FILE: <bc-40.htm> GENERAL INDEX [Navigate to MAIN MENU ]
[For
educational purposes only; do not review, quote or abstract]
BIOLOGICAL CONTROL OF AND BY ACARINA
(Contacts)
----Please CLICK
on
desired underlined categories [ to search for Subject Matter,
depress Ctrl/F ]:
|
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 is 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
parasites 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 MELVYL Library ] Alfeev,
N. I. 1940. the utilization of Hunterellus
hookeri How. for the control
of the ticks Ixodes ricinus L. and Ixodes persulcatus Sch. with reference to the peculiarities of
their metamorphosis under the conditions of the Province of Leningrad, p.
23-5. In: Pavlovsky (ed.),
2nd Conf. Parasitol. Problems, Nov. 1940, Akad. Nauk. S.S.S.R. Isv. André,
M. 1942. Sur l'Hemisarcoptes
malus Shimer (= coccisugus Lignières)
(Acarien). Bull. du Mus. Histoire Naturelle, 2eS 14: 173-80. Avidov,
Z., D. Blumberg & U. Gerson. 1968. Cheletogenes
ornatus (Acarina:
Cheyletidae), a predator of the chaff scale on citrus in Israel. Israel J.
Ent. 3: 77-94. Babayan,
G. A. & S. B. Oganesyan. 1979. Natural enemies of the Armenian mussel
scale (Lepidosaphes amlicola Borchs.) and
possibilities of conserving them in the presence of chemical treatments.
Biologischeskii Zhurnal Armenii 32: 194-99. [in Russian]. Baccetti,
B. 1960. Le cocciniglie Italiane delle Cupressaceae. Redia 45: 23-111. Baker,
E. W. & A. E. Pritchard. 1953. A guide to the spider mites of cotton.
Hilgardia 22: 203-34. Baker,
E. W. & G. W. Wharton. 192. An Introduction to Acarology. Macmillan Co.,
NY. 465 p. Banks,
N. 1906. A revision of the Tyroglyphidae of the United States. U.S. Dept.
Agr. Tech. Ser. No. 13. 34 p. Bartlett,
B. & P. DeBach. 1952. New natural enemies of avocado pests. Citrus Leaves
32(10): 16-17. Bedford,
E. C. G. 1949. Report of the plant pathologist, p. 11-19. In: Rept. of the Dept. Agr. for
the Year 1949, Bermuda Bd. Agr. Bellows,
T. S., Jr. & T. W. Fisher, (eds) 1999. Handbook of Biological Control:
Principles and Applications. Academic Press, San Diego, CA. 1046 p. Bennett,
F. D. & I. W. Hughes. 1959. Biological control of insect pests in
Bermuda. Bull. Ent. Res. 50: 423-36. Bishopp,
F. C. 1934. Records of hymenopterous parasites of ticks in the United States.
Wash. Ent. Soc. Proc. 35: 87-8. Böhm,
H. 1966. Ein Beitrag sur biologischen Bekämpfung von Spinnmilben in
Gewächshäusern. Pflanzenschutz Ber. 34: 65-77. Boudreaux,
H. B. 1963. Biological aspects of some phytophagous mites. Ann. Rev. Ent. 8:
137-54. Boyce,
A. M. 1936. The citrus red mite, Paratetranychus
citri McGregor, in
California and its control. J. Econ. Ent. 29: 125-30. Boyle,
W. W. 1957. On the mode of dissemination of the two-spotted spider mite, Tetranychus telarius (L.). Hawaii. Ent.
Soc. Proc. 16: 261-68. Bravenboer,
L. 1968. Biological control of mites in glasshouses. 2nd Internatl. Cong.
Acarol. Proc., Nottingham, England, July 19-25, 1967. p. 365-71. Bravenboer,
L. & G. Dosse. 1962. Phytoseiulus
riegeli Dosse als Prädator
einiger Schdmilben aus der Tetranychus
urticae Koch. Ent. Expt.
& Appl. 5: 291-304. Brickhill,
C. D. 1958. Biological studies of two species of tydeid mites from
California. Hilgardia 27: 601-20. Bruce,
W. A. 1983. Mites as biological control agents of stored product pests, p.
74-8. In: M. A. Hoy, G. L.
Cunningham & L. Knutson (eds.), Biological Control of Pests by Mites.
Univ. Calif. Div. Agr. & Nat. Res., Special Publ. 3304. Cagle,
L. R. 1949. Life history of the two-spotted spider mite. Va. Agr. Expt. Sta.
Tech. Bull. 113. 31 p. Chant,
D. A. 1961. An experiment in biological control of Tetranychus telarius
(L.) (Acarina: Tetranychidae) in a greenhouse, using Phytoseiulus persimilis
Athias-Henriot (Phytoseiidae). Canad. Ent. 93: 437-43. Chant,
D. A. 1963. Some mortality factors and the dynamics of orchard mites. Ent.
Soc. Canad. Mem. 32: 33-40. Chant,
D. A. 1965. Generic concepts in the family Phytoseiidae (Acarina:
Mesostigmata). Canad. Ent. 97: 351-74. Cooley,
R. A. 1928. Tick parasites. Mont. St. Bd. Ent. 7th Bien. Rept. p. 10-16. Cooley,
R. A. 1932. The Rocky Mountain wood tick. Mont. Agr. Expt. Sta. Bull. 268. 58
p. Cooley,
R. A. & G. M. Kohls. 9133. A summary of tick parasites. 5th Pac. Sci.
Cong. Proc. 5: 3375-81. Davis,
D. W. 1952a. Influence of population density on Tetranychus multisetis.
J. Econ. Ent. 45: 652-54. Davis,
D. W. 1952. Some effects of DDT on spider mites. J. Econ. Ent. 45: 1011-19. Davis,
D. W. 1956. The problem of species determination in spider mites. Proc. Utah
Acad. Sci. 33: 183-84. Davis,
D. W. 1959. Resistance to insecticides among Utah orchard pests. Proc. Utah
Acad. Sci. 36: 177. Davis,
D. W. 1965. New developments in orchard pest control. Proc. Utah Hort. Soc.
for 1964. p. 79-82. Davis,
D. W. 1967a. McDaniel mites and the resistance problem in Utah. Utah Farm
& Home Sci. 28(1): 26-7, 31. Davis,
D. W. 1967b. Biological control of orchard mites - a look into the future.
Proc. Utah Hort. Soc. for 1966: 43-7. Davis,
D. W. 1970b. Variations in the anatomy of Typhlodromus
occidentalis (Acarina:
Phytoseiidae). Ann. Ent. Soc. Amer. 63: 696-9. Davis,
D. W. 1970c. Integrated control of apple pests in Utah. Utah Science 31(2):
43-5, 48. Davis,
D. W. & G. L. Nielsen. 1958. The biology and distribution of the McDaniel
mite in Utah. Utah Acad. Sci. 35: 166. DeBach,
P. & S. Landi. 1961. New parasites of California red scale. Calif.
Citrograph 44: 290, 301, 303-04. Dosse,
G. 1958. Über einige neue Raubmilbenarten (Acar., Phytoseiidae).
Pflanzenschutz Ber 21: 44-61. Dosse,
G. 1959. Der Einfluss von Temperatur und Nahrung auf verschiedene
Raubmilbenarten und Hinweise auf die Möglichkeit einer biologischen
Bekämpfung von Spinnmilben in Gewächshäusern. 4th Internatl. Cong. Crop.
Protect. Proc., Hamburg 1957. p. 929-32. Dzhibladze,
K. N. 1969. A new species of predatory mite (Hemisarcoptidae) attacking the
orange scale, Cornuaspis beckii Newman (Homoptera,
Coccoidea), in Western Georgia. Ent. Rev. 48: 435-36. Ebeling,
W. 1948. Effect of citrus red mites on a California red scale population. J.
Econ. Ent. 41: 109. Ebeling,
W. 1950. Subtropical Entomology. Lithotype Proc. Co., San Francisco. 575 p. Ebeling,
W. 1959. Subtropical Fruit Pests. Univ. Calif. Div. Agr. Sci. Pub. 436 p. Ehara,
S. 1962. Notes on some predatory mites (Phytoseiidae and Stigmaeidae). Japan.
J. Appl. Ent. & Zool. 6: 53-60. Ehara,
S. 1964. The tetranychid mites of Japan. 1st Internatl. Cong. Acarol. Proc.,
Ft. Collins, Colo 1963. Acarologia 6: 409-13. English,
L. L. & G. F. Turnipseed. 1941. The influence of temperature and season
on the citrus red mite. J. Agr. Res. 62: 65-78. Ewing,
H. E. & R. L. Webster. 1912. Mites associated with the oyster-shell scale
(Lepidosaphes ulmi Linne). Psyche 19: 121-34. Fernandez,
R. V. 1973. Acaros de cítricos en la provincia de Tucumán. REvista Agron. del
Noroeste Argentina 9: 413-526. Flechtmann,
C. H. W. 1968. Hemisarcoptes
malus um predator do pulgao
lanigero do pesseguero. Resumos de le Reuniao Anual Sociedade Brasileira de
Entomologia. Piracicaba, Sao Paulo. p. 75. Fleschner,
C. A. 1953. Host-plant resistance as a factor influencing population density
of citrus red mites on orchard trees. J. Econ. Ent. 45: 687-95. Fleschner,
C. A. 1955. Natural mite enemies introduced from Guatemala. Citrus Leaves 35:
28. Fleschner,
C. A. 1958. Field approach to population studies of tetranychid mites on
citrus and avocado in California. 10th Internatl. Cong. Ent. Proc. 2: 669-74. Fleschner,
C. A., M. E. Badgley, D. W. Ricker & J. Hall. 1956. Air drift of spider
mites. J. Econ. Ent. 49: 624-27. Fukuda,
J. & N. Shinkaji. 1954. Experimental studies on the influence of
temperature and relative humidity upon the development of citrus red mite (Metatetranychus citri McGregor); (1) on the
influence of temperature and relative humidity upon the development of the eggs.
Nat. Tokai-kiaki Agr. Expt. Sta. Bull. 2: 160-69. [in Japanese w/ English
summary]. Gerson,
U. 1964. Parlatoria cinerea, a pest of citrus in
Israel. FAO Plant Prot. Bull. 12: 82-5. Gerson,
U. 1966. A redescription of Eupalopsis
maseriensis (Canestrini and
Fanzago) (Acarina: Eupalopsellidae). Israel. J. Zool. 15: 148-54. Gerson,
U. 1967a. The natural enemies of the chaff scale, Parlatoria pergandii
Comstock, in Israel. Entomophaga. 12: 97-109. Gerson,
U. 1967b. Observations on Hemisarcoptes
coccophagus Meyer
(Astigmata: Hemisarcoptidae), with a new synonym. Acarologia 9: 632-8. Gerson,
U. 1967c. Some cheyletid and pseudocheylid mites from Israel. Acarologia 9:
359-69. Gerson,
U. 1968b. Some raphignathoid mites from Israel. J. Nat. Hist. 2: 429-37. Gerson,
U. 1971. The mites associated with citrus in Israel. Israel J. Ent. 6: 5-22. Gerson,
U. 1973. The mites associated with armored scale insects. Proc. 35d
Internatl. Cong. Acarol., Prague. W. Junk, The Hague. p. 653-4. Gerson,
U. 1985. Other predaceous mites and spiders, p. 205-10. In: W. Helle & M. W. Sabelis (eds.), Spider Mites,
Their Biology, Natural Enemies and Control. Elsevier, Amsterdam, vol. 1B. Gerson,
U. 1992a. Biology and control of the broad mite, Polyphagotarsonemus latus
(Banks) (Acari: Tarsonemidae). Expt. Appl. Acarol. (in press). Gerson,
U. & D. Blumberg. 1969. Biological notes on the mite Saniosulus nudus.
J. Econ. Ent. 62: 729-30. Gerson,
U. & E. Cohen. 1989. Resurgences of spider mites (Acari: Tetranychidae)
induced by synthetic pyrethroids. Exp. Appl. Acarol. 6: 29-46. Gerson,
U. & Y. Rössler. 1982. Integrated citrus pest management in Israel, p.
150-54. In: P. J. Cameron et
al. (eds.), Proc. Australasian Workshop on Development & Implementation
of IPM, Govt. Printer, Auckland. Gerson,
U. & R. Schneider. 1981. Laboratory and field studies of the mite Hemisarcoptes coccophagus Meyer (Astigmata:
Hemisarcoptidae), a natural enemy of armored scale insects. Acarologia 22:
199-208. Gerson,
U. & R. Schneider. 1982. The hypopus of Hemisarcoptes coccophagus
Meyer (Acari: Astigmata: Hemisarcoptidae). Acarologia 23: 171-6. Gerson,
U. & R. L. Smiley. 1990. Acarine Biocontrol Agents: An Illustrated Key
and Manual. Chapman & Hall. 174 p. Gerson,
U. & V. Vacante. 1992. The use of indigenous acarine predators to control
citrus mite pests. Proc. VII Internatl. Citrus Congr. (in press). Gerson,
U. & M. van de Vrie. 1979. The potential of mites in the biological
control of mites and insect pests. Proc. 4th Intern. Cong. Acarol.,
Saalfelden. Akademiai Kiado, Budapest. p. 629-35. Gerson,
U., R. Kenneth & T. I. Muttath. 1979. Hirsutella
thompsonii, a fungal
pathogen of mites. II. Host-pathogen interactions. Ann. Appl. Biol. 91:
29-40. Gerson,
U., A. Venezian & D. Blumberg. 1983. Phytophagous mites on date palms in
Israel. Fruits 38: 133-35. Gerson,
U., B. M. O'Connor & M. A. Houck. 1990. Acari, p. 77-97. In: D. Rosen (ed.), Armored
Scale Insects, Their Biology, Natural Enemies and Control. World Crop Pests,
Vol. 4B. Elsevier Science. Gifford,
D. 1959. A sarcoptiform mite apparently new to Britain. Ent. Mon. Mag. 95: 1. Gould,
H. J., N. W. Hussey & W. J. Parr. 1968. Large scale commercial control of
T. urticae on cucumbers by the predatory mite Phytoseiulus riegeli. 2nd Internatl. Cong.
Acarol. Proc., Nottingham, England, Jul 19-25, 1967. p. 383-88. Gulmahamad,
H. & P. DeBach. 1978. Biological control of the San Jose scale Quadraspidiotus perniciosus (Comstock)
(Homoptera: Diaspididae) in southern California. Hilgardia 46: 205-38. Henderson,
C. F. & J. K. Holloway. 1942. Influence of leaf age and feeding injury on
the citrus red mite. J. Econ. Ent. 35: 683-86. Henneberry
& Boswell. 1963. [NOT IN McMurtry 1977b]. Huffaker,
C. B., C. E. Kennett & G. L. Finney. 1962. Biological control of olive
scale, Parlatoria oleae (Colvée) in California by
imported aphytis maculicornis (Masi)
(Hymenoptera: Aphelinidae). Hilgardia 32: 541-636. Hussey,
N. W. & W. J. Parr. 1965. Observations on the control of Tetranychus urticae Koch on cucumbers by
the predatory mite Phytoseiulus
riegeli Dosse. Ent. Expt.
& Appl. 8: 271-81. Jeppson,
L. R., C. A. Fleschner & J. O. Complin. 1957. Influence of season and
weather on citrus red mite populations on lemons in southern California. J.
Econ. Ent. 50: 293-300. Karsemeijer,
M. M. D. 1973. Observations on the enemies of the oyster shell scale Lepidosaphes ulmi, on apple in the
Netherlands. Netherlands J. Plant Path. 79: 122-24. Kaufmann,
T. 1977. Hemisarcoptes sp.
and biological control of the date palm scale, Parlatoria blanchardi
Targioni, in the Sahel region of Niger. Environ. Ent. 6: 882-84. Kenyon,
K. W., C. E. Yunker & I. M. Newell. 1965. Nasal mites (Halarachnidae) in
the sea otter. J. Parasitol. 51: 960. Kosztarab,
M. 1963. The armored scale insects of Ohio (Homoptera: Coccoidae:
Diaspididae). Ohio Biol. Surv. Bull. 2. 120 p. Laing,
J. 1968. Life history and life table of Phytoseiulus
persimilis Athias-Henriot.
Acarologia 10: 578-88. Langenscheidt,
M. 1966. Phytoseiulus riegeli Dosse, ein biologisches
Bekämpfungsmittel gegen Spinnmilben im Gewächshaus (Acari, Phytoseiidae).
Ztschr. f. Pflanzenschutz 73: 452-57. Larrouse,
F., A. G. King & S. B. Wolbach. 1928. The overwintering in Massachusetts
of Ixodiphagus caucurtei. Science 67: 351-53. Lignières,
J. 1893a. Etude zoologique et anatomique du Tyroglyphus malus
et de sa nymphe hypopiale. Mémoires de la Société Zool. de France 6: 5-15. Lignières,
J. 1893b. Etude zoologique et anatomique de l'Hemisarcoptes coccisugus.
Mémoires de la Société Zool. de France 6: 16-25. Lord,
F. T. 1947. The influence of spray programs on the fauna of apple orchards in
Nova Scotia. II. Oystershell scale. Canad. Ent. 79: 196-209. Lord,
F. T. & A. W. MacPhee. 1953. The influence of spray programs on the fauna
of apple orchards in Nova Scotia. VI. Low temperatures and the natural
control of the oystershell scale, Lepidosaphes
ulmi (L.) (Homoptera:
Coccidae). Canad. Ent. 85: 282-91. Mathys,
G. & E. Guignard. 1967. Enseignements recueillis au cours de neuf ans de
travaux avec Prospaltella perniciosi Tow., parasite du
pou San José (Quadraspidiotus
perniciosus Comst.).
Entomophaga 12: 212-22. McClanahan,
R. J. 1968. Influence of temperature on the reproductive potential of two
mite predators of the two-spotted spider mite. Canad. Ent. 100: 549-56. McCoy,
C. W. & L. G. Albrigo. 1975. Feeding injury to the orange caused by the
citrus rust mite, Phyllocoptruta
oleivora (Prostigmata:
Eriophyidae). Ann. Ent. Soc. Amer. 68: 289-97. McCoy,
C. W. & T. L. Couch. 1982. Microbial control of the citrus rust mite with
the mycoacaracide Mycar. Florida Ent. 65: 116-27. McCoy,
C. W. & A. M. Heimpel. 1980. Safety of potential mycoacaracide, Hirsutella thompsonii to vertebrates. Environ. Ent. 9: 24-49. McCoy,
C. W. & R. F. Kanavel. 1969. Isolation of Hirsutella thompsonii
from the citrus rust mite Phyllocoptruta
oleivora, and its
cultivation on various synthetic media. J. Invert. Pathol. 14: 386-90. McGregor,
E. A. 1950. Mites of the genus Neophyllobius.
Bull. So. Calif. Acad. Sci. 49: 55-70. McMurtry,
J. A. 1961. Current research on biological control of avocado insect and mite
pests. Calif. Avocado Soc. Yearbk. 45: 104-06. McMurtry,
J. A. 1963. Diaspidine scale insects as prey for certain phytoseiid mites, p.
151-4. In: Adv. Acarol.,
Cornell Univ. Press. McMurtry,
J. A. 1969. Biological control of citrus red mite in California. In: H. D. Chapman (ed.), Proc.
1st Intern. Citrus Symp., Univ. of Calif., Riverside. p. 855-62. McMurtry,
J. A. 1977a. Some predaceous mites [Phytoseiidae] on citrus in the
Mediterranean region. Entomophaga 22: 19-30. McMurtry,
J. A. 1977b. Acarina, p. 1-8. In:
C. P. Clausen (ed.), Introduced Parasites and Predators of Arthropod Pests
and Weeds: a World Review. Agr. Handbk. No. 480, U.S. Dept. Agr., Agr. Res.
Svc. 551 p. McMurtry,
J. A. 1978. Biological control of citrus mites. In: W. Grierson (ed.), "Proc. Intern. Soc.
Citriculture, Vol. 2: p. 855-62. McMurtry,
J. A. 1982. The use of phytoseiids for biological control: progress and
future prospects, p. 23-48. In:
M. A. Hoy (ed)., Recent Advances in Knowledge of the Phytoseiidae. Univ. of
Calif. Div. Agr. Sci. Publ. 3284, Berkeley, Calif. McMurtry,
J. A. 1983. Phytoseiid predators in orchard systems: A classical biological
control success story, p. 21-26. In:
M. A. Hoy, G. L. Cunningham & L. Knutson (eds.), Biological Control of
Pests by Mites. Univ. of Calif. Special Publ. 3304, Berkeley, CA. McMurtry,
J. A. 1985a. Avocado, p. 327-37. In:
W. Helle & M. W. Sabelis (eds.), Spider Mites, Their Biology, Natural Enemies
and Control. Vol. 1B. Elsevier Sci. Publ., Amsterdam. McMurtry,
J. A. 1985b. Citrus, p. 339-47. In:
W. Helle & M. W. Sabelis (eds.), Spider Mites, Their Biology, Natural
Enemies and Control. Vol. 1B. Elsevier Sci. Publ., Amsterdam. McMurtry,
J. A. 1989. Utilizing natural enemies to control pest mites on citrus and
avocado in California, USA. In:
Proc. VII Intern. Congress Acarol., 1986. McMurtry,
J. A. & H. G. Johnson. 1963. Progress report on the introduction of a
thrips parasite from the West Indies. Calif. Avocado Soc. Yearbk. 47: 48-51. McMurtry,
J. A. & H. G. Johnson. 1966. An ecological study of the spider mite Oligonychus punicae (Hirst) and its natural
enemies. Hilgardia 37: 363-402. McMurtry,
J. A. & G. T. Scriven. 1964a. Studies on the feeding, reproduction and
development of Amblyseius hibisci (Acarina: Phytoseiidae)
on various food substances. Ann. Ent. Soc. Amer. 57: 649-55. McMurtry,
J. A. & G. T. Scriven. 1964b. Biology of the predaceous mite Typhlodromus rickeri (Acarina: Phytoseiidae).
Ann. Ent. Soc. Amer. 57: 362-67. McMurtry,
J. A. & G. T. Scriven. 1965. Insectary production of phytoseiid mites. J.
Econ. Ent. 58: 282-4. McMurtry,
J. A. & G. T. Scriven. 1966. The influence of pollen and prey density on
the number of prey consumed by Amblyseius
hibisci (Chant) (Acarina:
Phytoseiidae). Ann. Ent. Soc. Amer. 59: 147-49. McMurtry,
J. A. & G. T. Scriven. 1968. Studies on predator-prey interactions
between Amblyseius hibisci and Oligonychus punicae: effects of host-plant conditioning
and limited quantities of alternate food. Ann. Ent. Soc. Amer. 61: 393-97. McMurtry,
J. A. & G. T. Scriven. 1975. Population increase of Phytoseiulus persimilis
on different insectary feeding programs. J. Econ. Ent. 68: 319-20. McMurtry,
J. A., H. G. Johnson & G. T. Scriven. 1969. Experiments to determine
effects of mass releases of Stethorus
picipes on the level of
infestation of the avocado brown mite. J. Econ. Ent. 62: 1216-21. McMurtry,
J. A., C. B. Huffaker & M. van de Vrie. 1970. Ecology of tetranychid
mites and their natural enemies: a review. I. Tetranychid enemies: their
biological characters and the impact of spray practices. Hilgardia 40(11):
331-90. McMurtry,
J. A., E. R. Oatman, P. A. Phillips & C. W. Wood. 1978. Establishment of Phytoseiulus persimilis (Acari:
Phytoseiidae) in southern California. Entomophaga 23: 175-179. McMurtry,
J. A., J. G. Shaw & H. G. Johnson. 1979. Citrus red mite populations in
relation to virus disease and predaceous mites in southern California.
Environ. Ent. 8: 160-64. McMurtry,
J. A., H. G. Johnson & M. H. Badii. 1984. Experiments to determine
effects of predator releases on populations of Oligonychus punicae
[Acarina: Tetranychidae] on avocado in California. Entomophaga 29: 11-19. McMurtry,
J. A., H. H. Badii & H. G. Johnson. 1984. The broad mite, Polyphagotarsonemus latus, as a potential prey for
phytoseiid mite in California. Entomophaga 29: 83-6. McMurtry,
J. A., H. G. Johnson & S. N. Newberger. 1991. Greenhouse thrips
parasitoid established on avocado in California. Calif. Agr. (in press). Metcalf,
R. L. & I. M. Newell. 1962. Investigation of the biochromes of mites.
Ann. Ent. Soc. Amer. 55: 350-53. Meyer,
M. K. P. 1962. Two new mite predators of the red scale (Aonidiella aurantii)
in South Africa. S. Afr. J. Agr. Sci. 5: 411-17. Michael,
A. D. 1903. British Tyroglyphidae, Vol. 2. The Ray Soc., London. 183 p. Mori,
H. & D. A. Chant. 1966a. The influence of prey density, relative
humidity, and starvation on the predaceous behavior of Phytoseiulus persimilis
Athias-Henriot (Acarina: Phytoseiidae). Canad. J. Zool. 44: 483-91. Mori,
H. & D. A. Chant. 9166b. The influence of humidity on the activity of Phytoseiulus persimilis Athias-Henriot and
its prey Tetranychus urticae (C. L. Koch) (Acarina:
Phytoseiidae, Tetranychidae). Canad. J. Zool. 44: 683-781. Morton,
F. A. 1928. Quantity production of tick parasites. Mont. St. Bd. Ent. 7th
Bien. Rept. p. 32-5. Muma,
M. H. 1954. Lady beetle predators of citrus mealybugs. Citrus Mag., Apr.
1954: 16-17. Muma,
M. H. 1955. Factors contributing to the natural control of citrus insects and
mites in Florida. J. Econ. Ent. 48: 432-38. Muma,
M. H. 1958. Predators and parasites of citrus mites in Florida. Proc. 10th
Intern. Congr. Ent. 4: 633-47. Muma,
M. H. 1961a. Mites associated with citrus in Florida. Fla. Agr. Expt. Sta.
Bull. 640. 39 p. Muma,
M. H. 1961b. The influence of cover crop cultivation on populations of
indigenous insects and mites in Florida citrus groves. Florida Ent. 44: 61-8. Muma,
M. H. 1964. Annotated list and keys to Phytoseiidae (Acarina: Mesostigmata)
associated with citrus in Florida. Fla. Agr. Expt. Sta. Tech. Bull. 685. 42
p. Muma,
M. H. 1966. Mists vs. mites. Sunshine State Agr. Res. Rept., Gainesville, FL.
11(3): 18-9. Muma,
M. H. 1969. Biological control of various insects and mites on Florida
citrus. In: H. D. Chapman
(ed.), Proc. 1st Intern. Citrus Symp., Riverside, Calif. 2: 863-70. Muma,
M. H. 1970. Natural control potential of Galendromus
floridanus (Acarina:
Phytoseiidae) on Florida citrus trees. Florida Ent. 53: 79-88. Muma,
M. H. 1975. Mites associated with citrus in Florida. Fla. Agr. Expt. Sta.
Bull. 640A. Muma,
M. H. & H. A. Denmark. 1970. Phytoseiidae of Florida. Fla. Dept. Agr.
Con. Serv., Gainesville. 1950 p. Muma,
M. H. & E. Katherine. 1949. Studies on populations of prairie spiders.
Ecology 30: 485-503. Munger,
F. 1963. Factors affecting growth and multiplication of the citrus red mite, Panonychus citri. Ann. Ent. Soc. Amer. 56: 867-74. Newell,
I. M. 1943. A new sironid from North America (Opiliones, Cyphophthalmi,
Sironidae). Trans. Amer. Microsc. Soc. 62(4): 416-22. Newell,
I. M. 1945a. The status of Thalassarachna
verrilli Packard 1871. Halacarus Grosse 1855, and Copidognathus Trouessart 1888
(Acari, Halacaridae). Trans. Amer. Microsc. Soc 64: 58-62. Newell,
I. M. 1945b. Hydrozetes
Berlese: The occurrence of the genus in North America and the phenomenon of
levitation (Acari, Oribatoidea). Trans. Conn. Acad. Arts Sci. 36: 253-75. Newell,
I. M. 1947a. A systematic and ecological study of the Halacaridae of Eastern
North America. Bull. Bingham Oceanogr. Coll. 19(3): 1-233. Newell,
I. M. 1947b. Studies on the morphology and systematics of the family
Halarachnidae Oudemans 1906 (Acari, Parasitoidea). Bull. Bingham Oceanogr.
Colln. 19(4): 234-66. [Contrib. No. 340 of Woods Hole Oceanographic Inst.]. Newell,
I. M. 1947c. Quantitative methods in biological and control studies of
orchard mites. J. Econ. Ent. 40: 683-9. Newell,
I. M. 1947d. The rediscovery and clarification of Siro acaroides (Ewing) 1923
(Opiliones, Cyphophthalmi, Sironidae). Trans. Amer. Microsc. Soc. 66(4):
354-65. Newell,
I. M. 1949. New genera and species of Halacaridae (Acari). Amer. Mus. Novit.
No. 1411. p. 1-22. Newell,
I. M. 1950a. Métodos de recolección de Halacaridae (Acari). Arthropoda
1(2/4): 375-6. Newell,
I. M. 1950b. New species of Copidognathus
from the Aleutians (Acari, Halacaridae). Amer. Mus. Novit. No. 1476: 1-19. Newell,
I. M. 1951a. New species of Agaue
and Thalassarachna from the
North Pacific (Acari, Halacaridae). Amer. Mus. Novit. No. 1489: 1-19. Newell,
I. M. 1951b. Copidognathus curtus Hall 1912 and other
species of Copidognathus
from western North America (Acari, Halacaridae). Amer Mus. Novit. No. 1499:
1-27. Newell,
I. M. 1951c. A comparative study of the mite fauna of the North American
Arctic, with special emphasis on the marine mites. A report to the Arctic
Institute of North America. 28 p. Newell,
I. M. 1951d. Further studies on Alaskan Halacaridae. Amer. Mus. Novit. No.
1536: 1-56. Newell,
I. M. 1953. The natural classification of the Rhombognathinae (Acari,
Halacaridae). Systematic Zool. 2(3): 119-35. Newell,
I. M. 1954. The Halacaridae or marine mites, p. 197-200. In: R. I. Smith et al. (eds.), Intertidal Invertebrates of
the Central California Coast. Univ. Calif. Press. Newell,
I. M. 1955. An autosegregator for use in collecting soil-inhabiting
arthropods. Trans. Amer. Microsc. Soc. 74(4): 389-92. Newell,
I. M. 1956a. The new genus Tetracondyla
in the Pacific (Acari: Oppiidae). Proc. Hawaii. Ent. Soc. 16(1): 113-21. Newell,
I. M. 1956b. A parasitic species of Copidognathus
(Acari: Halacaridae). proc. Hawaii. Ent. Soc. 16(1): 122-25. Newell,
I. M. 1956c. Pachygnathus notops Gosse 1855-- 100 years
later. Annals & Magazine of Nat. Hist., Ser. 12(9): 465-75. Newell,
I. M. 1957a. A new genus and species of Oribatei exhibiting external sexual
dimorphism. proc. Hawaii. Ent. Soc. 16(2): 298-306. Newell,
I. M. 1957b. Studies on the Johnstonianidae (Acari, Parasitengona). pacific
Sci. 11(4): 396-466. Newell,
I. M. 1958. Specific characters and character variants in adult and larvae of
the genus Paratrombium
Bruyant 1910 (Acari, Trombidiidae), with descriptions of two new species from
western North America. Pacific Sci. 12: 350-70. Newell,
I. M. 1959. Acari, p. 1080-1116. In:
W. T. Edmondson (ed.), Fresh Water Biology, Chapt. 42. Ward & Whipple
Publ. Newell,
I. M. 1960. Charadracarus
new genus, Charadracarinae new subfamily (Acari, Johnstonianidae), and the
status of Typhlothrombium
Berlese 1910. Pacific Science 14(2): 156-72. Newell,
I. M. 1963. Feeding habits in the genus Balaustium
(Acarina, Erythraeidae), with special reference to attacks on man. J.
Parasitol. 49: 498-502. Newell,
I. M. 1967a. Prostigmata: Halacaridae (Marine Mites). Reprinted from
Antarctic Research Series Vol. 10, Entomology of Antarctica, by J. Linsley
Gressit (ed.). p. 81-95. Newell,
I. M. 1967b. Abyssal Halacaridae (Acari) from the southeast Pacific. Pacific
Insects 9(4): 693-708. Newell,
I. M. 1970. Construction and use of tabular keys. Pacific Insects 12(1):
25-37. Newell,
I. M. 1971a. Problems in the study of subtidal Halacaridae (Acari), p. 103-7.
In: Proc. 1st Internatl.
Conf on Meiofauna. Smithsonian Contrib. Zool. 76. Newell,
I. M. 1971b. Halacaridae (Acari) collected during Cruse 17 of the R/V ANTON
BRUUN, in the southeastern Pacific Ocean. Scientific Results of the Southeast
Pacific Expedition. ANTON BRUUN Rept. No. 8: 1-58. Newell,
I. M. 1972. Tabular keys: further notes on their construction and use, p.
259-67. In: Growth by
Intussusception: Ecological Essays in Honor of G. Evelyn Hutchinson. Trans.
Conn. Acad. Arts & Sci. 44. Newell,
I. M. & M. Andre. 1959. Revision des especes de Rhombognathus (Halacariens Marins) decrites par Edouard L.
Trouessart. Acarologia 1(1): 124-46. Newell,
I. M. & R. E. Ryckman. 1964. Hirstiella
pyriformis sp. n. (Acari,
Pterygosomidae), a new parasite of lizards from Baja California. J.
Parasitol. 59: 163-71. Newell,
I. M. & R. E. Ryckman. 1966. Species of Pimeliaphilus [Acari:
Pterygosomidae] attacking insects, with particular reference to the species
parasitizing Triatominae [Hemiptera: Reduviidae]. Hilgardia 37(12): 403-36. Newell,
I. M. & R. E. Ryckman. 1969. Pimeliaphilus
zeledoni n. sp. (Acari,
Pterygosomidae), a parasite of Triatoma
dimidiata (Latr.)
(Hemiptera, Reduviidae). Bull. S. Calif. acad. Sci. 68(3): 138-44. Newell,
I. M. & L. Trevis, Jr. 1960. Angelothrombium
pandorae n. g., n. sp.
(Acari, Trombidiidae), and notes on the biology of the giant reduvelvet
mites. Ann. Ent. Soc. Amer. 53: 293-304. Newell,
I. M. & P. H. Vercammen-Grandjean. 1964. Pteridopus n. g. (Acari, Johnstonianidae) and a probable
auditory organ in a mite. Acarologia 6: 98-110. Oatman,
E. R. 1965. Predaceous mite controls two-spotted spider mite on strawberry.
Calif. Agr. 19: 6-7. Oatman,
E. R. & J. A. McMurtry. 1966. Biological control of the two-spotted
spider mite on strawberry in southern California. J. Econ. Ent. 59: 433-39. Oatman,
E. R., J. A. McMurtry, H. H. Shorey & V. Voth. 1967. Studies on
integrating Phytoseiulus persimilis releases, chemical
applications, cultural manipulation, and natural predation for control of the
two-spotted spider mite on strawberry in southern California. J. Econ. Ent.
60: 1344-51. Oldfield,
G. N., I. M. Newell & D. K. Reed. 1972. Insemination of protogynes of Aculus cornutus from spermatophores, and description of the sperm
cell. Ann. Ent. Soc. Amer. 65: 1080-4. Pickett,
A. D. 1965. The influence of spray programs on the fauna of apple orchards in
Nova Scotia. XIV. Supplement to II. Oystershell scale, Lepidosaphes ulmi
(L.). Canad. Ent. 97: 816-21. Pimentel,
D., M. W. Rumsey & F. A. Streams. 1960. Rearing tyroglyphid mites on Neurospora. Ann. Ent. Soc.
Amer. 53: 549. Quayle,
H. J. 1912. Red spiders and mites of citrus trees. Calif. Agr. Expt. Sta.
Bull. 234: 483-530. Richards,
A. M. 1962. The oyster-shell scale Quadraspidiotus
ostreaeformis (Curtis), in
the Christchurch district of New Zealand. New. Zeal. J. Agr. Res. 5: 95-100. Riley,
C. V. 1873. Fifth Annual Report on the Noxious, Beneficial, and other Insects
of the State of Missouri. Regan & Carger, Jefferson City, MO. Rosen,
D. 1967. Biological and integrated control of citrus pests in Israel. J.
Econ. Ent. 60: 1422-27. Samarasinghe,
S. & E. J. LeRoux. 1966. The biology and dynamics of oystershell scale, Lepidosaphes ulmi (L.) (Homoptera:
Coccidae), on apple in Quebec. Ann. Ent. Soc. Quebec 11: 206-92. Schmutterer,
H. 1959. Shildläuse oder Coccoidea. I. Dekelschildläuse oder Diaspididae. Die
Tierwelt Deutschlands. 45 Teil. Gustav Fischer Verlag, Jena. Sellers,
W. F. & G. G. Robinson. 1950. The effect of the miticide Neotran upon the
laboratory production of Aspidiotus
lataniae Signoret as a
coccinellid food. Canad. Ent. 82: 170-73. Shimer,
H. 1868. Notes on the "apple bark-louse" (Lepidosaphes conchiformis
Gmelin sp.) with a description of a supposed new Acarus. Trans. Amer. Ent. Soc. 1: 361-74. Simmonds,
F. J. 1958. The oleander scale, Pseudaulacaspis
pentagona (Targ.)
(Homoptera, Diaspididae) in Bermuda. Bermuda Dept. Agr. Bull. No. 31. Simmonds,
F. J. 1960. Biological control of the coconut scale, Aspidiotus destructor
Sign., in Principe, Portuguese West Africa. Bull. Ent. Res. 51: 223-37. Smirnoff,
W. A. 1957. La cochenille du palmier dattier (Parlatoria blanchardi
Targ.) en Afrique du Nord, comportement, importance économique, prédateurs et
lutte biologique. Entomophaga 2: 1-98. Smirnov,
E. & W. Polejaeff. 1933. Density of population and sterility of the
females in the coccid Lepidosaphes
ulmi L. J. Anim. Ecol. 3:
29-40. Smith,
C. N. & M. M. Cole. 1943. Studies of parasites of the American dog tick.
J. Econ. Ent. 36: 569-72. Smith,
C. N., M. M. Cole & H. K. Gouck. 1946. Biology and control of the
American dog tick. U.S. Dept. Agr. Tech. Bull. 905. 74 p. Smith,
F. F., T. J. Henneberry & A. L. Boswell. 1963. The pesticide tolerance of
Typhlodromus fallacis (Garman) and Phytoseiulus persimilis Athias-Henriot with
some observations on efficiency of P.
persimilis. J. Econ. Ent.
56: 274-78. Stoetzel,
M. B. & J. A. Davidson. 1971. Biology of the obscure scale, Melanaspis obscura (Homoptera: Diaspididae) on pin oak in Maryland.
Ann. Ent. Soc. Amer. 64: 45-50. Summerland,
S. A. & D. W. Hamilton. 1951. Hemisarcoptes
malus, a predator of Forbes
scale. J. Econ. Ent. 44: 818. Swirski,
E. & R. Schlechter. 1961. Some phytoseiid mites (Acarina: Phytoseiidae)
of Hong Kong, with a description of a new genus and seven new species. Israel
J. Agr. Res. 11: 97-117. Swirski,
E. & S. Amitai. 1961. Some phytoseiid mites (Acarina: Phytoseiidae) of
israel, with a description of two new species. Israel J. Agr. Res. 11:
193-202. Tanigoshi,
L. K. 1983. Advances in knowledge of the biology of the Phytoseiidae, p.
1-22. In: M. A. Hoy (ed.),
Recent Advances in Knowledge of the Phytoseiidae. Univ. Calif. Div. Agr.
Sci., Publ. 3284. p. 1-22. Tevis,
L., Jr. & I. M. Newell. 1962. Studies on the biology and seasonal cycle
of the giant red velvet mite, Dinothrombium
pandorae (Acari,
Trombidiidae). Ecology 43: 497-505. Thomas,
H. A. 1961. Vidia (Coleovidia) cooremani, new subgenus and new
species, and notes on the life history (Acarina: Saproglyphidae). Ann. Ent.
Soc. Amer. 54: 461-63. Turnbull,
A. L. & D. A. Chant. 1961. The practice and theory of biological control
of insects in Canada. Canad. J. Zool. 39: 697-753. Vaivanijkul,
P. & F. H. Haramoto. 1969. The biology of Pyemotes boylei
Krczal (Acarina: Pyemotidae). Proc. Hawaii. Ent. Soc. 20: 443-54. van
de Vrie, M. 1962. The influence of spray chemicals on predatory and phytophagous
mites on apple trees in laboratory and field trials in the Netherlands.
Entomophaga 7: 243-. van
de Vrie, M. 1964a. The distribution of phytophagous and predacious mites on
leaves and shoots of apple trees. Entomophaga 9: 233-8. van
de Vrie, M. 1964b. The effect of an experimental spray schedule on the
population of Metatetranychus
ulmi Koch and Typhlodromus pyri Scheuten. Entomophaga 9:
243-6. van
de Vrie, M. 1965. Problems and prospects in the integrated control of
phytophagous mites. Boll. Zool. Agr., Bachicolt., Milano, Ser. II. p. 275-83. van
de Vrie, M. 1966. Population sampling for integrated control. Proc. FAO Symp.
Integr. Pest Contr., Rome, 1965 2: 57-75. van
de Vrie, M. 1967a. Population sample counts on spider mites. Entomophaga,
Mém. Hors Sér. 3: 55. van
de Vrie, M. 1967b. The effect of some pesticides on the predatory bugs Anthocoris nemorum L. and Orius
spec. and on the woolly aphid parasite Aphelinus
mali Hald. Entomophaga, Mém.
Hors Sér. 3: 95-101. van
de Vrie, M. 1986. Apple. Chapter 3.2.4. p. 311-25. In: W. Helle & M. W. Sabelis (eds.), "Spider
Mites: Their Biology, Natural Enemies and Control. Vol. 1B. Elsevier,
Amsterdam. 458 p. van
de Vrie, M. & A. Boersma. 1970. The influence of the predaceous mite Typhlodromus potentillae (Garman) on the
development of Panonychus ulmi (Koch) on apple grown
under various nitrogen conditions. Entomophaga 15: 291-304. van
de Vrie, M. & D. Kropczynska. 1965. The influence of predatory mites on
the population development of Panonychus
ulmi (Koch) on apple. Boll.
Zool. Agr. Bachicolt. Ser. II7: 119-30. van
de Vrie, M. & D. Kropczynska. 1967. The influence of predatory mites on
the population development of Panonychus
ulmi Koch on apple.
Entomophaga, Mém. Hors Sér. 3: 77-84. van
de Vrie, M. & C. A. van den Anker. 1967. The Stuttgart funnel method to
estimate the effect of pesticides on the arthropod fauna of fruit trees.
Entomophaga, Mém. Hors Sér. 3: 21-4. van
de Vrie, M., J. A. McMurtry & C. B. Huffaker. 1972. Ecology of
tetranychid mites and their natural enemies: a review. III. Biology, ecology
and pest status, and host plant relations of tetranychids. Hilgardia 41:
343-432. Wafa,
A. K., M. A. Zaher & Z. R. Soliman. 1970. Life-history of the predator
mite Eutogenes africanus Wafa and Soliman
(Acarina: Cheyletidae). Bull. Soc. Ent. d'Egypte 54: 129-31. Watson,
T. F. 1964. Influence of host plant condition on population increase of Tetranychus telarius (Linnaeus) (Acarina:
Tetranychidae). Hilgardia 35: 273-322. Williams,
J. R. 1970. Studies on the biology, ecology and economic importance of the
sugar-cane scale insect, Aulacaspis
tegalensis (Zhnt.)
(Diaspididae), in Mauritius. Bull. Ent. Res. 60: 61-95. Willoughby,
P. A. & M. Kosztarab. 1974. Studies on the morphology and systematics of
scale insects. No. 7. Morphological and biological studies on two species of Chionaspis (Homoptera:
Coccoidea: Diaspididae). Research Div. #92, Virginia Poly. Inst. & St.
Univ., Blacksburg, VI. Wood,
H. P. 1911. Notes on the life history of the tick parasite, Hunterellus hookeri How. J. Econ. Ent. 4:
425-31. Wright,
K. A. & I. M. Newell. 1964. Some observations on the fine structure of
the midgut of the mite Anystis
sp. Ann. Ent. Soc. Amer. 57: 684-93. Zaher,
M. A. & Z. R. Soliman. 1971. Life-history of the predator mite, Cheletogenes ornatus (Canestrini and
Fanzago). Bull. Soc. Ent. d'Egypte 55: 85-9. |