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BioControl (2014) 59:7987 DOI 10.

1007/s10526-013-9549-4

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 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 sunower (cv. Choco Sun) were evaluated for their suitability as banker plants (BP) for Orius insidiosus (Hemiptera: Anthocoridae) in commercial greenhouses. Oviposition, egg hatch, nymphal development to adulthood, and population increase were quantied 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. Nymphal 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 combined 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
Handling Editor: Patrick De Clercq. M. O. Waite (&) Monsanto Canada Inc., Guelph, ON N1G 0B4, Canada e-mail: Meghann.o.garlough@monsanto.com C. D. Scott-Dupree School of Environmental Sciences, University of Guelph, Guelph, ON N1G 2W1, Canada M. Brownbridge 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

In greenhouse ornamental crops, western ower thrips (WFT), Frankliniella occidentalis Pergande (Thysanoptera: 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 efcacious 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 oriculture greenhouses.

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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 owers 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 owering 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 sufcient 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 chrysanthemums, 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 owers, there are often few WFT in the vegetative stages of plant development. Secondly, a lack of alternative food sources as ornamentals typically are not owering 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 owering 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 sunower (cv. Choco Sun) plants to support a population of Orius insidiosus in commercial greenhouses. The plant species/cultivars were selected based upon growers observations 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 requirements 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/cultivars 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 rst 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 Vineland, ON Canada under growing conditions of 21 1 C, RH 70 5 %, 16: 8 h (L:D). Plant

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Evaluation of seven plant species/cultivars Table 1 Plant species/cultivars evaluated as potential banker plants for Orius insidiosus Common name Marigold, cv. Lemon Gem Castor Bean Ornamental Pepper, cv. Black Pearl Ornamental Pepper, cv. Purple Flash Gerbera Daisy, cv. Festival Species Tagetes patula Ricinus communis Capsicum annuum Capsicum annuum Gerbera jamesonii Source Veseys Seeds Ltd, Charlottetown, PE Richters Herbs, Goodwood, ON Stokes Seeds Ltd., Thorold, ON Stokes Seeds Ltd., Thorold, ON Orchard Park Growers, St. Catharines, ON Lakeshore Inc., Jordan Station, ON Feverfew Sunower, cv. Choco Sun Tanacetum parthenium Helianthus annuus Richters Herbs, Goodwood, ON Stokes Seeds Ltd., Thorold, ON

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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 (Benecial 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 owers, with the exception of gerbera and sunower in which each cup contained one ower. 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 lled 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-lled 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)]. After 48 h, the cages were removed from the chamber and the adults were removed from the cages. The number

All source locations are within Canada

species used in the BP trials were grown from seed with the exception of gerbera, which was purchased as a owering plant from commercial growers who did not use chemical insecticides. Seeds were hand sown two months prior to trial initiation into seeding trays lled with ProMix potting medium (Premier Tech ` re-du-Loup QC Canada). One Horticulture, Rivie 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 substrate 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 the Vineland Research and Innovation Centre. Adults were transferred to cages which were constructed from plastic containers (750 ml disposable Ziploc GladWare 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 provide ventilation. A handful of buckwheat hulls was placed into each container to serve as a refuge for the insects.

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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 (ve 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. Paralm 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 lter paper, which was placed on top of a vacuum ask and the ethanol was withdrawn. The lter paper was transferred 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 lter paper during the plant washing process. The oviposition assays were conducted in three blocks and replicated eight times per plant species/cultivar. Development of rst 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 were used in this study to ensure that all individuals had the same nutritional starting point. These rst 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. Catharines, 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 lled 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)] and nymphs 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 owering 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 ow rate) and fruit and dead owers removed weekly (ensuring no O. insidiosus were removed on plant matter). Each treatment received ten female and ve 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.

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Mean number of eggs Mean number of nymphs a a a a A A A ab A A b A A

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 separated using Tukeys multiple means comparison. The mean of the residuals was equal to zero and a Shapiro Wilk test conrmed that the residuals were approximately normally distributed. A Type I error rate (a) of 0.05 was used to test for signicance. 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 comparison. A Type I error rate (a) of 0.05 was used to test for signicance. A repeated measures ANOVA using the Mixed procedure, SAS v. 9.2 (SAS Institute, Cary, NC, USA), with variance partitioned into the xed 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 conrmed that the residuals were approximately normally distributed. Differences between means were determined using Tukeys multiple means comparison using a signicance level of a = 0.05.
Mean # of eggs or nymphs/ plant

50

40

30

20

10

Plant species/cultivar

Fig. 1 Mean number (SE) of Orius insidiosus eggs laid by ve 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 signicantly different according to Tukeys test (P [ 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). Signicantly fewer eggs (mean SE) were found on sunowers (14.9 7.1 eggs plant-1; P = 0.042) than marigold (49.9 6.4; P \ 0.0001). Differences in the number of eggs laid by O. insidiosus on marigold, gerbera, Black Pearl pepper, Purple Flash pepper, feverfew, and castor bean, were not signicant. 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 signicantly differ on sunowers 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 sunowers. Nymphal development time from rst 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 signicantly 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 significant difference in development times on feverfew, castor bean, Black Pearl pepper and Purple Flash pepper (Fig. 2). There were signicant 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 signicantly affected by plant species (F = 25.64; df = 5, 201; P \ 0.0001), week (F = 26.10; df = 4, 201; P \

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84 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 Gerbera daisy Black pearl pepper Purple ash pepper Feverfew Castor bean Lemon gem marigold Choco sun sunower
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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 rst 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 sunowers was signicantly lower than on plants other than gerbera. The number of eggs counted on the Choco Sun sunowers, 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 signicant 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 sunowers. This could be attributed to a severe powdery mildew (Blumeria graminis) infection that occurred on the sunower plants during the experiment. Sunowers were selected as a BP candidate as large Orius populations can be found in eld sunowers during summer months. However, owing to their susceptibility to powdery mildew, sunower 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 specic locations on a plant favour

na 31 32 40 45 41 140 15

Survival (%) 58.1 50.0 45.0 42.2 36.6 10.7 0.0

Initial number of nymphs

10 9
a

bc

c ab

Mean development time (days)

8 7 6 5 4 3 2 1 0

Plant species/cultivar

Fig. 2 Mean development time (days SE) of Orius insidiosus nymphs (\24 h old) to reach the adult stage when reared on six potential banker plant species/cultivars in a growth chamber at 25 1 C, R.H. 70 5 %, 16:8 h L:D. Means followed by the same letter are not signicantly different from each other by Tukeys test (P [ 0.05). No nymphs reached the adult stage on the sunowers (cv. Choco Sun) and these replicates were excluded from statistical analysis

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 signicantly higher (P \ 0.0001) than on any other plant species (Fig. 3). Marigold supported signicantly lower populations of O. insidiosus throughout (P \ 0.0001).

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Mean Number of Orius insidiosus Nymphs & Adults

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65 60 55 50 45 40 a 35 30 25 20 c 15 10 5 0 May 12 a b c b bc c d May 26 d d d c b b b b d d June 23 July 7 b b c c d d b b a a a Purple Flash Castor Bean Gerbera Black Pearl Feverfew Marigold

June 9

Fig. 3 Mean number (SE) of Orius insidiosus nymphs and adults 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 signicantly from each other by Tukeys test (P \ 0.05)

nymphal performance and survival (Coll 1996; Lundgren 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 development 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 signicantly different among plants. The fastest and slowest times differed by one day. Orius insidiosus nymphs requiring one day longer to reach adulthood is unlikely to be as biologically signicant 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 prole of marigold pollen is not sufcient 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 nal research component evaluated the plant species/cultivars for their ability to support O. insidiosus 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 laboratory 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 signicantly shorter than nymphs reared on the other plants. However, the nymphal development bioassays were carried out in ideal conditions with one pollen-producing gerbera ower per nymph. In contrast, the greenhouse bioassay was conducted over a ten week period and the limited number of large owers which are normally produced per gerbera plant (2.7 owers plant-1 observed, n = 8) was probably insufcient to support

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O. insidiosus population growth over the trial period. Weekly maintenance and pruning were conducted to encourage new growth and re-owering, but over ten weeks, no new owers were produced on the gerbera plants from weeks 7 through 10. In comparison, feverfew and marigold plants produce many small owers (22.9 and 31.8 owers 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 insufcient nutritional 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 owers were consistently present on Purple Flash peppers (23.3 owers plant-1, n = 8) and the number of O. insidiosus on this plant was signicantly higher than on any other plants, including the Black Pearl pepper (7.9 owers plant-1, n = 8), over the ten week observation period. Defensive properties of pollen also inuence the ability of O. insidiosus to utilize pollen from different plant species/cultivars. Sporopollenin, which is a component of the outer wall of a pollen grain, is highly resistant to acids and enzymatic degradation (Schuel 1992). Pollen containing higher levels of sporopollenin 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 accessing 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 population 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 benets of placing Black Pearl pepper BPs in commercial hoop houses following 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 owers. They concluded that BPs might be best suited for indoor greenhouse crops. In commercial greenhouse ornamental crops, Purple 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 greenhouse ornamental crops.
Acknowledgments This work was conducted at the Vineland Research and Innovation Centre, a not for prot 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.

References
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Author Biographies
M. O. Waite This research is part of the M.Sc. project of Meghann Waite which focuses on identifying new strategies to increase the control and cost efciency of the biological control agent, Orius insidiosus Say, in greenhouse ornamentals. C. D. Scott-Dupree Her research focuses on the integrated management of insect pests as well as the impact of agroecosystems on non-target benecial arthropods. M. Brownbridge His research projects include the development 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 the Niagara region, specializing in integrated pest management solutions for greenhouse oriculture crops.

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