition
started after another 25 days, indicating the total egg to egg cycle was
about 3 months at 28°C. During this study, female mites consumed an average
of 90 crawlers during their adult lives, which lasted an average of 43 days
(Gerson et al. 1990). Cheletogenes ornatus was
reared on eggs of the olive scale, Parlatoria
oleae (Colvée) (Zaher &
Soliman 1971). It was reported that the predator's complete development took
about 25 days at 29°C. Mites in that study produced an average of 16.8 eggs
per female, and each female consumed ca. 170 scale insect eggs (males 125)
and lived for 16.6 days. Such differences in life cycle parameters obtained
in the two laboratory studies of this mite have also been reported for other
species (Gerson 1985). Female survival is dependent on the ambient RH, and at
28°C, mites kept at 0% RH lived only 3 days, with the survival time at 21, 50
and 80% RH being 12.5, 14.5 and 26 days, respectively (Avidov et al. 1968).
Starved females (at high RH and 28°C) survived an average of 16 days (range
1-33) (Gerson et al. 1990). Exposure of C. ornatus
females to citrus leaves dipped in several pesticides showed that the
fungicide zineb had little effect on mite survival. The acaricide
chlorobenzilate, however, was very toxic, causing almost total mortality 24 h
post-treatment (Avidov et al. 1968). Field studies indicated that this
predator was much more common on citrus bark (where diaspidids flourish) than
on leaves or fruit. Mite numbers were usually low during winter, rising in
summer and peaking during autumn. These observations, along with the
laboratory data noted above, indicate that C. ornatus
has two summer generations on citrus in Israel. Reproduction ceases during
winter, probably in connection with female diapause. Available information indicates that C. ornatus has a low rate of increase, a pronounced winter
ebb and is difficult to rear in the laboratory. But it is a hardy species
capable of survival under adverse conditions, and it is also the dominant
acarine predator of armored scale insects on citrus. Avidov et al. (1968) recommended that efforts be directed at conserving the
predator in the field. Data on another diaspidid-feeding cheyletid
presented by Wafa et al. (1970) show that adult females and males of Eutogenes africanus Wafa & Soliman consumed an average of 186
and 156 eggs of Parlatoria oleae, respectively. The life
cycle at 29°C required ca. 31 days, and each female deposited an average of
16 eggs. Other cheyletids observed feeding on armored scale insect crawlers
in the field include Hemicheyletia
bakeri (Ehara) which feeds on the yellow scale, Aonidiella citrina
(Coquillett) in Florida (Muma 1975) and Cheletominum
berlesei (Oudemans) on the
latania scale, Hemiberlesia lataniae (Signoret) in
California (Ebeling 1959) and on Parlatoria
spp in Israel (Gerson 1967a). Cheletominus
berlesei has also been
observed feeding on Hemisarcoptes
mites associated with Lepidosaphes
beckii in California (Gerson
et al. 1990), with numbers of Hemisarcoptes
negatively correlated with Cheletominum
density. Additional cheyletid species, some as yet undescribed were observed
to feed on various diaspidids on fruit trees in New Zealand and the Cook
Islands (Gerson et al. 1990). Eupalopsellidae.--This family of prostigmatid mites is characterized by very
long palpi and chelicerae, a rather reduced palpal thumb-claw complex and the
modification of the pretarsal empodia into two pairs of capitate raylets.
Species in two genera are known to feed on diaspidids (Gerson et al. 1990). Saniosulus nudus
Summers is an active predator of crawlers of Parlatoria spp. on citrus in Israel. The prey is held by
the mite's anterior legs as the predator inserts its cheliceral stylets into
the crawler's body. Feeding may proceed for 30-40 min until the dried prey
remains are pushed off the chelicerae. All active stages of this species feed
on diaspidid eggs and crawlers. Second-stage nymphs and adults are also
attacked but do not appear to be seriously affected (Gerson & Blumberg
1969). Observations once a month in a citrus grove
indicated that populations of S.
nudus on bark peaked during
late summer and then declined (Gerson 1967a). These mites have been
subsequently observed feeding on various other species of diaspidids in
Israel (Gerson et al. 1990). The species was experimentally cultured on
Florida red scale, Chrysomphalus
aonidum (L.), reared on
green lemon fruits. The generation time of S. nudus
was ca. 3 weeks at 24°C and 2 weeks at 28°C, the latter being less than half
the time required for diaspidid generations. Each female produced 40-50 eggs,
regardless of prior mating. Copulation itself is rather prolonged, with the
female dragging the male around behind her. if introduced into laboratory
cultures of armored scale insects, S.
nudus may affect them to the
extent that control measures must be implemented (Gerson & Blumbeg 1969).
Eupalopsis maseriensis
(Canestrini & Fanzago) has also been collected from citrus bark in Israel
(Gerson 1966). It is a rare predator, whose feeding habits are similar to
those of S. nudus. Phytoseiidae.--This family among free-living mesostigmatid mites is
characterized by having 20 or fewer pairs of dorsal setae. Some species are
efficient predators of phytophagous mites and have been intensively studied
(Tanigoshi 1983). Several species of Phytoseiidae were collected near armored
scale insects (Baccetti 1960, Muma 1975) but their role in such communities
is uncertain. Typhlodromus baccetti Lombardini was a
constant associate of juniper scales, Carulaspis
spp., in Tuscany, Italy (Baccetti 1960). Mites gain access under the scales'
shields, where they feed on the eggs. The predator overwinters as an egg,
matures in May and undergoes two summer generations. It was considered a
scale-insect predator of some importance. Other phytoseiid species have been
observed to feed, oviposit and complete their life cycles when offered
diaspidid crawlers as food in the laboratory (Tanigoshi 1983). Whether such
diets are also used in the field, and to what extent, remains unknown (Gerson
et al. 1990). Other Predators / Parasites.-- Gerson et al. (1990) enumerate several other families in
the Prostigmata, which are generally polyphagous predators or parasite, with
diaspidids sometimes being included in their diets: Anystidae.--Species in this family are fast runners which move about in
a corkscrew or figure eight pattern (Muma 1975). These relatively primitive
prostigmatid mites possess a palpal thumb-claw complex in which the
palptarsus extends well beyond the tibial claw. Anystis agilis
Banks was observed by Muma (1975) to feed on crawlers of purple scale, Lepidosaphes beckii, in Florida. Ewing &
Webster (1912) noted that this mite is a common predator of oyster-shell
scale, Lepidosaphes ulmi, crawlers and eggs. Bdellidae.--Mites in this family have an elongate rostrum, with long
palpi which terminate in strong setae and lack a palpal thumb-claw complex.
Ewing & Webster (1912) reported species of Bdella and Cyta
associated with and probably feeding on L.
ulmi, and Muma (1975)
reported Bdella distincta (Baker & Blalock)
feeding on eggs and crawlers of L.
beckii. The latter species
appeared to be widely distributed in unsprayed citrus groves in Florida
(Gerson et al. 1990). Cunaxidae.--This family is morphologically similar to the closely
related Bdellidae, differing in the form of the palpi which are raptorial and
end in a claw. A species of Cunaxoides
was reported by Baker & Wharton (1952) to feed on diaspidids. Erythraeidae.--Species in this family are usually parasitic on various
arthropods during their larval instar. The nymphs and adults are predaceous.
The family may be distinguished by having numerous body setae, a palpal
thumb-claw complex, and long, straight cheliceral stylets. Species in the
genus Balaustium feed on
various diets, from flower pollen to various insects including diaspidid
crawlers (Gerson et al. 1990). These mites are also known to bite humans
(Newell 1963). Pyemotidae.--These mites are usually parasites of arthropods. Adult
females have reduced palpi, capitate prodorsal sensillae, and a series of
segment-like plates on the dorsal opisthosoma. Females are frequently
physogastric, swelling enormously as they feed. Many pyemotid species of
polyphagous parasites, feeding on a wide variety of arthropod hosts, often
Lepidoptera or Coleoptera. Vaivanijkul & Haramoto (1969) reported that Pyemotes boylei Krczal parasitized Diaspis echinocacti
(Bouché) in Hawaii. An undetermined species was found to parasitize females
of Lindingaspis rossi (Maskell) in New Zealand.
Rates of parasitism of the latter species ranged from 12-15% (Gerson et al.
1990). Stigmaeidae.--Mites in this family have an ovoid or elongate dorsum that
is usually covered by plate-like sclerites. They have a palpal thumb-claw
complex and short, stylet-like chelicerae, but lack peritremes. Agistemus terminalis (Quayle) is a predator of the arrowhead scale, Unaspis yanonensis (Kuwana) in Japan (Ehara 2962). Another
species, Agistemus floridanus Gonzalez, feeds on
crawlers of A. aurantii in Florida (Muma
1975). Associated Species.--Gerson et al. (1990) discuss mites of various taxa which
are sometimes encountered under the shields of dead scale insects or may be
found among live diaspidids without actually harming them. The most frequent
and widespread associates are species of the asigmatid genus Thyreophagus (Acaridae). These
are often erroneously called T.
entomophagus Laboulbène, but
probably represent T. angustus (Banks) or related
species. The cigar-shaped, milk-colored mites have a strongly reduced dorsal
setation, but retain pretarsal empodial claws. The confusion of these mites
with Hemisarcoptes in early
literature is common. Ewing & Webster (1912) claimed that these mites
were found only under shields of dead Lepidosaphes
ulmi, feeding exclusively on
dead material. Other records from this habitat include those of Kosztarab
(1963) and Muma (1975) from the U.S. and Williams (1970) from Mauritius.
Gerson (1971) found Thyreophagus
under various diaspidids in Israel and Canada, and reared them for several
generations on a fungal diet. These mites have been commonly collected from
various diaspidid species in the U.S. (Gerson et al. 1990), where gut-content
analysis indicated fungi making up a large portion of their diet. They also
have been collected from dead armored scales in New Zealand (Gerson et al.
1990). The deutonymph described as Thyreophagus
(= Monieziella) brevipes by Banks (1906)
probably represents that of Hemisarcoptes
malus. Another genus of astigmatid mites sometimes
found in association with diaspidids is Tyrophagus (Acaridae) reported by
Williams (1970) to be numerous among older scale masses of Aulacaspis tegalensis (Zehntner) on sugar cane in Mauritius. These
mites are common saprophages in many situations and commonly contaminate
laboratory cultures of other mite species. A number of mites whose normal habitat is
the bark of trees has been reported in association with scale insects.
Species in several families of the order Cryptostigmata (= Oribatei) were
reported in association with various armored scales in Ohio (Kosztarab 1963).
These associations are probably accidental, however (Ewing & Webster
1912). Species of the prostigmatid family Tydeidae are quite ubiquitous mites
sometimes associated with diaspidids. Ewing & Webster (1912) often found Triophtydeus (= Tydeus) coccophagus (Ewing) with L. ulmi
and commented, "That this mite is predaceous upon scale insect or its
eggs, there is but little doubt." But, in their words, "the case
here is not so conclusive." Brickhill (1958) demonstrated that tydeids
may complete their development and oviposit while offered spider mite eggs
alone, but all eggs that had been fed on subsequently hatched. It is possible
that even if tydeids, which generally feed on honeydew and sooty mold fungi,
occasionally try to pierce diaspidid eggs, the latter remain undamaged. Gerson et al. (1990) mentioned some
perplexing observations in regard to associations with plant feeding mites.
Ebeling (1948) noted that settlement of the citrus red mite, Panonychus citri (McGregor) (Tetranychidae) on citrus leaves rendered
the latter unsuitable for crawlers of A.
aurantii. This adverse
effect was observed two days after mite settlement, and no crawlers survived
on leaves which had been colonized by the mite 12 days or more. Gerson et al.
(1983) found that the palm infesting tenuipalpid Taoiella indica
Hirst may place its eggs only within colonies of the parlatoria date scale, Parlatoria blanchardi. Such eggs were found in 60.3% of scale
colonies examined. Gerson et al. (1990) concluded that a consideration
of feeding modes allows a separation of those mites having some control
potential into two groups, namely predators and parasites. Species of Hemisarcoptes and Pyemotes may be considered a
parasite since host death, if occurring, usually occurs after long term
feeding. All other important mites species are predators. Available data
strongly suggest that at present species of the former group appear to be
more promising as agents for the control of armored scale insects. This
conclusion is based not only on the Hemisarcoptes
data, but also on encouraging results of Bruce (1983) in regards to Pyemotes species used against
stored food pests. Such results also serve to remind us that Acari can and should
be used much more vigorously against insect pests. The overview of Acari as
natural enemies of armored scale insects should serve to emphasize the
paucity of information concerning such relationships. Claims that mites
actually control diaspidid populations in the field are scarce, and mostly
refer to species of Hemisarcoptes.
It is sad that all too few mite enemies of armored scale insects have been
recognized, collected, reported and considered by researchers, although some
notable exceptions occur (Ewing & Webster 1912, Gerson et al. 1990).
Gerson & van de Vrie (1979) noted that acarine enemies of armored scale
insects are a group in regard to which most work is still done at the first,
preliminary stage: recognizing the natural enemy. Acarina
in the Marine Environment Mites inhabit broad habitats in the marine environment, where
some species are associated with and/or attack other marine organisms,
including marine mammals. Kenyon (1965) and Newell (1943-73) have specialized
in these mites which include a vast array of species. There are only limited
ongoing studies of their economic importance and use as biological control
organisms. REFERENCES: [Additional references may be found at
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