Use of predatory mites in commercial

biocontrol: current status and future

prospects
Markus Knappa , Yvonne van Houtena , Elmer van Baala , Thomas Groota
a
Koppert Biological Systems, P.O. Box 155, 2650 AD Berkel en Rodenrijs, the Netherlands.

ABSTRACT
Predatory mites play the leading role in commercial augmentative biological control.
They are mainly used in protected vegetable and ornamental cultivation systems to control
phytophagous mites, thrips and whiteflies. Use in open-field systems and in animal
husbandry is still limited. Phytoseiidae species are by far the most important group of
commercially available mite biocontrol agents with about 20 species offered worldwide.
Out of these, Amblyseius swirskii, Phytoseiulus persimilis, Neoseiulus cucumeris and
Neoseiulus californicus are the most important ones, covering together about two thirds of
the entire arthropod biocontrol agent market. The widespread use of these leaf-inhabiting
predatory mites has stimulated research into their biology and we now have substantial
knowledge on, for instance, the interaction between different predatory mite species, that
helps to improve biocontrol programmes. Soil predatory mites, for example Stratiolaelaps
scimitus (Laelapidae) or Macrocheles robustulus (Macrochelidae) for the control of sciarid
fly larvae and thrips pupae are much less frequently used and also much less researched.
This makes further development of biocontrol strategies using these mites more difficult.
Currently, there appears to be no reliable method to quantify the abundance of these mites
in soil samples. In studies at our laboratory, the frequently used Berlese-Tullgren funnels
gave very variable results. We observed that soil predatory mites can even multiply during
the extraction process. In addition to the control of plant pests, predatory mites can also be
used to control parasites of animals like the poultry red mite, Dermanyssus gallinae. Good
results have been obtained applying a combination of the predatory mites Androlaelaps
casalis (Laelapidae) and Cheyletus eruditus (Cheyletidae) in laying hen stables. This paper
provides an overview on the current status of commercial biological control using predatory
mites and identifies research needs to make the currently available mite biocontrol agents
even more successful and extend biological control with mites to other areas.

Keywords biological control, Phytoseiidae, soil predatory mites, poultry mites

Received 17 November 2016 Introduction

Accepted 29 March 2018
Published 28 September 2018 Biological control is a key ecosystem service and an underlying pillar of integrated pest
Corresponding author management (IPM) (Naranjo et al. 2015). In many parts of the world, augmentative biological
Markus Knapp: [email protected] control of insect and mite pests is common practice as part of the IPM strategy in greenhouse
Academic editor cultivation of vegetables, fruits and ornamentals and 80% of the revenue created by commercial
Serge Kreiter biological control is created in protected cultivation (Pilkington et al. 2010). Predatory mites,
mainly from the family Phytoseiidae play the leading role among the biocontrol agents used.
DOI This paper will (1) give an overview on the importance of predatory mites in biocontrol;
10.24349/acarologia/20184275 (2) show research needs to improve biological control, mainly with regard to soil-inhabiting
predatory mites and present some preliminary experiments on this; (3) describe the potential of
Copyright
predatory mites in the control of poultry red mite, Dermanyssus gallinae (DeGeer).
Knapp M. et al.
Distributed under
Creative Commons CC-BY 4.0
How to cite this article Knapp M. et al. (2018), Use of predatory mites in commercial biocontrol: current status and
future prospects. Acarologia 58(Suppl): 72-82; DOI 10.24349/acarologia/20184275
 Table 1 The most important arthropod biocontrol agents (by turnover) used in augmentative biolog-
ical control (modified after van Lenteren 2012).

Species Family Target(s) Year of first use

Amblyseius swirskii Phytoseiidae Whiteflies, thrips, mites 2005
Phytoseiulus persimilis Phytoseiidae Spider mites 1968
Neoseiulus californicus Phytoseiidae Mites 1985
Macrolophus pygmaeus Miridae Whiteflies 1994
Encarsia formosa Aphelinidae Whiteflies 1926
Orius laevigatus Anthocoridae Thrips 1993
Nesidiocoris tenuis Miridae Whiteflies, Tuta absoluta 2003
Neoseiulus cucumeris Phytoseiidae Thrips 1985
Eretmocerus eremicus Aphelinidae Whiteflies 1995
Aphidius colemani Braconidae Aphids 1991

The importance of predatory mites in biological control

The market for arthropod biological control agents has been growing with a rate of around 15%
per year in the last decades (Ravensberg 2015). In 1997, the worldwide turnover of natural
enemies was only about US$ 30 million (Bolckmans 1999), this increased to about US$ 50
million in 2000 (van Lenteren 2007) and to about US$ 400 million in 2010 (van Lenteren
2012); in 2014 it was about US$ 600 million.
The total number of arthropod species commercially available for augmentative biological
control worldwide was about 125 in 2000 (van Lenteren 2000). In the next decade about
100 species were added. Of the 219 species listed by van Lenteren (2012) more than half
were Hymenoptera (52.2%) followed by Acari (13.1%), Coleoptera (12.2%) and Heteroptera
(8.3%). The 25 most frequently used species make up more than 90% of the turnover. The vast
majority of them is used to control whiteflies, thrips, spider mites and aphids (van Lenteren
2012). This is also clearly to see in the ranking of the most important natural enemies. In
the 1990s, 25% of the market was taken by the whitefly parasitoid Encarsia formosa Gahan,
followed by the spider mite predator Phytoseiulus persimilis Athias-Henriot (12%) and the
thrips predator Neoseiulus cucumeris (Oudemans) (12%) (Bolckmans 1999). This situation
has changed in the last years. Currently more than 60% of the market is accounted for by
phytoseiid predatory mites with Amblyseius swirskii (Athias-Henriot), P. persimilis, Neoseiulus
californicus (McGregor) and N. cucumeris as the major species (Table 1). One of the major
drivers of this change was the introduction of A. swirskii into the market in 2006. This species
can control thrips and whiteflies simultaneously. It was the first predatory mite available that
was effective against whiteflies (Nomikou et al. 2001), which until then were mainly controlled
by parasitoids. The major advantages of A. swirskii compared to parasitoids are (1) it has a
generalist feeding habit and can establish and multiply on pollen and other prey and therefore
be introduced preventively, before the target pests are present in many crops; (2) it is easy
to produce because it is small, multiplies quickly and can be reared on stored product mites,
and (3) it can control several pests simultaneously. These characteristics made it possible
that A. swirskii developed into the most widely used biocontrol agent in protected cultivation
worldwide within only 10 years (Calvo et al. 2015).

The majority of the predatory mites commercially available in Europe are belonging to the
Phytoseiidae, followed by Laelapidae and one species each of Macrochelidae and Cheyletidae
(Table 2). The rise of phytoseiids in importance for biological control in turn increased the
interest in applied aspects of phytoseiid biology and a growing number of publications on, for

Knapp M. et al. (2018), Acarologia 58(Suppl): 72-82; DOI 10.24349/acarologia/20184275 73

 Table 2 Commercially available mite species used in biological control in Europe (modified after
van Lenteren 2012).

Year of first
Species Target(s)
commercial use
Phytoseiidae
Amblydromalus limonicus Thrips, whiteflies 2011
Amblyseius andersoni Mites, small insects 1995
Amblyseius swirskii Thrips, whiteflies 2005
Euseius gallicus Thrips, whiteflies 2013
Iphiseius degenerans Thrips 1993
Neoseiulus barkeri Thrips 1981
Neoseiulus californicus Spider mites 1985
Neoseiulus cucumeris Thrips 1985
Neoseiulus fallacis Spider mites 1997
Phytoseiulus persimilis Spider mites 1968
Transeius montdorensis Thrips, whiteflies 2003
Typhlodromus pyri Mites 1990

Laelapidae
Androlaelaps casalis Poultry red mite 2012
Gaeolaelaps aculeifer Sciarids, thrips 1995
Stratiolaelaps scimitus Sciarids, thrips 1995

Macrochelidae
Macrocheles robustulus Sciarids, thrips 2010

Cheyletidae
Cheyletus eruditus Poultry red mite, stored product pests 1985

instance influence of prey abundance and diversity on performance (Messelink et al. 2008),
intra-guild interaction (Abad-Moyano et al. 2010; Buitenhuis et al. 2010), influence of host
plants (Beard and Walter 2001; van Houten et al. 2013; Buitenhuis et al. 2014) or on the
evaluation of biocontrol using phytoseiids in commercial crops (Gacheri et al. 2015; Sampson
and Kirk 2016) help biocontrol agent producers and biocontrol practitioners to understand the
behaviour of phytoseiids and to further improve biological control.

Compared to the phytoseiids, soil-dwelling predatory mites so far play a minor role
in biological control. The four species currently on the market, Gaeolaelaps aculeifer
(Canestrini), Gaeolaelaps gillespiei Beaulieu, Stratiolaelaps scimitus (Womersley) (Laelapidae)
and Macrocheles robustulus (Berlese) (Macrochelidae) probably account for less than 2% of the
turnover with natural enemies. This group is also much less researched than the Phytoseiidae;
the number of records in the Web of Science core collection is 2109 for Phytoseiidae, 229 for
Laelapidae and 109 for Macrochelidae (check date 16 June 2016). The biology and use in
biological control of these soil-dwelling species was recently reviewed (Moreira and Moraes
2015; De Azevedo et al. 2015). All of them can feed on a wide range of prey, however
few studies have been conducted to validate their effectivity in field experiments and results

Knapp M. et al. (2018), Acarologia 58(Suppl): 72-82; DOI 10.24349/acarologia/20184275 74

 are not consistent (see e.g. Manners et al. 2013; Pozzebon et al. 2015). Despite this, a
significant number of greenhouse growers regularly apply soil-dwelling predatory mites to
control sciarid larvae and thrips. One of the challenges in semi-field and field studies is the
quantitative assessment of the density of the soil-dwelling predatory mites released. Usually
Berlese-Tullgren funnels are used for this purpose (Freire et al. 2007; Grosman et al. 2011),
however the size and design of these funnels can vary widely and data on their extraction
efficiency are lacking. Therefore, we have conducted some preliminary experiments on the
extraction efficiency of a commercially available Berlese-Tullgren funnel (see below).

Preliminary experiments on the extraction efficiency of

Berlese-Tullgren funnels
Berlese-Tullgren funnels are widely used to extract arthropods from soil or vegetation samples
(Edwards 1991, André et al. 2002). However, they have mostly been used for ecological or
taxonomic studies and their extraction efficiency for specific soil arthropods is very variable.
There are very few studies investigating extraction efficiency by recapture of known numbers
of specific arthropods added to defaunated soil or other medium (Brady 1969; André et al.
2002).
For our experiments we used a commercially available Berlese-Tullgren apparatus consist-
ing of 12 funnels (Burkard Scientific, Uxbridge, UK). This apparatus is supplied with 25 W
light bulbs by the manufacturer. As the extractions took very long in preliminary trials, we also
tested 40 W bulbs in some experiments. The removable upper part of the funnels is equipped
with a 2 mm mesh at the bottom. To reduce the amount of debris falling into the sampling vials
we added a finer mesh (800 µm) and put 10 g of buckwheat husks on top of this mesh before
adding potting soil. The collection vials fixed at the lower end of the funnels were filled with
70% alcohol. We used standard potting soil (50% coco peat, 15% white peat, 35% frozen black
peat: pH 5.5, e.c. 1.0), which was sterilized to kill any potential pre-existing fauna and then
re-moistened to a water content of approximately 70% (w/w). Experiments were conducted
with the soil-dwelling predatory mite species M. robustulus and G. aculeifer. Both species were
reared in the laboratory at 25 °C; M. robustulus was reared on Carpoglyphus lactis (Linnaeus)
and G. aculeifer on Acarus siro (Linnaeus).

Moisture content of soil samples

To determine the reduction in moisture content of the samples in the funnels with 25 W and
40 W bulbs, 500 mL of the potting soil (without mites) was filled in the removable upper part
of funnels. This resulted in a layer of approximately 7.5 cm height. Thereafter the removable
containers were placed into the Berlese apparatus equipped with 25 W and 40 W light bulbs,
respectively. The moisture content was determined with a moisture analyser (Sartorius MA 160,
Sartorius Lab Instruments, Göttingen, Germany) immediately before the start of the experiment
and thereafter by weighing the samples regularly for 9 days. At the end of the trial period the
moisture content was determined again with the moisture analyser.
The water content was 71.5% when the experiment started and declined faster under the
40 W bulb than under the 25 W bulb. In both treatments the moisture content did not decline
further after day 7 (Figure 1). The moisture content at the end of the trial was 8.1% and 9.2%
with 40 W and 25 W, respectively.

Extraction efficiency of M. robustulus with and without C. lactis

Thirty adult predatory mites (80% females) were added to 500 mL moist potting soil and
carefully mixed in a plastic bowl. In the treatments with C. lactis several thousand C. lactis
were added as prey for the M. robustulus. This was done because samples from the field
usually also contain prey for the predatory mites. Thereafter the samples were put on top of the

Knapp M. et al. (2018), Acarologia 58(Suppl): 72-82; DOI 10.24349/acarologia/20184275 75

 Figure 1 Development of the moisture content in potting soil samples in Berlese-Tullgren funnels
with 25 W and 40 W lamps.

buckwheat in the removable containers of the Berlese-Tullgren apparatus equipped with 40 W

lights. The containers were placed into the apparatus and the lights were switched on after 10
minutes. Three containers (replications) were prepared per treatment.
Without food 63% of the mites were extracted after 7 days. No more mites came out of the
sample during the next two days. When C. lactis was provided as food, more mites came out
of the samples than initially were put in. On day 5, an average of 52.7 mites had been collected
in the collection vials; 50% of them were immatures (Figure 2).

Extraction efficiency of G. aculeifer with and without A. siro

The setup was similar to the experiment with M. robustulus with the following exceptions. The
Berlese-Tullgren funnel was equipped with 25 W lamps; A. siro was used as prey mite instead
of C. lactis and 20 adult G. aculeifer (80% females) were used per funnel in the trial without
prey mites, whereas 30 mites were used in the experiment with prey mites.
Without prey mites, 85% of the mites were recaptured, when A. siro was added G. aculeifer
multiplied similarly to M. robustulus. More mites were extracted than introduced and the
proportion of immatures was 49%. In contrast to the trial with M. robustulus also larvae were
found in the extraction vials (Figure 3).
The most striking result of these experiments was that the predatory mites can multiply in
the Berlese-Tullgren funnel if food is available. Moisture and temperature in certain areas of
the sample seem to be conducive to the development of both predatory mite species. Egg to
adult development of M. robustulus can be extremely fast; it takes about 2 days at 25-29 °C
(Filipponi and Mosna 1968). Immature development of G. aculeifer is slower, however this
species can develop from egg to adult in 8 to 10 days between 27 and 30 °C; the eggs hatch
after around 3 days under these conditions (Amin et al. 2014).
Furthermore there are indications that the extraction of G. aculeifer is more efficient than
the extraction of M. robustulus if no food is available. Some M. robustulus adults also left the
soil sample by walking out of the top of the funnels. This was verified by putting double sided
tape on the outer rim of the funnels, after we observed that prey mites were walking on the
rim of the funnels. A few M. robustulus adults were found on this tape. As M. robustulus is

Knapp M. et al. (2018), Acarologia 58(Suppl): 72-82; DOI 10.24349/acarologia/20184275 76

 A

Figure 2 Number of Macrocheles robustulus collected in the collection vials without prey mites (A)
and with prey mites (B). The experiment was started with 30 adult M. robustulus (80% females) per
funnel. Vertical bars indicate standard errors for the total number of mites (nymphs and adults).

phoretic on beetles (Glida et al., 2003; Philips, 2009), these mites might have walked to the
upper surface in search a phoriont.
To our knowledge there is no other study investigating the re-capture of known numbers
of released soil-dwelling predatory mites with Berlese-Tullgren funnels. Brady (1969)
investigated the extraction efficiency of Caloglyphus berlesei (Michael), A. siro and Cheyletus
trouessarti (Oudemans) from defaunated poultry litter; he recorded recovery rates of 86-97%
for adults. Until a reliable extraction method has been developed, counts of soil-dwelling
predatory mites extracted with Berlese-Tullgren funnels from comparative experiments with
different species have to be interpreted with great care (see e.g. Messelink and van Holstein-Saj
2008; Grosman et al. 2011; Pijnakker et al. 2011).

Knapp M. et al. (2018), Acarologia 58(Suppl): 72-82; DOI 10.24349/acarologia/20184275 77

 A

Figure 3 Number of Gaeolaelaps aculeifer collected in the collection vials without prey mites (A)
and with prey mites (B). The experiment without prey mites (A) was started with 20 adult G. ac-
uleifer (80% females) per funnel; the experiment with prey mites (B) was started with 30 G. aculeifer
(80% females) per funnel. Vertical bars indicate standard errors for the total number of mites (larvae,
nymphs and adults).

Biological control of the poultry red mite, Dermanyssus

gallinae
The poultry red mite D. gallinae poses a serious threat to egg laying hens worldwide. Economic
costs associated with control and production losses have been estimated at € 130 million per
year for the European Union egg production industry. Dermanyssus gallinae has typically been
controlled by using synthetic acaricides but resistance of the mites to acaricides and legislative
restrictions on the use of acaricides make this approach more and more difficult (Sparango et al.
2014). The behaviour of the mites poses an additional challenge to their control. The majority
of the life cycle is spent off the host where the mites hide in large numbers in secluded areas like

Knapp M. et al. (2018), Acarologia 58(Suppl): 72-82; DOI 10.24349/acarologia/20184275 78

 cracks formed by timber joints or in metal profiles supporting the perches. Due to the difficulty
to conduct experiments on life chicken few data are available on the life cycle of D. gallinae.
Maurer and Baumgärtner (1992) found that immature development takes 7 days at 25 °C and
calculated an intrinsic rate of increase (rm ) of 0.120 at this temperature. Dermanyssus gallinae
can develop between 5 and 40 °C but conditions for juvenile development are most favourable
between 30 and 37 °C (Maurer and Baumgärtner 1992; Maurer and Hertzberg 2001).
Research on the control of D. gallinae with predatory mites has started later than research on
using predatory mites to control plant pests. Maurer (1993) discovered that Cheyletus eruditus
(Schrank) is associated with D. gallinae in Swiss poultry houses and can feed on its larvae.
However, initial trials with this predatory mite species in poultry houses yielded disappointing
results (Maurer and Hertzberg, 2001).
In a new effort to find biocontrol agents for poultry red mite Lesna et al. (2009) surveyed
the mite fauna in starling nests and chicken houses in the Netherlands. The predators C. eruditus
and A. casalis were frequently found both in starling nests and in chicken houses. Blattisocius
keegani Fox was also common in chicken houses but not in starling nests. Small experiments
in the laboratory confirmed that A. casalis and G. aculeifer predated on D. gallinae. The latter
predator was rare in starling nests and absent in chicken houses, but is available commercially
as a biocontrol agent. Later Ali et al. (2014) demonstrated that S. scimitus also predates the
poultry red mite in laboratory trials.
In small laying hen cages A. casalis and S. scimitus reduced the D. gallinae population
compared to untreated cages but A. casalis was more efficient than S. scimitus. However, both
species were not able to completely eradicate the poultry red mites from the cages (Lesna et al.
2012).
The hunting behaviour of C. eruditus and A. casalis is different. Cheyletus eruditus is
a “sit and wait”/ ambush predator. It rarely actively stalks its prey (Thind and Ford 2006).
Androlaelaps casalis is hunting for prey very actively (Barker 1968). At 25.6 °C, 75-100%
relative humidity and with Glycophagus domesticus (De Geer) as prey, development time of A.
casalis from egg to adult is about 8 days (Barker 1968).
Although C. eruditus has been known to occur in poultry houses for long (Brady 1970)
it has mainly been investigated as biological control agent of stored product mites (Zdarkova
1998). Maurer (1993) calculated an intrinsic rate of increase (rm ) of 0.144, which is higher than
the rm of D. gallinae (0.12, Maurer and Baumgärtner 1992) and a doubling time of 4.8 days for
C. eruditus on D. gallinae at 25 °C. With Lepidoglyphus destructor (Schrank) as prey, egg to
adult development of C. eruditus took 15.4 days, females laid 133 eggs (5.4 per day) and the
rm was 0.178 (Barker 1991).
Releases of mass-reared C. eruditus as well as A. casalis in chicken houses have not led to a
reliable control of D. gallinae (Maurer and Hertzberg 2001; Koppert unpubl. data). Therefore,
a strategy was developed based on the combined use of these two predatory mites. In aviary
systems, C. eruditus is released after cleaning the laying hen houses between two production
cycles, one week before the new hens are introduced, by spreading the mite together with a
carrier in the chicken houses. This strategy has been used in grain stores in the Czech Republic
in the past (Zdarkova and Horak 1990) and gives the mites time to find a hiding place and to
feed on any remaining poultry red mites. As the carrier is very attractive to the chicken and
C. eruditus tends to remain in the carrier for some time most predatory mites are eaten by the
hens quickly after spreading if the release is carried out when chicken are already present. The
release of A. casalis starts two weeks after the laying hens have arrived and some dung and dirt
has accumulated. Androlaelaps casalis is released by installing bottles containing predatory
mites, prey mites and a carrier, close to typical hiding places of D. gallinae in the structure of
the chicken house, or more humid places (dung accumulations). The predatory mites disperse
through the perforated lid of the bottles. At the same time D. gallinae are also entering the
bottles in search for hiding places and are subsequently fed on by the population of predatory
mites inside. These bottles can sustain a large population of predatory mites for more than 8
weeks; thereafter they are replaced. This system takes into account observations of Lesna et

Knapp M. et al. (2018), Acarologia 58(Suppl): 72-82; DOI 10.24349/acarologia/20184275 79

 al. (2012) who recorded that most poultry red mites were found higher up in the cage structure
whereas the predatory mites A. casalis and S. scimitus were mainly found on the cage floor, and
recommended to create artificial environments in which predatory mites can thrive and which
can be positioned higher up in the structural elements of cage and aviary systems.

Research needs
Biological control using leaf-inhabiting phytoseiid predatory mites has been booming during
the last decades. This was made possible through close collaboration between researchers
and biocontrol companies. Nevertheless, there are many challenges remaining, for instance
developing better systems for ornamentals where tolerance levels for pests are very low,
possibly by using providing alternative food sources for the predators (see e.g. Buitenhuis et
al. 2015; Delisle et al. 2015; Janssen and Sabelis 2015) or finding generalist predatory mites
that can successfully control pests on tomatoes (see e.g. van Houten et al. 2013).
The potential for the use of soil-dwelling predatory mites could be as high as for leaf-
inhabiting predatory mites. Many species of soil-dwelling predatory mites are known to feed
on important plant pests (Carrillo et al. 2015). However, very little is known about them besides
that they prey on a certain pest species and to realize their potential we need collaborative
efforts to develop further understanding of the biology and ecology of these mites and their
interaction with other soil organisms. We have shown that currently even reliable methods
to quantify the density of soil-dwelling predatory mites are lacking. Development of such a
method would be an important first step.
Control of poultry red mite has received more and more attention during the last few years.
Initial results with A. casalis and C. eruditus are promising in aviary systems but satisfactory
control will only be possible in combination with other measures.
Generally, mites have a huge potential in biological control. There small size and fast
population development makes them relatively easy to rear and to transport to the end users.
Further improvements of mass-rearing and introduction techniques will make biological control
using predatory mites even more robust and reliable as well as more affordable for growers.

Acknowledgements
We thank Alma van der Heiden for her assistance with the soil mite extraction experiments.

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