9
Evaluation of seven plant species/cultivars for their suitability as banker plants for Orius insidiosus (Say) M. O. Waite C. D. Scott-Dupree M. Brownbridge R. Buitenhuis G. Murphy Received: 8 May 2013 / Accepted: 17 October 2013 / Published online: 26 October 2013 Ó International Organization for Biological Control (IOBC) 2013 Abstract Marigold (cv. Lemon Gem), castor bean, ornamental pepper (cv. Black Pearl and Purple Flash), gerbera daisy (cv. Festival), feverfew, and sunflower (cv. Choco Sun) were evaluated for their suitability as banker plants (BP) for Orius insidiosus (Hemiptera: Anthocoridae) in commercial greenhouses. Oviposi- tion, egg hatch, nymphal development to adulthood, and population increase were quantified in laboratory trials. Assessments of oviposition and egg hatch indicated that all plants tested were equally accepted by O. insidiosus. Nymphal development to adulthood and survival tests indicated that gerbera may be a suitable BP as survival was the highest (58.1 %), whereas marigold would not be an acceptable BP as only 10.7 % of nymphs survived to adulthood. Nym- phal development time differed by only one day among all plants. In greenhouse cage experiments, Purple Flash pepper supported the greatest population growth over a ten week period. Based on the com- bined results from all tests, Purple Flash pepper appears to have the greatest potential as a BP species for O. insidiosus. Keywords Orius insidiosus Á Hemiptera Á Anthocoridae Á Frankliniella occidentalis Á Banker plant Á Biological control Introduction In greenhouse ornamental crops, western flower thrips (WFT), Frankliniella occidentalis Pergande (Thysa- noptera: Thripidae), is one of the most economically important and challenging pests to control, causing aesthetic feeding damage to plants and vectoring tospoviruses (German et al. 1992; Tommasini and Maini 1995; Daughtrey et al. 1997; Brødsgaard 2004; Bosco et al. 2008). Insecticide resistance, worker health and safety considerations, and a lack of efficacious insecticides have provided incentives to increase the use of biological control agents to manage WFT globally. However, lack of access to effective chemical control options has driven an accelerated use of biological control in Canadian floriculture greenhouses. Handling Editor: Patrick De Clercq. M. O. Waite (&) Monsanto Canada Inc., Guelph, ON N1G 0B4, Canada e-mail: [email protected] C. D. Scott-Dupree School of Environmental Sciences, University of Guelph, Guelph, ON N1G 2W1, Canada M. Brownbridge Á R. Buitenhuis Horticultural Production Systems, Vineland Research and Innovation Centre, Vineland, ON L0R 2E0, Canada G. Murphy Ontario Ministry of Agriculture, Food and Rural Affairs, Vineland, ON L0R 2E0, Canada 123 BioControl (2014) 59:79–87 DOI 10.1007/s10526-013-9549-4

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  • Evaluation of seven plant species/cultivars for theirsuitability as banker plants for Orius insidiosus (Say)

    M. O. Waite C. D. Scott-Dupree

    M. Brownbridge R. Buitenhuis G. Murphy

    Received: 8 May 2013 / Accepted: 17 October 2013 / Published online: 26 October 2013

    International Organization for Biological Control (IOBC) 2013

    Abstract Marigold (cv. Lemon Gem), castor bean,

    ornamental pepper (cv. Black Pearl and Purple Flash),

    gerbera daisy (cv. Festival), feverfew, and sunflower

    (cv. Choco Sun) were evaluated for their suitability as

    banker plants (BP) for Orius insidiosus (Hemiptera:

    Anthocoridae) in commercial greenhouses. Oviposi-

    tion, egg hatch, nymphal development to adulthood,

    and population increase were quantified in laboratory

    trials. Assessments of oviposition and egg hatch

    indicated that all plants tested were equally accepted

    by O. insidiosus. Nymphal development to adulthood

    and survival tests indicated that gerbera may be a

    suitable BP as survival was the highest (58.1 %),

    whereas marigold would not be an acceptable BP as

    only 10.7 % of nymphs survived to adulthood. Nym-

    phal development time differed by only one day

    among all plants. In greenhouse cage experiments,

    Purple Flash pepper supported the greatest population

    growth over a ten week period. Based on the com-

    bined results from all tests, Purple Flash pepper

    appears to have the greatest potential as a BP species

    for O. insidiosus.

    Keywords Orius insidiosus Hemiptera Anthocoridae Frankliniella occidentalis Banker plant Biological control

    Introduction

    In greenhouse ornamental crops, western flower thrips

    (WFT), Frankliniella occidentalis Pergande (Thysa-

    noptera: Thripidae), is one of the most economically

    important and challenging pests to control, causing

    aesthetic feeding damage to plants and vectoring

    tospoviruses (German et al. 1992; Tommasini and

    Maini 1995; Daughtrey et al. 1997; Brdsgaard 2004;

    Bosco et al. 2008). Insecticide resistance, worker health

    and safety considerations, and a lack of efficacious

    insecticides have provided incentives to increase the use

    of biological control agents to manage WFT globally.

    However, lack of access to effective chemical control

    options has driven an accelerated use of biological

    control in Canadian floriculture greenhouses.

    Handling Editor: Patrick De Clercq.

    M. O. Waite (&)Monsanto Canada Inc., Guelph, ON N1G 0B4, Canada

    e-mail: [email protected]

    C. D. Scott-Dupree

    School of Environmental Sciences, University of Guelph,

    Guelph, ON N1G 2W1, Canada

    M. Brownbridge R. BuitenhuisHorticultural Production Systems, Vineland Research and

    Innovation Centre, Vineland, ON L0R 2E0, Canada

    G. Murphy

    Ontario Ministry of Agriculture, Food and Rural Affairs,

    Vineland, ON L0R 2E0, Canada

    123

    BioControl (2014) 59:7987

    DOI 10.1007/s10526-013-9549-4

  • The biological control agent, Orius insidiosus Say

    (Hemiptera: Anthocoridae), is an omnivore and has

    been documented as feeding upon a variety of arthropod

    pests in addition to pollen and plant sap (Almer et al.

    1998; Deligeorgidis 2002; Osborne et al. 2004).

    Successful control of WFT using augmentative releases

    of O. insidiosus has been achieved in greenhouse

    vegetable crops such as sweet peppers, where the pollen

    in the flowers provides a supplemental source of food

    that aids predator survival and establishment (Chambers

    et al. 1993). Populations of O. insidiosus can persist for

    an extended period in the absence of prey providing they

    are able to access alternative food sources. For example,

    populations were maintained for six months on flower-

    ing sweet peppers with low to absent populations of

    WFT (van den Meiracker and Ramakers 1991; van

    Lenteren and Loomans 1999). This suggests that

    inoculative releases of O. insidiosus may be sufficient

    to control thrips over an entire growing season if the

    predator is released on pollen-producing plants.

    The use of O. insidiosus to control WFT in

    greenhouse ornamental crops, such as chrysanthe-

    mums, has been limited to date as the predatory bug can

    be slow to establish and exert controla critical factor

    in short-term crops. Relatively poor establishment of

    O. insidiosus is achieved in ornamentals as a result of

    several factors. First, a lack of prey: as a result of the

    low tolerance for thrips and the preference of WFT for

    flowers, there are often few WFT in the vegetative

    stages of plant development. Secondly, a lack of

    alternative food sources as ornamentals typically are

    not flowering in the production area of greenhouses

    until just before they are shipped so pollen is not readily

    available. Lastly, eggs of O. insidiosus are removed

    from the greenhouse when the plants are shipped,

    severely limiting population growth in production

    areas. Repeated releases of O. insidiosus purchased

    from commercial insectaries are thus necessary to

    maintain an effective O. insidiosus population in the

    greenhouse, a strategy that is not economically viable.

    Determining a suitable flowering banker plant (BP)

    system for O. insidiosus could improve opportunities to

    use the predator to control WFT by providing a source of

    supplementary food in the form of pollen (Frank 2010;

    Huang et al. 2011). Establishing such a system in the

    greenhouse would allow O. insidiosus to establish and

    increase its population, offering growers the option of

    preventative introductions. An effective BP should

    provide a location for feeding and reproduction, as well

    as allowing nymphs to reach the adult stage quickly,

    ensuring a high survival rate, and supporting population

    growth. The objective of this study was to evaluate the

    potential of marigold (cv. Lemon Gem), castor bean,

    ornamental pepper (cv. Black Pearl and Purple Flash),

    gerbera daisy (cv. Festival), feverfew, and sunflower

    (cv. Choco Sun) plants to support a population of Orius

    insidiosus in commercial greenhouses. The plant spe-

    cies/cultivars were selected based upon growers obser-

    vations of wild O. insidiosus specimens within and

    adjacent to greenhouses and a literature review of plants

    known to have favourable characteristics for supporting

    O. insidiosus and similarities in their growth require-

    ments as ornamental crops. The Black Pearl ornamental

    pepper is currently used as a BP by some commercial

    growers at a rate of 100 BPs per acre (pers. obs, G.M.) as

    it has been shown to support O. insidiosus in the absence

    of prey (Wong and Frank 2012, 2013). Studies by Wong

    and Frank (2012, 2013) indicate that pollen from Black

    Pearl peppers can result in larger populations of O.

    insidiosus by increasing the longevity of the predator

    when prey are absent, reduce development time and

    increase likelihood of survival to adult. While it has been

    shown that pollen from the Black Pearl pepper plants

    provides a suitable source of nutrition for O. insidiosus,

    growers using Black Pearl pepper BPs have had varying

    degrees of success in establishing populations of O.

    insidiosus. Other plant species or cultivars may be more

    suitable as BPs. Three tests were conducted to assess

    different attributes of the different plant species/culti-

    vars with regard to their comparative suitability to serve

    as BPs, namely:

    (a) Acceptance by O. insidiosus for oviposition and

    rate of egg hatch.

    (b) Orius insidiosus development time and survival

    from first instar to adult.

    (c) Effect of host plant on O. insidiosus population

    growth.

    Methods

    Plants

    All plants (Table 1) were grown in greenhouses at the

    Vineland Research and Innovation Centre in Vine-

    land, ON Canada under growing conditions of

    21 1 C, RH 70 5 %, 16: 8 h (L:D). Plant

    80 M. O. Waite et al.

    123

  • species used in the BP trials were grown from seed

    with the exception of gerbera, which was purchased as

    a flowering plant from commercial growers who did

    not use chemical insecticides. Seeds were hand sown

    two months prior to trial initiation into seeding trays

    filled with ProMix potting medium (Premier Tech

    Horticulture, Rivie`re-du-Loup QC Canada). One -

    month later, seedlings were transferred to 15 cm diam

    plastic pots containing the same growing mix. All

    plants were watered once daily by hand using a water

    and fertilizer (202020, NPK) mixture until sub-

    strate was uniformly moist.

    Insects

    Orius insidiosus adults were obtained from Biobest

    Biological Systems Canada Ltd. (Leamington ON

    Canada). A colony was maintained in a growth cabinet

    [25 1 C, 70 5 % R.H., 16:8 h (L:D)] at theVineland Research and Innovation Centre. Adults

    were transferred to cages which were constructed from

    plastic containers (750 ml disposable Ziploc Glad-

    Ware container) in which two holes (2 cm diameter)

    had been cut into opposite sides of the container that

    were covered with mesh screen (180 lm) to provideventilation. A handful of buckwheat hulls was placed

    into each container to serve as a refuge for the insects.

    Two green bean pods (Phaseolus vulgaris L.) were

    placed into each cage as an oviposition substrate

    (Richards and Schmidt 1996), along with a kidney

    bean leaf to provide moisture. Frozen Ephestia

    kuehniella Zeller eggs (Beneficial Insectary, Redding

    CA) adhered to a (4 9 2 cm) Post-it note were also

    provided as a food source. Adults were allowed to feed

    ad libidum throughout. Every three days, the green

    beans were transferred to a new cage to produce

    discrete even-aged cohorts and egg strips, beans and

    leaves were replaced as needed. Only adult cages

    received green beans for oviposition.

    Bioassays

    Evaluation of plant species/cultivars as oviposition

    substrate for Orius insidiosus

    Adults were removed from the rearing colony using an

    aspirator and their sex determined by observing the

    abdomen (209, Olympus SZ61) which is symmetrical

    for females and asymmetrical for males as their

    genitalia are sickle-shape (Slater 2005). Five adult

    females, which were 7 1 days old to ensure that

    they were sexually mature and had mated, were

    collected, placed in containers and starved for 24 h.

    The females were placed into cages containing one

    stem from an individual test plant species/cultivar.

    Each plant stem carried multiple flowers, with the

    exception of gerbera and sunflower in which each cup

    contained one flower. The base of each stem was

    wrapped in cotton batting and fed through a hole in the

    lid of a 2 oz Solo cup. The cup was filled with water

    to prevent the plant from desiccating. The cotton

    batting placed around the stem of plants served to

    prevent adults from entering the water-filled cup. The

    entire set up was then placed into an 8 oz Solo dish

    and a bottle cage positioned on top of each dish. A

    bottle cage was constructed from the bottom portion of

    a 2 l plastic bottle. To provide ventilation and prevent

    build-up of condensation in the cages, 2 9 5 cm

    diameter holes were made in the sides of the bottle

    using a punch that was heated over a Bunsen burner.

    Mesh-screening was glued over the holes using a hot

    glue gun. The cages were held in a growth chamber

    [25 1 C, 70 5 % R.H., 16:8 h (L:D)]. After48 h, the cages were removed from the chamber and

    the adults were removed from the cages. The number

    Table 1 Plant species/cultivars evaluated as potential bankerplants for Orius insidiosus

    Common name Species Source

    Marigold, cv.

    Lemon Gem

    Tagetes

    patula

    Veseys Seeds Ltd,

    Charlottetown, PE

    Castor Bean Ricinus

    communis

    Richters Herbs,

    Goodwood, ON

    Ornamental Pepper,

    cv. Black Pearl

    Capsicum

    annuum

    Stokes Seeds Ltd.,

    Thorold, ON

    Ornamental Pepper,

    cv. Purple Flash

    Capsicum

    annuum

    Stokes Seeds Ltd.,

    Thorold, ON

    Gerbera Daisy, cv.

    Festival

    Gerbera

    jamesonii

    Orchard Park Growers,

    St. Catharines, ON

    Lakeshore Inc., Jordan

    Station, ON

    Feverfew Tanacetum

    parthenium

    Richters Herbs,

    Goodwood, ON

    Sunflower, cv.

    Choco Sun

    Helianthus

    annuus

    Stokes Seeds Ltd.,

    Thorold, ON

    All source locations are within Canada

    Evaluation of seven plant species/cultivars 81

    123

  • of eggs laid on each stem and the oviposition sites

    were recorded. Stems were returned to their respective

    cages and returned to the growth chamber.

    Egg viability was assessed by counting the number

    of nymphs as a proportion of eggs laid on each stem

    using a plant washing technique. Stems were removed

    from the bottle cages on Day 6 (five days after the

    females were introduced into the cages) and placed

    into 100 ml plastic jars with lids containing an E.

    kuehniella egg strip (2 9 2 cm) as a food source for

    emerging nymphs. Jars were then returned to the

    growth chamber. On Day 8, the egg strips were

    removed from the jars and 40 ml of 70 % ethanol was

    added. Parafilm was placed onto the top of the jar,

    and the jars shaken for 60 s. The jars were emptied

    into a funnel lined with a Whatman no. 4 filter paper,

    which was placed on top of a vacuum flask and the

    ethanol was withdrawn. The filter paper was trans-

    ferred to a Petri dish and observed under a microscope

    (209, Olympus SZ61) to count the numbers of

    nymphs. The stem was also examined under a

    microscope to locate any nymphs which did not

    transfer to the filter paper during the plant washing

    process. The oviposition assays were conducted in

    three blocks and replicated eight times per plant

    species/cultivar.

    Development of first instar Orius insidiosus nymphs

    to the adult stage on selected plant species/cultivars

    Orius insidiosus adults (7 1 days old) were

    removed from the rearing colony using an aspirator

    and their sex was determined as described above.

    Twenty females were collected and transferred for

    24 h to a plastic cage containing four green bean pods

    for oviposition. The eggs laid on the green beans in the

    plastic cages were observed daily for hatching to

    ensure nymphs used in the development assays were

    \24 h old. Nymphs hatched from these eggs wereused in this study to ensure that all individuals had the

    same nutritional starting point. These first instars were

    transferred to development cages using a moistened

    paint brush. Development cages were constructed by

    cutting a small hole in the centre of the bottom of a

    60 ml Solo cup (Kaumeyer Paper Ltd., St. Catha-

    rines, ON). The Solo cup was then placed inside a

    250 ml plastic DART Conex Classic cup. To

    provide ventilation, two 2 cm diameter holes were

    cut into the sides of the 250 ml DART cup and mesh

    screening was glued over the holes. A stem from an

    individual test plant species/cultivar was inserted

    through the hole cut into the Solo cup and wrapped

    in cotton to prevent nymphs from escaping. The

    bottom of the DART cup was filled with water and the

    Solo cup containing an individual plant cutting from

    each plant species/cultivar was placed inside. A

    nymph was placed on the plant and the lid was placed

    on the cup. Cages were placed in a growth chamber

    [25 1 C, 70 5 % R.H., 16:8 h (L:D)] andnymphs were monitored daily for development,

    recorded as the number of days to adulthood and

    survival. Observations were continued until the adult

    stage was reached or the nymph died. The experiment

    was repeated until at least 15 nymphs had reached the

    adult stage on each BP species.

    The effect of host plant on Orius insidiosus population

    growth

    The suitability of plant species/cultivars as host plants

    was evaluated based upon the ability of an Orius

    population to increase in cage bioassays conducted in a

    greenhouse at the Vineland Research and Innovation

    Centre. Eight individual potted flowering plants, one of

    each species/cultivar, were randomly assigned to a

    dome cage (60 cm960 cm960 cm; Model 2120F,

    MegaView Science Co. Ltd; Taichung, Taiwan).

    Plants were watered twice daily for 2 min using a drip

    irrigation system (2 l h-1 flow rate) and fruit and dead

    flowers removed weekly (ensuring no O. insidiosus

    were removed on plant matter). Each treatment

    received ten female and five male adults (7 1 days

    old) which had been collected from the rearing colony.

    Populations of O. insidiosus on each plant were

    sampled bi-weekly for ten weeks. Sampling was

    conducted by placing a white tray inside the dome

    cage and tapping the plant vigorously over the tray to

    dislodge nymphs and adults. Dislodged individuals

    were collected using an aspirator. The seams and

    ceiling of the cage were inspected for additional adults

    which may not have landed on the white tray, and were

    also collected using the aspirator. The collected

    nymphs and adults were counted, the number recorded,

    and insects returned to the appropriate cage. The

    greenhouse bioassays were conducted in three blocks

    with eight replicates for each plant species/cultivar.

    82 M. O. Waite et al.

    123

  • Data analysis

    Oviposition and number of hatched nymphs (as a

    proportion of eggs laid) data were subjected to an

    analysis of variance (ANOVA) using PROC GLM in

    SAS v. 9.2 (SAS Institute, Cary, NC, USA). Means for

    the number of eggs laid and the numbers of hatched

    nymphs were analyzed independently and were sep-

    arated using Tukeys multiple means comparison. The

    mean of the residuals was equal to zero and a Shapiro

    Wilk test confirmed that the residuals were approxi-

    mately normally distributed. A Type I error rate (a) of0.05 was used to test for significance.

    Nymphal development data was subjected to an

    analysis of variance (ANOVA) using PROC GLM in

    SAS v. 9.2 (SAS Institute, Cary, NC, USA). Means

    were separated using Tukeys multiple means com-

    parison. A Type I error rate (a) of 0.05 was used to testfor significance.

    A repeated measures ANOVA using the Mixed

    procedure, SAS v. 9.2 (SAS Institute, Cary, NC, USA),

    with variance partitioned into the fixed effect treatment

    and the random effect block, was used to test the effect

    of host plant on population growth. The total number of

    O. insidiosus individuals counted every two weeks

    was analyzed to determine population growth over

    time (ten week sampling period). The ShapiroWilk

    test confirmed that the residuals were approximately

    normally distributed. Differences between means were

    determined using Tukeys multiple means comparison

    using a significance level of a = 0.05.

    Results

    Plant species had an effect on the oviposition of O.

    insidiosus (F = 2.50; df = 6, 46; P = 0.035) (Fig. 1).

    Significantly fewer eggs (mean SE) were found on

    sunflowers (14.9 7.1 eggs plant-1; P = 0.042)

    than marigold (49.9 6.4; P \ 0.0001). Differencesin the number of eggs laid by O. insidiosus on

    marigold, gerbera, Black Pearl pepper, Purple Flash

    pepper, feverfew, and castor bean, were not signifi-

    cant. Plant species/cultivar did not have an effect on

    the emergence of nymphs (F = 1.17; df = 6, 38;

    P = 0.34) (Fig. 1). The mean number of nymphs as a

    proportion of emergence from eggs laid did not

    significantly differ on sunflowers from the other six

    potential BP species.

    Nymphs reared on gerbera had the highest survival

    rate at 58.1 %. In contrast, only 10.7 % of nymphs

    reared on marigold reached the adult stage (Table 2).

    No nymphs developed to adulthood on sunflowers.

    Nymphal development time from first instar to adult

    was affected by plant species (F = 3.47; df = 5, 77;

    P = 0.007) (Fig. 2). Development time for nymphs

    reared on gerbera (8.3 0.13 days) was significantly

    shorter than for those reared on marigold (9.2 0.16

    days) and Black Pearl pepper (9.2 0.23 days) and

    Purple Flash (9.1 0.21 days). There was no signif-

    icant difference in development times on feverfew,

    castor bean, Black Pearl pepper and Purple Flash

    pepper (Fig. 2).

    There were significant differences in the size of an

    O. insidiosus population developing on the different

    plant species/cultivars (F = 23.35; df = 5, 201;

    P \ 0.0001; Fig 3). Population size was significantlyaffected by plant species (F = 25.64; df = 5, 201;

    P \ 0.0001), week (F = 26.10; df = 4, 201; P \

    0

    10

    20

    30

    40

    50

    60

    Plant species/cultivar

    Mean number of eggsMean number of nymphs

    a

    a

    a

    a

    a

    b

    ab

    Mea

    n #

    of e

    ggs

    or n

    ymph

    s/ p

    lant

    A A

    AA A

    A

    A

    Fig. 1 Mean number (SE) of Orius insidiosus eggs laid byfive females in 48 h and subsequent hatched nymphs on seven

    potential banker plant species/cultivars in no-choice tests in a

    growth chamber at 25 1 C, RH 70 5 %, 16:8 h L:D(n = 8). Means for the number of eggs laid and the numbers of

    hatched nymphs were analyzed independently and are indicated

    by lower case (eggs) or upper case (nymphs) letters. Means

    indicated by the same upper or lower case are not significantly

    different according to Tukeys test (P [ 0.05)

    Evaluation of seven plant species/cultivars 83

    123

  • 0.0001), and the plant 9 week interaction (F = 5.00;

    df = 20, 201; P \ 0.0001). In each assessment week,the population of O. insidiosus on Purple Flash pepper

    plants was significantly higher (P \ 0.0001) than onany other plant species (Fig. 3). Marigold supported

    significantly lower populations of O. insidiosus

    throughout (P \ 0.0001).

    Discussion

    Based on the results from the current study, Purple

    Flash ornamental pepper is the best candidate for use

    as a BP for O. insidiosus, having characteristics that

    were superior to all other species/cultivars tested. It

    appears that the Purple Flash pepper is highly suitable

    as a banker plant for O. insidiosus, promoting

    oviposition, nymphal development and survival, and

    supporting population development to levels that were

    greater than on any other species/cultivar.

    The first stage of the evaluation was done to

    determine the suitability of host plants for oviposition

    by O. insidiosus as a measurement of both the quality

    of plants as oviposition substrate and by their quality

    as food. Oviposition preferences are based upon

    physical characteristics of a plant, such as trichome

    density and epidermal thickness (Coll 1996; Lundgren

    and Fergen 2006; Seagraves and Lundgren 2010).

    Oviposition was equivalent on Lemon Gem marigold,

    castor bean, feverfew, Black Pearl pepper, Purple

    Flash pepper and gerbera daisy. The number of eggs

    laid on Choco Sun sunflowers was significantly lower

    than on plants other than gerbera. The number of eggs

    counted on the Choco Sun sunflowers, however, was

    inaccurate as fewer eggs were counted than nymphs

    recovered. Errors in egg counts were likely a result of

    O. insidiosus oviposition habits: the insect lays eggs in

    concealed locations (e.g., sepals) and consequently

    many eggs were missed. This is supported by the fact

    that there are no significant differences between the

    numbers of emerged nymphs found on each plant

    species.

    In nymphal development and survival trials, no

    nymphs reached the adult stage on the Choco Sun

    sunflowers. This could be attributed to a severe powdery

    mildew (Blumeria graminis) infection that occurred on

    the sunflower plants during the experiment. Sunflowers

    were selected as a BP candidate as large Orius

    populations can be found in field sunflowers during

    summer months. However, owing to their susceptibility

    to powdery mildew, sunflower cannot be recommended

    as a BP for O insidiosus in greenhouses. BPs, like crop

    plants, are susceptible to pests and diseases, and this has

    to be considered as a factor in the selection of

    appropriate candidate species (Huang et al. 2011).

    Several studies have determined that O. insidiosus

    females select oviposition sites based upon plant

    species, and specific locations on a plant favour

    Table 2 Survival (%) of Orius insidiosus nymphs (\24 h old)reared on seven potential banker plant species at 25 1 C,R.H. 70 5 %, 16:8 h (L:D)

    Banker plant na Survival (%)

    Gerbera daisy 31 58.1

    Black pearl pepper 32 50.0

    Purple flash pepper 40 45.0

    Feverfew 45 42.2

    Castor bean 41 36.6

    Lemon gem marigold 140 10.7

    Choco sun sunflower 15 0.0

    a Initial number of nymphs

    bc c

    ab c c

    0

    1

    2

    3

    4

    5

    6

    7

    8

    9

    10

    Mea

    n de

    velo

    pmen

    t tim

    e (d

    ays)

    Plant species/cultivar

    a

    Fig. 2 Mean development time (days SE) of Orius insidio-sus nymphs (\24 h old) to reach the adult stage when reared onsix potential banker plant species/cultivars in a growth chamber

    at 25 1 C, R.H. 70 5 %, 16:8 h L:D. Means followed bythe same letter are not significantly different from each other by

    Tukeys test (P [ 0.05). No nymphs reached the adult stage onthe sunflowers (cv. Choco Sun) and these replicates were

    excluded from statistical analysis

    84 M. O. Waite et al.

    123

  • nymphal performance and survival (Coll 1996; Lund-

    gren and Fergen 2006; Lundgren 2011). As nymphs do

    not tend to move among plants and are thus restricted

    to the resources available on the plant on which they

    hatch, females must choose a plant that will support

    their development and survival. Survival and devel-

    opment time (from egg hatch to adult) are thus

    important measures of BP suitability. While each plant

    tested was equally suitable for oviposition, nymphal

    survival and development varied according to the BP

    species/cultivar tested. Nymphal development time

    was significantly different among plants. The fastest

    and slowest times differed by one day. Orius insidio-

    sus nymphs requiring one day longer to reach adult-

    hood is unlikely to be as biologically significant as

    plant species/cultivar, which had a much greater effect

    on the size of an O. insidiosus population, as observed

    in the ten week greenhouse trial. Nymphal survival on

    the test BP species/cultivars varied widely. Only

    10.7 % of nymphs reached the adult stage on marigold

    plants. Marigolds have been suggested as a potential

    banker plant for Orius sp., based on their observed

    abundance on the plants. However, the high numbers

    of Orius sp. observed in these trials may have been due

    to feeding upon thrips rather than pollen (Baggen et al.

    1999; Silveira et al. 2009). Buergi (2007) concluded

    that the nutritional profile of marigold pollen is not

    sufficient to support O. insidiosus reproduction.

    Marigold would be an excellent choice for female O.

    insidiosus to lay eggs if the plant was infested with

    prey and hence the suitability of the pollen would not

    be an important factor. However, among the criteria

    for selection of a BP for O. insidiosus is that it must

    provide an alternative source of food when prey is

    limited or absent. This renders marigold a poor

    candidate (Huang et al. 2011).

    The final research component evaluated the plant

    species/cultivars for their ability to support O. insid-

    iosus population growth over a ten week period. This

    provided more practical information on the potential

    of BPs in a commercial ornamental greenhouse as O.

    insidiosus were reared for multiple generations under

    greenhouse conditions. From the results of the labo-

    ratory bioassays, it might have been expected that

    gerbera daisy would be a good choice as a BP. Orius

    insidiosus nymphs reared on gerbera daisy had a

    survival rate of 58.1 % and development time

    (8.3 0.21 days) was significantly shorter than

    nymphs reared on the other plants. However, the

    nymphal development bioassays were carried out in

    ideal conditions with one pollen-producing gerbera

    flower per nymph. In contrast, the greenhouse bioas-

    say was conducted over a ten week period and the

    limited number of large flowers which are normally

    produced per gerbera plant (2.7 flowers plant-1

    observed, n = 8) was probably insufficient to support

    0

    5

    10

    15

    20

    25

    30

    35

    40

    45

    50

    55

    60

    65

    May 12 May 26 June 9 June 23 July 7Mea

    n N

    umbe

    r of O

    rius

    insi

    dios

    us N

    ymph

    s &

    Adul

    ts

    Purple Flash Black Pearl

    Castor Bean Feverfew

    Gerbera Marigold

    bb bc

    c

    d

    b

    b

    a

    d

    c c

    b b

    a

    c d d

    b bb

    a

    d d

    a

    bc c

    d d

    a

    Fig. 3 Mean number (SE) of Orius insidiosus nymphs andadults reared on six different host plants in a greenhouse

    bioassay (n = 8) conducted in Vineland ON, Canada, May 6th

    to July 7th, 2011. Means followed by the same letter within the

    same sampling period do not differ significantly from each other

    by Tukeys test (P \ 0.05)

    Evaluation of seven plant species/cultivars 85

    123

  • O. insidiosus population growth over the trial period.

    Weekly maintenance and pruning were conducted to

    encourage new growth and re-flowering, but over

    ten weeks, no new flowers were produced on the

    gerbera plants from weeks 7 through 10. In compar-

    ison, feverfew and marigold plants produce many

    small flowers (22.9 and 31.8 flowers plant-1

    observed, respectively). However, in both plants the

    number of O. insidiosus also declined despite the

    increased availability of pollen. The decline of O.

    insidiosus populations on feverfew and marigold may

    be attributed to the pollen having insufficient nutri-

    tional value (e.g., amino acid and lipids content) to

    sustain development of O. insidiosus rather than it

    simply being a factor of the quantity of pollen

    available (Buergi 2007; Schuel 1992). Future studies

    should address the quantity of pollen of the plant

    species/cultivars being tested in addition to pollen

    quality. Many flowers were consistently present on

    Purple Flash peppers (23.3 flowers plant-1, n = 8)

    and the number of O. insidiosus on this plant was

    significantly higher than on any other plants, including

    the Black Pearl pepper (7.9 flowers plant-1, n = 8),

    over the ten week observation period. Defensive

    properties of pollen also influence the ability of O.

    insidiosus to utilize pollen from different plant

    species/cultivars. Sporopollenin, which is a compo-

    nent of the outer wall of a pollen grain, is highly

    resistant to acids and enzymatic degradation (Schuel

    1992). Pollen containing higher levels of sporopol-

    lenin are, thus, likely to provide little nutritive value

    and will not promote survival and development. Orius

    insidiosus feeds on pollen in a similar manner to the

    way it feeds on prey, i.e. it inserts its rostral tip into the

    grain and extracts the inner contents as opposed to

    ingesting the entire grain (Fauvel 1974). While certain

    pollen may have a high intrinsic nutritive value, the

    thickness of the pollen wall and the sporopollenin

    content of the wall may prevent O. insidiosus access-

    ing these nutrients. So, while certain pollen sources

    may have high nutritive value, O. insidiosus may be

    unable to access these nutrients rendering the pollen

    unacceptable as a supplemental food.

    The key attribute measured for a suitable BP is its

    ability to support long term population growth of the

    biological control agents. Results of the current study

    suggest that the Purple Flash pepper has the greatest

    potential for use as a BP for O. insidiosus as it has

    superior characteristics for population growth

    compared with other species/cultivars tested. The

    Purple Flash ornamental pepper provided equally

    acceptable oviposition locations in comparison to all

    other plants tested, predator development time was

    equivalent to all other plants tested (9.1 0.21 days)

    with the exception of gerbera, and it supports popu-

    lation growth over multiple generations. Populations

    were consistently higher on Purple Flash than on any

    of the other plants tested in the greenhouse bioassays.

    In addition, the Purple Flash cultivar is a smaller, more

    compact plant that is better suited to placement on

    benches in ornamental greenhouses than the Black

    Pearl. Future studies should investigate the suitability

    of other ornamental pepper cultivars in addition to

    Purple Flash and Black Pearl, and it is important to

    assess their performance in a commercial greenhouse.

    When evaluating potential benefits of placing Black

    Pearl pepper BPs in commercial hoop houses follow-

    ing augmentative releases of O. insidiosus, Wong and

    Frank (2012) found that addition of the BPs did not

    improve the level of pest control obtained. Spiders

    colonized the BPs and reduced O. insidiosus access to

    pollen in the flowers. They concluded that BPs might

    be best suited for indoor greenhouse crops.

    In commercial greenhouse ornamental crops, Pur-

    ple Flash BPs supported constant populations of O.

    insidiosus (pers. obs, R.B.). Additional studies are now

    required to determine the dispersal of O. insidiosus

    through the greenhouse from the BPs, as well as

    determining effects on their searching behaviour when

    prey is scarce. The Purple Flash ornamental pepper is a

    promising BP for O. insidiosus that could enhance

    utilization of the predator to control WFT in green-

    house ornamental crops.

    Acknowledgments This work was conducted at the VinelandResearch and Innovation Centre, a not for profit organization

    dedicated to horticultural science and innovation, located in

    Canadas Niagara Region. We thank Biobest Canada Ltd. for

    supply of O. insidiosus, Angela Brommit, Rebecca Eerkes, Erik

    Glemser, and Andrew McFarlane for technical support. This

    research was funded through the OMAFRA-University of

    Guelph Research Program and a MITACS grant sponsored by

    Eco Habitat Agri-Services to M.O.W.

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    Author Biographies

    M. O. Waite This research is part of the M.Sc. project ofMeghann Waite which focuses on identifying new strategies to

    increase the control and cost efficiency of the biological control

    agent, Orius insidiosus Say, in greenhouse ornamentals.

    C. D. Scott-Dupree Her research focuses on the integratedmanagement of insect pests as well as the impact of

    agroecosystems on non-target beneficial arthropods.

    M. Brownbridge His research projects include the develop-ment and integration of microbial biocontrol strategies for

    insect pests and weeds.

    R. Buitenhuis She studies the use of predators, parasitoids,entomopathogenic nematodes, trap plants, and banker plants

    for the control of insect and mite pests.

    G. Murphy He provides technical support to growers in theNiagara region, specializing in integrated pest management

    solutions for greenhouse floriculture crops.

    Evaluation of seven plant species/cultivars 87

    123

    Evaluation of seven plant species/cultivars for their suitability as banker plants for Orius insidiosus (Say)AbstractIntroductionMethodsPlantsInsectsBioassaysEvaluation of plant species/cultivars as oviposition substrate for Orius insidiosusDevelopment of first instar Orius insidiosus nymphs to the adult stage on selected plant species/cultivarsThe effect of host plant on Orius insidiosus population growth

    Data analysis

    ResultsDiscussionAcknowledgmentsReferences