Response of the Brown Stink Bug (: ) to the Aggregation Pheromone, Methyl (2E,42)-Decadienoate1

T. C. Leskey and H. W. Hogmire3

USDA—ARS, Appalachian Fruit Research Station, Kearneysville, West Virginia USA

J. Entomol. Sci. 42(4): 548-557 (October 2007) Abstract Field-based experiments were conducted to evaluate the response of the abundant brown stink bug, Eusch,stus servus (Say), to commercially available pheromone lures containing methyl (2E,42)-decadienoate deployed in association with yellow pyramid traps. servus aggregated over a zone of at least 3.14 m 2 based on significantly greater numbers located on mullein plants located 1 m from baited traps compared with plants at 5 and 10 m. At this distance, —96% of all adults located on mullein plants were not subsequently captured by baited traps. However, the presence of mullein plants near baited traps did not significantly reduce baited trap captures. Even if mullein plants were not present, baited trap captures remained statistically identical. Based on all adults captured in baited traps and located on surrounding mullein plants, 50% of all individuals that entered plots were captured in traps.

Key Words stink bug, aggregation pheromone, monitoring, traps, apple, peach

Several stink bug species, including the brown stink bug, Euschistus servus (Say), are serious pests of stone fruit (Hogmire 1995) and are becoming increasingly im- portant pests of apple (Brown 2003, Pfeiffer 2005). The potential for stink bugs to become even more problematic could be influenced by additional cancellations or restrictions placed on broad spectrum insecticides based on the Food Quality Pro- tection Act. Economic damage caused by stink bugs in fruit orchards could increase as narrow-spectrum insecticides replace broad spectrum materials for control of key insect pests. To effectively manage stink bugs in a narrow-spectrum, reduced-spray environment, it is imperative that treatments be triggered by a monitoring system designed to detect increases in stink bug abundance or activity. Approaches evaluated to monitor stink bug activity in fruit orchards have included: (1) beating tray samples (Leskey and Hogmire 2005, Ohlendorf 1999); (2) sweep net samples (Leskey and Hogmire 2005); (3) examination of broadleaf weed hosts (Ohlendorf 1999); (4) direct observation of fruit injury (Ohlendorf 1999); and (5) trap- ping samples. Trap types evaluated in fruit orchards have included: (1) plastic jar traps deployed from branches in tree canopies (Leskey and Hogmire 2005); (2) tube traps constructed with wire mesh cone funnels at either end (Krupke et al. 2001, Ohlendorf 1999); and (3) yellow pyramid traps deployed on the ground between trees

Received 05 December 2006; accepted for publication 17 January 2007. 2Address inquiries (email: [email protected]). West Virginia University, Tree Fruit Research and Education center, P0 Box 609, Kearneysville, WV 25430

548 LESKEY AND HOGMIRE: Stink Bug Responses to Pheromone 549

(Hogmire and Leskey 2006, Leskey and Hogmire 2005, Johnson et al. 2002) and on horizontal limbs within tree canopies (Hogmire and Leskey 2006) in the border row of fruit orchards. These traps are often baited with methyl (2E,4Z)-decadienOate, the major male- produced volatile produced by a number of Nearctic Euschistus sop. (Aldrich et al. 1991). In field tests conducted in Maryland, the presence of this compound in asso- ciation with traps increased the number of Euschistus spp. captured in or near baited traps (Aldrich et al. 1991). In Washington, E. conspersus, an important pest in apple orchards in western North America, was attracted to the immediate area surrounding traps baited with methyl (2E, 42)-decadienoate, but few individuals were captured in traps themselves (Krupke et al. 2001). When these same lures were attached to of E. mullein plants, Verbascum thapsus L., a favored host plant, greater numbers conspersus were recovered from these plants compared with unbaited plants (Krupke et al. 2001). In mid-Atlantic fruit orchards, the addition of this odor stimulus in traps was proven effective at significantly increasing captures of the more prevalent brown (Leskey and Hogmire stink bug, E. servus, and the dusky stink bug, E. tristigmus 2005), as well as increasing the number of individuals observed within the vicinity of baited traps (Leskey and Hogmire, unpubl. data). These field observations raise the possibility that some Euschistus spp. attracted by the presence of methyl (2E,42)-decadienoate lures may never be captured in baited traps. If this is the case, trap-based monitoring approaches could underesti- mate the size of the wild population. Therefore, we conducted a series of field-based experiments which evaluated the response of the most abundant Euschistus spp. in midAtlantic fruit orchards, the brown stink bug, E. servus, to methyl (2E, 4Z)- decadienoate to learn: (1) if adults aggregate in a zone surrounding pyramid traps baited with commercially available pheromone lures; (2) what proportion of adults attracted by the presence of pheromone lures in baited traps were subsequently captured; and (3) if the presence of favored host plants near baited traps acted as a harborage and reduced the likelihood of trap capture.

Materials and Methods

Aggregation zone. Six experimental plots were established in a 5-ha open field area planted in mixed fescue and surrounded by hedgerows, small wood lots, and apple orchard blocks at the Appalachian Fruit Research Station (AFRS) in Kear- neysville, WV. In addition, 6 plots were established in a 3-ha open field area planted in mixed fescue and surrounded by a small pine woodlot and peach orchards at the West Virginia Univ. Tree Fruit Research and Education Center (WVU) in Kear- neysville, WV. Groundcover was trimmed to a height of -10 cm for these studies. Each plot was 20 x 20 m and separated from other plots by at least 20 m. A single pyramid trap constructed of yellow Coroplast (AIN Plastics, VA Beach, VA) with a modified jar top (Hogmire and Leskey 2006), was baited with lures containing 200 mg of methyl (2E,4Z)-decadienOate (Advanced Pheromone Technologies, Inc., Mary- hurst, OR, now APTl y , Inc., Portland, OR) or left unbaited and deployed at the center of each plot. Four potted mullein plants were placed in each of 3 circular subplots of increasing diameter (2, 10, and 20 m) around each trap. Plants were spaced equi- distantly from one another in each circular subplot, with plants in 2 and 20 m subplots placed at each cardinal direction and those at 10 m off-set by 450 (Fig. 1). Thus, plants were located 1, 5, and 10 m from traps based on 2, 10, and 20 m circular subplots. -

550 J. Entomoj. Sci. Vol. 42, No. 4 (2007)

20m

Fig. 1. Single experimental plot with centrally located pyramid trap surrounded by four potted mullein plants in each circular subplot of increasing diameter (2, 10, and 20 m).

Three plots containing baited traps and 3 plots containing unbaited traps were in- stalled on 19 July 2005 and sampled for 5 consecutive days at AFRS. At WVU, traps and plants were installed on 18 July 2005 and sampled for 6 consecutive days. The number of bugs located on each plant was counted, and bugs were left on plants. Bugs in traps were counted and removed daily. Nontransformed data, as the homo- geneity-of-variances assumption was met in all cases, were analyzed using the GLM procedure (SAS Institute 2001) to construct analysis of variance (ANOVA) tables for cumulative numbers of bugs located over the entire sampling period on plants at different distances from baited and unbaited traps. The model included the following class variables: presence of bait (BAIT), distance from traps (DIS) and the interaction term (BAITxDIS). Data were subsequently analyzed using the GLM procedure (SAS Institute 2001) to construct analysis of variance (ANOVA) tables for the number of servos E. recovered from plants at different distances from baited traps over the entire sampling period. The number of bugs recovered from baited and unbaited traps across all plots was compared using a two sample f-test (SAS Institute 2001). Adult attraction and capture. Six plots were established at AFRS and WVU with baited and unbaited traps and mullein plants identical to protocols described above. Plants and traps were deployed from 26-30 July 2005 at AFRS and from 25-29 July 2005 at WVU. After an initial 48-h period to allow bugs to move into plots and colonize mullein plants, all adult male and female bugs were located on plants, counted, and 551 LESKEY AND HOGMIRE: Stink Bug Responses to Pheromone marked on their scutellum with a unique dot of paint using DecoColor TM pens (Uchida of America Corp. Torrance, CA) to indicate sex and plot location. All marked bugs were left on plants. The number of marked and unmarked bugs subsequently located on each plant was recorded 24 and 48 h later. Bugs in traps were counted and removed daily. The number of bugs recovered from baited and unbaited traps across all plots was compared using a two sample t-test (SAS Institute 2001). Presence of host plants. Six plots were established at AFRS and WVU based on protocols described previously. In this case, the pyramid trap deployed at the center of every plot was baited. In 3 plots, mullein plants were not deployed (no potential host plant harborages available), whereas in the other 3 plots mullein plants were posi- tioned as previously described (host plant harborages available). Plants and traps were deployed from 9-12 Aug and again from 24-27 Aug 2005 at AFRS, and from 2-6 Aug 2005 at WVU. In plots with mullein plants, the number of bugs located on each plant was counted, and bugs were left on plants. Bugs in baited traps were counted and removed daily. The number of bugs recovered from baited traps in plots with and without mullein plants was compared using a two sample t-test (SAS Institute 2001).

Results

Aggregation zone. At AFAS, stink bug presence on mullein plants was significant (P= 0.05), DIS (P= 0.05), and (F= 5.32; df = 5,12; P= 0.0083) with the effect of BAIT the interaction term (BAITxDIS) (P = 0.02) all being significant. Overall, significantly more E. servus were located on plants in plots with baited traps (67.0 ± 20.2 SE) compared with unbaited traps (33.1 ± 8.8 SE). Significantly more E. sen/us were recovered from plants 1 m from baited traps compared with plants at all other dis- tances from traps, with the exception of plants 10 m from unbaited traps (Table 1). The mean number of stink bugs ± SE captured in baited traps (55.3 ± 15.5 SE) was significantly greater than unbaited traps (11.3 ± 4.3 SE) (t = 2.787, df = 4, P < 0.05). At WVU, stink bug presence on mullein plants was significant (F= 8.64; df = 5,12; < 0.01), and the interaction term <0.01) with the effect of BAIT (P <0.01), DIS (P P E. sen/us (BAITxDIS) (P = 0.0066) all being significant. Overall, significantly more were located on mullein plants in plots with baited traps (26.3 ± 8.2 SE) compared with unbaited traps (5.2 ± 2.4 SE). Significantly more E. seus were recovered from plants 1 m from baited traps compared with plants at all other distances from traps (Table 1). The mean number of stink bugs ± SE captured in baited traps (11.3 ± 2.8 SE) was significantly greater than unbaited traps (1.3 ± 0.8 SE) (t = 3.354; df = 4; P< 0.05). E. senius were cap- Adult attraction and capture. At AFRS, significantly more tured in baited traps (17.0 ± 4.9 SE) compared with unbaited traps (3.0 ± 2.1 SE) (t= 2.62; df = 4; p = 0.05) over the entire 96 h period. In plots containing baited traps, 35% of all adults were captured by traps; whereas, 65% were located on mullein plants during the initial colonization period. In plots containing unbaited traps, 27.2% of all adults recorded were captured by traps, whereas 72.8% were located on mullein plants. A total of 102 adults was located on mullein plants and marked in plots = 24) traps, respectively. Twenty-four hours containing baited (n = 78) and unbaited (n after being marked, 68.7% of males and 56.7% of females had left mullein plants located 1 m from baited traps (Table 2). This percentage increased to 93.7% for males and 86.7% for females after 48 h. Dispersal from mullein plants located 10 m

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Table 1. Mean (± SE) number of adult E. servus located on mullein plants at 1, 5, and 10 m from baited or unbaited pyramid traps in an open field in experimental plots at AFRS and at WVU

Site Treatment Distance Mean ± SE AFRS Baited 1 135.3 ± 34.Oa Baited 5m 37.7 ± 10.7b Baited 10 m 28.0 ± 10.3b Unbaited 1 27.3 ± 10.9b Unbaited 5m 22.0± 11.0b Unbaited 10m 50.0 ± 21.6ab wvu Baited 1 54.3 ± 10.Oa Baited 5m 15.3 ± 7.5b Baited 10 m 9.3 ± 7.Ob Unbaited 1 3.0 ± 3.Ob Unbaited 5m 9.0 ± 6.5b Unbaited 10 m 3.7 ± 2.7b Means in within a column within a site followed by a different letter are significantly different HSD test). (P = 0.05; Tukeys

from baited traps reached 100% for marked males and females after 48 h (Table 2). Dispersal from plants located 1 m from unbaited traps reached 100% for males after 24 h and for females after 48 h (Table 2). Among all marked adults in plots containing baited traps, only 3.8% were subsequently captured in baited traps, whereas 96.2% were not. Those captured consisted of 3 females marked on plants located 1 m from baited traps (10.3% of all individuals marked on plants located 1 m from baited traps). None of the marked adults within plots containing unbaited traps were subsequently captured. Plant-to-plant within-plot movement was recorded for 6 marked adults (7.6% of the marked population) in plots with baited traps, and for 5 marked adults (20.8% of the marked population) in plots with unbaited traps over the 48 h period. Plot-to-plot movement was recorded for 1 individual; 1 adult male marked on a plant 5 m from an unbaited trap was subsequently captured in a neighboring plot on a plant 1 m from a baited trap. At WVU, significantly more E. servus were captured in baited traps (10.0 ± 2.0 SE) compared with unbaited traps (0.7 ± 0.7 SE) (t= 4.427; df = 4; P= 0.01) over the entire 72 h period. In plots containing baited traps, 44% of all adults were captured by traps; whereas 56% were located on mullein plants during the initial colonization period. In plots containing unbaited traps, 20% of all adults recorded were captured by traps, whereas 80% were located on mullein plants. A total of 27 adults was located on mullein plants and marked in plots containing baited (n = 19) and unbaited (n = 8) traps, respectively. Twenty-four hours after being marked, 75.0% of males and 33.3% of females had left mullein plants located 1 m from baited traps (Table 2). This percentage Increased to 93.7% for males and remained the same for females after

553 LESKEY AND HOGMIRE: Stink Bug Responses to Pheromone

Table 2. Total number (N) of male and female E. servus that were marked on mullein plants located 1, 5, and 10 m from baited or unbaited pyramid traps in an open field, and the percentage (%) remaining, and total number of wild, unmarked adults found 24 and 48 h later in experi- mental plots at AFRS and WVU

Male Female

24h 48 h Site Treatment Distance N 24 h 48h N 30 43.3%,10 13.3%,4 AFRS Baited 1 m 16 31.3%,2 6.3%,0 25.0%,2 Baited 5m 11 9.1%,1 9.1%, 1 8 50.0%,5 6 16.7%,2 0.0%,l Baited 10m 7 14.3%,1 0.0%,0 6 33.3%,2 0.0%,2 Unbaited 1 m 1 0.0%,0 0.0%,0 5 0.0%,2 0.0%, 1 Unbaited 5 m 2 0.0%,l 0.0%,l 20.0%,l Unbaited 10 m 5 0.0%,2 20.0%,2 S 0.0%,1 66.6%,10 WVU Baited 1 4 25.0%,6 6.3%,4 3 66.6%,7 S 20.0%,2 20.0%,4 Baited 5 6 66.6%,l 16.7%,2 —,0 Baited 10 m 0 —,0 1 50.0%,l Unbaited 1 0 —,1 —,2 2 50.0%,0 4 50.0%,2 25.0%,6 Unbaited 5 2 50.0%,3 50.0%,3 —,1 —,0 Unbaited 10 m 0 —,0 —,0 0

The total number of marked individuals, the percentage (%) of marked individuals recovered, and the total number of wild individuals found on mullein plants.

48 h. Dispersal from mullein plants located 5 m from baited traps reached 83.3% for marked males and 80.0% for marked females after 48 h. No adults were marked on mullein plants located 10 m from baited traps (Table 2). In plots with unbaited traps, dispersal reached 50% for males and 75% for females after 48 h for those that were marked on plants located 5 m from unbaited traps. No adults were marked on mullein plants located 1 and 10 m from unbaited traps (Table 2). Among all marked adults in plots containing baited and unbaited traps, none were subsequently captured in traps. Plant-to-plant within-plot movement was recorded for 2 marked adults (10.5% of the marked population) in plots with baited traps and none with unbaited traps. No plot- to-plot movement was recorded. captured per Presence of host plants. At AFRS, the mean number of E. servus baited trap was not significantly different (t= 0.142; df = 4; P= 0.89) for traps in plots with mullein plants present (16.00 ± 4.36 SE) compared with plots without mullein plants (15.33 ± 1.76 SE). Each day, 2.00 ± 0.55 SE and 1.92 ± 0.22 SE stink bugs were captured per trap in plots with and without mullein plants, respectively. The number of bugs observed per mullein plant per day was 0.45 bugs/plant (or 5.4 bugs per plot per day). At distances of 1, 5, and 10 m, the number of bugs observed per mullein plant per day was 0.53, 0.48, and 0.36 bugs/plant, respectively. At WVU, the mean number of E. servus captured per baited trap was not signifi- MR

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cantly different (t= 1.992; df =4; P= 0.12) for traps in plots with mullein plants present (7.3 ± 3.0 SE) compared with plots without mullein plants (14.7 ± 2.2 SE). Each day, 1.8 ± 0.08 and 3.7 ± 0.8 stink bugs were captured per trap in plots with and without mullein plants, respectively. The number of bugs observed per mullein plant per day was 0.11 bugs/plant (or 1.33 bugs per plot per day). At distances of 1, 5, and 10 m, the number of bugs observed per mullein plant per day was 0.15, 0.19, and 0.00 bugs/plant, respectively.

Discussion

Euschistus servus aggregated over a zone of at least 3.14 m 2 surrounding pyramid traps baited with commercially available pheromone lures containing 200 + mg methyl (2E, 42)-decadienoate based on increased numbers recovered from mullein plants located 1 m from baited traps compared with plants at 5 and 10 m from baited traps (Table 1). Methyl (2E, 4Z)-decadienoate appears to act as an attractant, eliciting both male and female stink bug orientation to the stimulus source based on increased captures in baited traps compared with unbaited traps. However, this chemical stimu- lus may also induce arrestment prior to trap entry, as few marked stink bugs located on mullein plants 1 m from baited traps were subsequently captured in traps. Kennedy (1978) indicated that an attractant can become an arrestant when the chemical stimu- lus is at a high concentration and as the insect nears the stimulus source. Perhaps, as E. servus oriented to the source of methyl (2E, 42)-decadienoate, they may have encountered a concentration sufficient to slow or stop movement prior to reaching traps. In our studies, >96% of all adults located on mullein plants were not subse- quently captured in baited traps. Krupke et al. (2001) observed E. conspersus in large numbers in the immediate area surrounding traps [baited with a formulation of methyl (2E, 4Z)-decadienoate identical to that reported here], but captured few adults in traps. If release rates were lowered, thereby reducing the concentration of stimulus encountered, would more stink bugs reach the traps? This is a possibility, although this question was not evaluated here. Gravimetric studies in our laboratory revealed that methyl (2E, 4Z)-decadienoate lures released on average - .453 ug/h for day 1 and then —160 ug/h for the next 38 d [similar to results reported by Krupke et al. (2001) in which lures released at 300 ug/h for day 1 and -. 150 ug/hr for days 6-13]. On the other hand, E. servus found on mullein plants surrounding baited traps may have responded to secondary stimuli rather than being arrested by high concentra- tions of methyl (2E, 42)-decadienoate. In other stink bug species, such as Nezara virdula (L.), vibrational signals on host plants provide directional cues for mate loca- tion and stimulate searching behavior (çokl et al. 1999). Pheromones likely mediate long-range mate searching for some pentatomid species, including Thyanta spp. (McBrien et al. 2002a), but vibrational cues may become more important over shorter distances (McBrien et al. 2002b). In general, females will call with a species-specific repertoire of pulses which is followed by a male-produced courtship song (Moraes et al. 2005). Similarly, methyl (2E, 42)-decadienoate attracts Euschistus spp. and may mediate mate-finding over longer distances. Aggregations of E. conspersus adults formed and mating occurred on mullein plants baited with this chemical stimulus (Krupke et al. 2006). Furthermore, it is known that both male and female E. consper- sus produce vibrational-borne signals (McBrien and Millar 2003) as do E. heros (Moraes et al. 2005). Like E. conspersus, we observed aggregation formation and mating on mullein plants within plots containing methyl (2E, 4 7)-decadienoate lures. LESKEY AND HOGMIRE: Stink Bug Responses to Pheromone 555

In general, adults aggregated on plants late in the afternoon, began mating, and remained in copula until the following morning (Leskey and Hogmire, unpubi. data). Perhaps, this male-produced aggregation pheromone mediates long-range mate find- ing for Euschistus spp., but once on host plants, other cues such as vibrational signals or visual stimuli elicit short-range mating behaviors. Identification of vibrational songs for E. servus could improve our ability to successfully attract and trap these species. In a practical sense, the question becomes how to deploy methyl (2E, 42)- decadienoate in cropping systems. Methyl (2E,42)-decadienOate lures serve to ag- Krupke et al. 2001) over gregate E. servus (results reported here) and E. conspersus ( a zone larger than that occupied by a baited trap. However, our results demonstrated that baited pyramid traps captured on average at least 4x as many adults as were found on individual mullein plants. However, recovery from individual mullein plants could be much higher compared with baited traps if plants were baited directly, as reported by Krupke et al. (2001). More importantly, baited trap or plant samples should reflect the overall relative size of the wild population if they are to be useful in pest management decision-making. Cullen and Zalom (2006) found no significant relationships between baited trap captures (commercially available jar trap with in- verted funnels) and canopy shake samples of E. conspersus in tomato fields, indi- cating that trap captures did not necessarily reflect the population density. However, they deployed a trap design that likely was ineffective at capturing E. conspersus. Leskey and Hogmire (2005) compared the jar trap with the pyramid trap and dem- onstrated that pyramid traps captured significantly more Euschistus spp. in mid- Atlantic fruit orchards. Captures in baited pyramid traps also reflected species abun- dance in apple orchards based on significant positive Pearsons correlations between biweekly trap and tree beating samples in particular, although this relationship was not significant in peach orchards (Leskey and Hogmire 2005). To establish a treatment threshold for stink bugs in fruit orchards, it is important to determine if relative numbers of trap captures reflect the amount of fruit injury in- curred. This could be difficult to establish with a trap-based approach because so many individuals remain uncaptured and adults aggregate in a zone extending sev- eral meters beyond a baited trap. However, this documented aggregation response could provide a novel trap-tree approach for monitoring Euschistus spp., similar to what has been developed for the plum curculio, Conotrachelus nenuphar ( Herbst), another direct pest of tree fruit. Deployment of the aggregation pheromone of plum curculio and a synthetic fruit volatile within canopies of apple trees led to aggregated and increased injury in so-called trap trees compared with unbaited tree canopies in the perimeter row of apple orchards (Prokopy et al. 2003). This trap tree monitoring technique, aggregating injury within particular odor-baited sentinel trees for visual inspection, was used to develop a protocol for determining the need for and timing of insecticide applications after petal fall for plum curculio in New England apple or- chards (Prokopy et al. 2004). Similarly, mullein plants baited with methyl (2E, 42)- decadienoate were suggested as a potential monitoring tool for E. conspersus (Krupke et al. 2001), although in this case presence of adults on a noncrop plant would be monitored rather than injury on the crop itself. A trap-tree approach may be effective for E. senius particularly because this species aggregates in large numbers within the vicinity of a pheromone lure. However, comparisons with a trap-based approach and correlations with injury would need to be assessed to determine which methodology may best serve as the most effective decision-making tool. 556 J. Entomol. Sci. Vol. 42, No. 4 (2007)

Acknowledgments

The authors thank Stephen Berg, Deborah Blue, Emma Bowers, Torri Hancock, Tim Win- field, and Starker Wright for excellent technical assistance and Drs. Peter Shearer and Chris Bergh for reviewing an earlier version of this manuscript. Mention of trade names or commercial products in this publication is solely for the purpose of providing specific information and does not imply recommendation or endorsement by the U.S. Department of Agriculture or West Virginia University.

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