Aust. J. Zool.

, 1974, 22, 505-17

The Introduction of Bdellodes lapidaria

(Acari :Bdellidae) from Australia into
South Africa for the Biological Control of
Sminthurus viridis (Collembola)

M. M. H. WallaceAand M. C.' WaltersB

A Division of Entomology, CSIRO, Private Bag, P.O., Wembley, W.A. 6014.
Department of Agricultural Technical Services, Winter Rainfall Region, Stellenbosch, South Africa.

Abstract
The lucerne flea, S, viridis, was introduced into South Africa with subterranean clover seed imported
from Western Australia in 1939. By 1959 the infestation had spread over 50,000 ha and methods of
control were examined.
The predatory mite B. lapidaria was known to be an effective biological control agent against
S. viridis in Australian pastures. From 1963 to 1966 over 75,000 mites were collected in Western
Australia and sent by air for immediate release in selected sites in South Africa (Cape Province).
The mites were shown to be established in 1965 and multiplied rapidly, enabling redistribution to
additional sites.
The influence of this predator on S. viridis numbers is now being monitored. Preliminary data
indicate that B. lapidaria is capable of influencing flea densities under South African conditions.
A definite relationship between the predator and sminthurid Collembola such as S. viridis slnd
Bourletiella arvalis is indicated, although B. lapidaria appears to be able to utilize other foods during
periods of sminthurid scarcity.

Introduction
The lucerne flea, Sminthurus viridis (L.) (Collembola), was first collected in the
Cape Province of South Africa in 1941 near Somerset West. At the time it was not
realized that the pest was, in fact, S. viridis, and it was not until 1954, when damage
to lucerne near Caledon was noted, that its identity was suspected. Even so a positive
identification was delayed until 1957. In the meantime the pest had spread rapidly
and by 1959 Walters (1968) estimated the total area infested at about 50,000 ha and
new infestations were continually being discovered.
It was at that time that work in Western Australia was beginning to reveal the
full effect of the predatory mite, Bdellodes lapidaria Kramer (Bdellidae), on numbers
of S. viridis in pastures (Wallace 1967). In the light of that knowledge the South
African Department of Agriculture decided to look into the possibility of importing
B. lapidaria into South Africa to aid in the biological control of S. viridis there.
This joint project, begun in 1963, and resulting in the successful establishment
of the predator in South Africa, is described in this paper.

Collection of Bdellodes lapidaria in Western Australia

Collection Sites
B. lapidaria is widespread in the south-western corner of Western Australia
(Wallace and Mahon 1971) but, like its principal host, S. viridis, it is very patchy,
 M. M. H. Wallace and M. C. Walters

and high-density populations are local and sporadic, and often of short duration.
To locate areas suitable for collecting large numbers it was necessary to carry out a
detailed survey immediately prior to the planned collecting date.
Suitable collecting areas convenient to Perth were available about 100 km to the
east near Toodyay, Northam and York, on the western boundary of the wool-growing
areas. High-density populations of both S. viridis and B. lapidaria were found there
without difficulty. Climatically the collection site and the proposed release sites near
Stellenbosch and Caledon in Cape Province are similar, both being basically Mediter-
ranean in character with cool moist winters and warm dry summers (Fig. 1). The
average growing season in each extends from about May through to October. Thus
the mites could be expected to be well adapted for survival at the release sites in South
Africa.
Caledon site Stellenbosch site Northam

Fig. 1. Climate diagrams of the collection site of Bdellodes lapidavia in Western Australia (Northam)
and of the release sites in South Africa (Caledon and Stellenbosch).

Methods of Collection
Two methods of collection were adopted in the field. The first attempted to take advantage of the
mites' tendency to shelter under pieces of stick, bark, etc. Many pieces of rough board cut from old
packing cases were scattered over the pasture in the selected area. They were picked up carefully
15-20 min later and mites clinging to them were knocked on to enamel plates from which they were
sucked up into small plastic tubes with a mouth-operated aspirator.
The second method consisted of sweeping through the pasture with an enamel plate about 25 cm
in diameter, and sucking the mites from this into tubes as before. In general, this method yielded
greater numbers of mites, especially in low-density populations, but it had the disadvantage that
collections initially contained large numbers of unwanted insects and mites, from which B. lapidaria
had to be sorted.
Late in 1966, a third method of collection was tried, by means of a power-operated vacuum-type
machine developed from air-conditioning equipment commonly used in large buildings. This proved
highly successful and led to the development of the portable equipment later used in Morocco and
France for the collection of bdellid mites and described fully by Wallace (1972).
The maximum hourly collection rate achieved by any method was about 2000 mites per person
in a densely populated area. An average figure would be closer to 500 per person hourly.

Methods of Sorting and Shipment

In 1963 and 1964 the methods adopted were as follows:
On return to the laboratory a black sleeve was placed over the field collection tube and the active
mites were induced to move into a clean tube by the open end being turned towards the light of a
window; the clean tube was then put into a jar of carbon dioxide until the mites became inactive.
They were then tipped into a shallow examination tray into which there was a steady flow of carbon
dioxide. Under the dissecting microscope all unwanted insects, mites and plant material were
removed, and identification of the bdellid mites was checked. The mites were then tipped into a
larger container with a gauze bottom, standing on a wire frame, which allowed the carbon dioxide
to flow out. When the mites had fully recovered they were transferred to the shipment cages.
Bdellodes lapidavia in South Africa

These cages were made from plastic sample tubes, 10 by 3 cm diameter. A 2-cm hole was drilled
in the bottom of each tube and in each cap. A dry plug of peat moss covered with cellophane was
rammed into the bottom of the cage, and a piece of rough cardboard placed in the cage to provide
extra surface area and oviposition sites. Between 100 and 120 mites were then placed in each cage
and a plug of peat moss moistened in distilled water and covered with cellophane was pushed into the
top of the cage and held in place by the screw cap.
For shipment the cages were put into cardboard boxes surrounded by loosely packed, crumpled
newspaper, and taken to the airport immediately before departure of the aircraft.
Owing to the poor survival of mites in these shipments, especially in 1964, the methods were
changed in 1965 in the light of experience gained in the shipment of mites from Europe to Australia.
The high mortalities seemed to be due mainly to the carbon dioxide treatment during sorting and
poorly aerated cages with too much water condensation. Accordingly, in 1965 and 1966, once the
field collection was in the laboratory, the contents of the tubes were tipped on to the centre of a large
sheet of white plastic spread out on a flat table. The active bdellid mites quickly ran out from the
small pile of debris in the centre of the sheet and as they moved on to clean areas they were sucked up
individually into the despatch cages. These were then examined with a lens and any unwanted
organisms sucked accidentally into the cages were removed.

Fig. 2. Cage for transporting living mites by air.

The shipment cages were again made from plastic sample tubes (Fig. 2) but the moist plug now
consisted of a wad of saturated facial tissue pressed into an inside-fitting cap. A piece of cellophane
was placed over the cap before it was pressed into place. A strip of dry tissue stretched over the
cellophane provided a suitable oviposition site. At the other end of the cage a piece of tissue was
placed over the perforated cap before it too was pressed into place. The cage thus had a moist end
and a well-aerated dry end.
These improved methods of sorting and shipment proved highly successful, as shown by the
survival figures from 4 September 1965 onwards, in Table 1.

Liberation of Bdellodes lapidavin in South Africa

Introduction Sites
The first releases of B. lapidaria in South Africa were carried out at Caledon during
1963 and 1964. They did not result in successful establishment of the species. Sub-
sequent releases at Stellenbosch proved successful and were followed by establishment
of the species at Somerset West and Caledon through both imported mites and the
progeny of the Stellenbosch population. The release sites employed, described below,
are indicated in Fig. 3 which also shows the distribution of S. viridis in South Africa
 M. M. H. Wallace and M. C. Walters

as at the end of 1970. The sites at which the various consignments were released are
indicated in Table 1.
Caledon (Site A). This site had previously been employed in a study of the seasonal
history of the lucerne flea, conducted during 1961 and 1962 (Walters 1968), and was
selected on the grounds that prior knowledge of seasonal fluctuations in lucerne flea
numbers would allow more precise evaluation of the influence of the imported preda-
tors. The soil was a loam with pH 6.3; the pasture consisted of a mixture of lucerne,
subterranean clover and various grass species. The site consisted of three separate
paddocks, each 45 by 90 m, exposed to different grazing practices, viz. ungrazed,
normally grazed and heavily grazed by sheep as described by Walters (1968).

Table 1. Details of collections of Bdellodes lapidaria sent to South Africa from 1963 to 1966
-
-

Date Number Appro~. Viability Number of Site

of sent survival eggs laid of
despatch ( %) in transitA release

High 2500 A, Caledon

Low 950 A, Caledon
Low 80 A, Caledon
Low 850 A, Caledon
Low 260 A, Caledon
Low 1260 A, Caledon
Modelrate 1520 A, Caledon
High 1950 C, Stellenbosch
High 6000 C, Stellenbosch
High 4300 C, Stellenbosch
High 2400 B, Caledon
High 2900 D, Somerset West
High 5100 B, Caledon

Total 78,210 30,070

A Estimate.

Each paddock was divided into halves isolated from each other and from the
adjoining paddocks by I-m strips of bare soil soaked periodically with old motor oil
and sprayed with herbicides. The imported bdellids were released on one half of each
paddock, the other half acting as a control plot.
Caledon (Site B). The failure to establish B, lapidaria at site A during 1963 and
1964, the successful establishment of the predator at Stellenbosch during 1965, and
the sale of the area containing site A to new owners led to an alternative release site
being selected at Caledon for use during 1966. Site B was employed for the release
of bdellid mites received from Australia during 1966 and of the progeny of the
population established at Stellenbosch during 1965. It was situated approximately
1 km from site A in a paddock of 40 ha. The soil was a loam with pH 6.1 ; the
pasture was established in 1964 and consisted of lucerne, subterranean clover (a
mixture of Clare, Mount Barker, Geraldton and Dwalganup cultivars), Festuca
arundinacea (Kentucky 31), Lolium rigidum (Wimmera ryegrass), L. multijorum
westerwoldicum and L. perenne (Ariki rye), and Erodium moschatum. The release site
consisted of an area 14 m square isolated from the surrounding pasture by a 1-m wide
vegetation-free border strip, and a fence which was removed the following year.
Bdellodes lapidaria in South Africa

Stellenbosch (Site C). All bdellid consignments received from Australia during
1965 were released in a small (0.2 ha) paddock on the Welgevallen Experimental
Farm at Stellenbosch.
The pasture consisted of a mixture of lucerne and white clover, Trifolium repens,
which had been established during 1964. The paddock had previously been planted
to kikuyu grass, Pennisetum clandestinum, and this grass dominated the pasture by
the end of 1967. The paddock was irrigated during the summer preceding release
but this was discontinued during the 1965-66 summer as it was feared that it might
prove detrimental to the survival of aestivating eggs of B. lapidaria. Irrigation was,

Fig. 3. Map of the Western Cape Province showing the distribution of Sminthurus viridis
in 1970 (stippling) and the sites at which Bdellodes lapidaria were released (e).

however, resumed at the end of the 1967 season as it was found that adjacent
irrigated paddocks colonized by B. lapidaria in 1966 contained active mites throughout
the 1966-67 summer. The cessation of irrigation during 1965 caused the disappearance
of the white clover component of the pasture during the ensuing summer. This was
replaced by lucerne and subterranean clover which was hand-sown in rows during the
autumn of 1966. The soil at this site was a sandy loam with p H 5.2.
All consignments were released at the central point of the paddock and the central
14-m-square area was fenced. The entire paddock was surrounded by a I-m border
strip.
Lucerne flea numbers were maintained at a high level on site C during the 1965
rainy season by means of weekly mass releases of fleas collected elsewhere. The aim
 M. M. H. Wallace and M. C . Walters

of these was to provide optimum conditions for further population increase of B.

lapidaria. They were discontinued during 1966 when it became apparent that another
sminthurid, Bourletiella arvalis Fitch, which occurred throughout the year in greater
numbers than S . viridis on irrigated pastures at Welgevallen, was an' acceptable
alternative prey to B. lapidaria. During the winter the prey numbers were further
augmented by the seasonal appearance of Sminthurinus terrestris Womersley, also
readily eaten by B. lapidaria.
Somerset West (Site D). This site was situated on the farm 'Vergelegen' near
Somerset West. The climate was very similar to that at the Stellenbosch site with
regard to both temperature and rainfall. The soil was a sandy loam with pH 6.0;
the vegetation was dominated by lucerne with Phalaris tuberosa as the only other
perennial of importance. The paddock, of 15 ha, was irrigated during the summer
months. Annual weeds such as Erodium and Hordeum murinum were present during
the colder months when lucerne growth was much retarded.
An area 14 m square was fenced and isolated from the rest of the paddock by a
1-m-wide border strip. There were large numbers of S. viridis at the time of release,
and B. arvalis and S. terrestris were also present.
Liberation Pvoceduves
Prior arrangements with customs and phytosanitary officials allowed rapid clearance of the
consignments on arrival, and liberation of the mites took place within 48-72 h of their departure from
Perth.
The consignments were unpacked in a quarantine room reserved for this purpose. After close
examination the packing material was sealed in plastic bags and burnt. The transparent sample tubes
were carefully examined by two persons for the presence of unwanted organisms before being packed
into new containers and transported to the release site. Screening of the collected material in Australia
proved so effective that no exotic organisms were detected in any of the mite consignments received
in South Africa.
An abundant supply of prey at the release site was ensured by prior collection of large numbers
of lucerne fleas which were released immediately before the bdellid mites. This was intended to
encourage initial concentration of the mites at the point of release and thus the establishment of a
breeding population.
The mites in each sample tube were shaken onto a sheet of white paper by removal of the upper
stopper and inversion of the tube which was then lightly rapped with a stick in order to dislodge them.
The survival rate of the mites in each tube was estimated by counting the dead or moribund indi-
viduals on the paper and remaining in the tube. These results are incorporated in Table 1.
The viability of the mites was estimated by shaking them onto a second sheet of paper containing
large numbers of S. vividis. Immediate and general feeding was regarded as indicating a high degree
of viability; more restricted feeding activity was regarded as indicating moderate viability while a
general failure to commence feeding immediately indicated low viability (Table 1).
A feature of several of the consignments received was the large numbers of eggs laid by the mites
while en route to South Africa. The pieces of cardboard or strips of tissue on which the eggs were
usually laid in the shipment cages were removed from the cages at the time of release and placed in
sheltered positions under small pieces of plank and bark provided as shelter for the bdellids at the
point of release. The estimated numbers of eggs received in each consignment are shown in Table 1.

Post-release Observations
Prey specijkity of Bdellodes lapidaria. Field observation at release sites indicated
that Collembola other than S . viridis were utilized by B. lapidaria. The sminthurids
B. arvalis and S. terrestris were readily fed upon in the field; the former species, in
fact, was the main prey species utilized at the Stellenbosch release site. Predation on
Entomobrya nivalis (L.) was also observed in the field. When confined in petri dishes
Bdellodes lapidaria in South Africa

in the laboratory B. lapidaria would also feed upon Isotoma sp., but would not attack
the podurids Hypogastrura armata (Nicolet), H. manubrialis (Tullb.) or Brachystomella
parvula (Schaffer).
Secondary release sites. During the first few seasons after establishment, the only
release site at which rapid increase of B. lapidaria allowed the collection of large
numbers of mites for release elsewhere was site C at Stellenbosch. The mites were
collected by hand, using soft camel-hair brushes, and were transported to the release
site on the day they were collected, so as to ensure maximum viability on release.
They were all released at a single point and adequate food was assured by the simul-
taneous release of large numbers of lucerne fleas. No barriers were provided. All
the secondary release sites utilized were situated in dryland pastures consisting of
lucerne, subterranean clover and various perennial pasture grasses.
The first release of locally collected predators took place on 30 July 1966, when
800 mites were released at site B. Subsequent releases of imported mites at this site
made it impossible to determine whether the locally collected mites had been successful
in establishing themselves or not.
Further collections of B. lapidaria at site C during 1967 and 1968 allowed populations
to be established at two new sites at Caledon. A total of approximately 800 mites
was released at each site during the course of the two seasons and resulted in per-
manent establishment in both cases.
Monitoring of Predator and Prey Populations
Regular quantitative surveys were commenced at site A in 1964, but ceased in 1966
when it became clear that B. lapidaria had not been established.
Sites B and C were inspected frequently from 1966 and quantitative surveys were
recommenced in 1970 when it became evident that B. lapidaria was firmly established.
Monitoring of site D was restricted to frequent inspection and indicated successful
establishment.

Results
Caledon (Site A)
The survey method described by Walters (1968) was employed for this purpose,
and population densities were measured every 4 weeks. On each date 10 sample
units were taken at random from the central 100 sq m of each plot. The results of
these surveys reflected the same influence of grazing on lucerne flea numbers as was
noted during 1961 and 1962 (Walters 1968). No B. lapidaria were found in 1964 or
subsequent years.
StelIenbosch (Site C )
The first indication of successful establishment was the discovery, on 12 January
1966, of viable eggs on pieces of plank at the release site. Active mites appeared
during April 1966 and increased so rapidly in numbers that it proved possible to
collect 800 mites for release at Caledon on 30 July 1966.
B. lapidaria spread beyond the confines of site C during 1966. Active mites were found
on the neighbouring irrigated paddocks throughout the following and subsequent dry
seasons, which indicated that moist conditions during summer were not as detrimental
to their chances of survival as had previously been feared.
 M. M. H. Wallace and M. C . Walters

When monitoring was recommenced in January 1970, B. lapidaria had extended

its distribution over an area of approximately 15 ha of the experimental farm around
site C. The release paddock was not regarded as suitable for monitoring, due to its
small size and the complete dominance of the plant cover by P. clandestinum. A
paddock of 0 . 5 ha, 300 m from the release site, was then used. The soil was a sandy
loam with pH 4.8. The vegetation cover consisted of Trijiolium repens (ladino),
lucerne and various grasses including Phalaris tuberosum, Paspalum dilatatum,
Lolium perenne, Setaria nigrirostris and Pennisetum clandestinum. The last subsequently
grew rapidly and by January 1972 was the dominant component of the pasture.
The paddock was irrigated throughout the dry season at a rate of 38 mm of water
per week and was grazed by sheep.

Fig. 4. Changes in the relative population densities of sminthurids,

chiefly Bourletiella arvalis -( ) and bdellids, chiefly Bdellodes
lapidaria (. .... ), at site C from January 1970 to January 1972.

The main bdellid species present on the monitored paddock was B, lapidaria.
Two other species, Neonzolgus capillatus (introduced from Morocco during 1969)
and Cyta latirostris, also occurred in the samples but in such low numbers that their
effect on the Collembola was probably insignificant. The chief prey species was
B. arvalis, as the lucerne flea occurred only in small numbers, perhaps because of the
influence of summer irrigation and high temperatures on the viability of diapause eggs
or survival of larvae (Wallace 1968). B. arvalis was the most common sminthurid on
irrigated Trfolium repens pastures in the Western Cape and appeared to be well
adapted to conditions in such situations. During the winter months S. terrestris was
present in large numbers, considerably augmenting the food supply of the predators.
E. nivalis was present throughout the year and could have provided readily available
alternative prey during periods of sminthurid scarcity.
The monitoring of bdellid and Collembola populations was conducted with the
aid of a portable power-driven collecting apparatus similar to that described by
Wallace (1972). Forty permanent sampling points, arranged in four parallel rows of
10 points each, were situated at the intersections of a 10-m grid drawn over the central
Bdellodes lapidaria in South Africa

0 . 4 ha of the paddock. At each point 11 samples were collected every 4 weeks

commencing on 20 January 1970 (Fig. 4).
A trend for low prey densities to follow periods of high bdellid numbers and high
prey densities to follow low bdellid numbers is evident. The specific interaction
between numbers of B. arvalis and of B. lapidaria is illustrated in Fig. 5, where high
densities (> 100 per sq m) of the prey were recorded only when the predator numbers
4 weeks previously were less than 2 per sq m.
There is also a strong indication in Fig. 4 of a response by the predator to high
prey densities. Thus a high density of Sminthuridae in March 1970 was followed 4
weeks later by a peak of bdellid numbers. Similar responses, but delayed for up to
12 weeks, were noted following high prey densities in October 1970 and October-
December 1971. Higher temperatures experienced in March-April than in the late
spring may have brought about the more rapid response at that time.
r
Fig. 5 Fig. 6

400r

No. o f Bdellodes lapidaria per sq m No. o f Bdellodes lapidaria per sq m

Fig. 5. Relation between numbers of Bourletiella arvalis and Bdellodes lapidaria at site C from
January 1970 to January 1972 inclusive. B. arvalis numbers are those recorded 4 weeks later than the
corresponding numbers of B. lapidaria.
Fig. 6. Relation between numbers of Bourletiella arvalis and numbers of Bdellodes lapidaria at 40
different sampling points on site C on 22 April 1970.

A closer study of the data collected on each sampling date also showed an inter-
action between prey numbers and predator numbers in space on the same date. On
certain sampling dates high prey numbers at sampling points tended to coincide with
low predator numbers at those same points and vice versa. On analysis of log,,, and
log,,, transformed data, significant negative regressions of B. arvalis numbers on
numbers of B. lapidaria were found to occur on 23 February 1970 (*), 23 March 1970
(**), 20 April 1970 (**), and 8 September 1970 (**). These relationships, as they
obtained on 20 April 1970, are illustrated in Fig. 6, where each point on the graph
represents a sampling point at the monitoring site. This provides further evidence of an
influence of B. lapidaria on B. arvalis numbers.

Caledon (Site B)
The B. lapidaria population which had been established at site B during 1966 was
regularly inspected during the period 1966-69. The low numbers of bdellids led
to reluctance to disturb the situation and perhaps decrease the chances of survival
 M. M. H. Wallace and M. C. Walters

of the population. During late 1969 individuals of B. lapidaria were found over 200 m
from the point of release, which indicated that the population had become established
over a wide area; it was decided to commence more intensive monitoring of predator
and pest populations during 1970.
Regular surveys of lucerne flea and bdellid populations were conducted at site B
and in two other paddocks (N, and N,) approximately 1 .5 km from site B. Site N, was
used for the release of another imported bdellid, Neomolgus capillatus from Morocco
and southern France, during 1969, while site N, was intended as a control to both
the others. The N. capillatus population at site N, showed little sign of activity
during 1970, 1971 or 1972, only 4, 25 and 13 bdellids being recovered during the
respective years, so that their effect on S. viridis numbers was probably negligible.
Hence both sites N, and N, can be regarded as controls for population densities
of B. lapidaria and lucerne flea at site B.

-
6
Z
- 50

- 40
- 30
Fig. 7. Changes in numbers of
- 20 Sminthurus viridis -( ) and
Bdellodes lapidaria (. .. . .
) at site B in

Site B Site N1
I,,{y
,J ,A J
Site N2
,\,O
1970-71, compared with those of
two neighbouring sites, N, and N,.

The plots were sampled at 4-week intervals, by the same method as that employed
at Stellenbosch during 1970 and 1971 (Fig. 7).
S. viridis and B, lapidaria first appeared in response to heavy rains (52-3 mm)
during late February 1970. When a sample was taken 3 weeks later the active stages
of both flea and predator had disappeared in response to unfavourably warm and dry
conditions. Further rains in May brought about a second hatching of eggs and the
Bdellodes lapidaria in South Africa

population of active stages then continued through until late November. In 1971
S. viridis was continuously present from 26 April to 8 November, and in 1972 from
24 May to 6 November.
The lucerne flea population curves for sites N, and N, differed little in form during
1970-1972. The differences in peak densities recorded on N, (509) and N, (100)
during 1970 may have been influenced by an anystid mite, Chaussieria sp., which
occurred at N, in exceptionally high numbers in 1970, and was almost absent during
1971 and 1972.
The lucerne flea density curve for site B during 1970 was basically similar to those
for N, and N,, with maximum numbers attained later in the season. Numbers
decreased rapidly at all three sites after October, with the advent of drier conditions.
An increase in the number of B. lapidaria at site B occurred between 12 October and
9 November 1970, providing a good supply of oversummering eggs and a rapid
increase in B. lapidaria numbers the following year. Such late increases in numbers of
B. lapidaria occurred on a larger scale during 1971 and 1972 and would appear to be
typical of the ecology of the species in this locality. Similar trends were noted in
Western Australia by Wallace (1967).
A comparison of the lucerne flea density curve for 1971 in site B with those for sites
N, and N, reveals that maximum flea numbers occurred earlier in site B than in the
other two sites. It is probable that the earlier decline in site B was due to predation by
B. lapidaria, since large numbers of mites were recorded in the early winter (June)
when they would be able to exert their maximum influence. Wallace (1967) showed
clearly that B. lapidaria was able to affect lucerne flea numbers substantially only when
the predator was sufficiently abundant early in the season (before the end of June).
Although large numbers of B. lapidaria appeared late in the 1972 season, they were
too late to have any substantial effect on the population density curve of S. viridis.
Only small numbers of predators were recorded early in the season; they were unable
to exert any observable influence on S. viridis numbers. In Western Australia, in
districts where B. lapidaria is established, the lucerne flea reaches pest densities in
local areas only once in every three or four years. A similar sequence of events is
probably now being initiated in the South African site.
The continued increase in B. lapidaria numbers at site B during 1971 and 1972,
at a stage when more predators than lucerne fleas were being collected, may have been
due to delayed hatching of eggs and good survival of adults, assisted by alternative
prey in the form of entomobryid Collembola. For example on 12 October 1971, 223
B. lapidaria were collected in company with only 91 S. viridis and a total of 801
entomobryids. On 6 November 1972, when 345 B. lapidaria were collected, the only
Collembola recorded were 858 entomobryids. Wallace (1967) noted an ability of
B. lapidaria to maintain low densities under Western Australian conditions when
S. viridis was present in only very small numbers. The presence of alternative foods
was regarded as a possible explanation for the phenomenon. This ability of B.
lapidaria to persist at relatively high densities in the relative scarcity of its preferred
prey is important in determining its efficacy as a biological control agent.

Discussion
Predatory mites have not been widely used in biological control programmes against
insect and mite pests, but, nevertheless, a number of highly successful projects have
 M. M. H. Wallace and M. C. Walters

been undertaken. Phytoseiidae have been used in experiments on the control of red
spider mites in glasshouses (Bravenboer 1969), in orchards (Muma 1969), and on
melons where good commercial control was achieved (Gould et al. 1969). Macro-
chelidae are being used in attempts for biological control of flies in animal manure
(Axtell 1969), while perhaps the most successful experiments were those described by
Huffaker and Kennett (1956), who employed phytoseiid mites for the control of the
cyclamen mite in strawberries. The small size of predatory mites places certain limits
on their use as biological control agents, so that they would normally be restricted to
preying upon phytophagous microarthropods, as in the majority of cases cited above,
or to the egg and early immature stages of larger pests.
As indicated by Doutt and DeBach (1964), there is a general belief amongst
biological control workers that specific parasites and predators are more likely to be
successful in biological control than more general feeders. A natural predator
specific to a given prey is inherently closely attuned to the prey species and thus highly
responsive to prey density changes. On the other hand, any outside agency which
tends to depress prey numbers may have a serious adverse effect on the specific
predator and thus drastically upset the delicate balance. A more general predator may
be able to maintain itself on other prey species during adverse periods, whilst still
maintaining a useful biological control of the main prey species. Wallace (1967)
showed that B. lapidaria belonged in the latter category. It responded effectively to
changes in the density of its principal prey, S. viridis, but was able to maintain its
numbers at a low level on other prey when S. viridis densities were excessively low, as
happened periodically through a combination of predatory action and adverse
weather. The present study indicates a wide range of prey species within the Collem-
bola as being acceptable to B. lapidaria. A preference for Sminthuridae as prey was
apparent, although prey species from without this group apparently served to tide
the predator over periods of sminthurid scarcity. The apparent preference for S.
viridis, which was observed in Australia by Wallace, and which occurs in dryland
pastures in South Africa, may have been due to the fact that S. viridis was numerically
by far the most dominant sminthurid in such situations. In situations where other
sminthurid species, such as B. arcalis, are numerically dominant, for example in
irrigated pastures at Stellenbosch, the interaction between B. lapidaria and S. viridis
now appears to be replaced by one between B. lapidaria and the dominant sminthurid.
This vermtility of B. lapidaria enhances its value, as it can exert a controlling influence
not only on S. viridis, but also on other phytopathogous sminthurids. B. arvalis and
S. terrestris, although not major pests in South Africa, are nevertheless often trouble-
some due to their attacks on newly emerged seedlings of a wide variety of plants,
and cause limited damage to pastures.
The ability of B. lapidaria to survive on a variety of hosts probably increases the
chances of it being successfully introduced into new localities and countries. The
high densities of B. arvalis at Stellenbosch must undoubtedly have helped to promote
the establishment of the species in South Africa.
.
DeBach (1964) in his introductory chapter noted that ' . . the end result of an
outstanding example of biological control is not spectacular and is likely to go
unnoticed and unappreciated because the formerly abundant organism has been
reduced to a rare species which is attacked by rare natural enemies'. The control
exerted by B. lapidaria on S. viridis in Western Australia is almost in that category,
Bdellodes lapidaria in South Africa

since the effect of the predator is to limit the pest, in areas where it has the potential
for annual outbreaks, to minor and local outbreaks every 3-5 yr (Wallace 1967).
Even so, local outbreaks can periodically reach levels where treatment with insecticides
may be necessary, but only on a very minor scale.
The situation in South Africa prior to the introduction of B. lapidaria was that
annual outbreaks of S. viridis on a relatively large scale were becoming more wide-
spread each year. Although B. lapidaria will probably have little influence on the rate
of spread of S. viridis, the project will be deemed successful if the annual outbreaks
are reduced to local occurrences only.

Acknowledgments
Assistance with the collection, sorting and shipment of mites in Western Australia
was provided by Messrs J. A. Mahon, E. Holm, G. S. McCutcheon and W. J. Penstone,
Division of Entomology, CSIRO. Assistance with the liberation and monitoring
programme in South Africa was provided by Mr E. P. Volschenk, Department of
Agricultural Technical Services, Winter Rainfall Region, Stellenbosch, South Africa.
We are grateful to many colleagues both in Australia and in South Africa for advice
on the project and for criticism of the manuscript.

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Manuscript received 8 May 1974