Dawn M. Olson

Nitrogen fertilization rate affects feeding, larval performance, and oviposition preference of the beet armyworm, Spodoptera exigua, on cotton

Nitrogen (N) is one of the most critical chemical elements for plant and animal growth, exerting a variety of bottom‐up effects. Development and oviposition of the beet armyworm, Spodoptera exigua (Hübner) (Lepidoptera: Noctuidae), were studied in relation to varying N fertilization levels (42, 112, 196, and 280 p.p.m.) in cotton [Gossypium hirsutum L. (Malvaceae)]. Low N fertilization of cotton plants led to reduced plant biomass and a lower percentage of N in leaf blades and in leaf petioles. Development of S. exigua larvae fed with plants with reduced N applications (42 and 112 p.p.m.) was prolonged relative to treatments receiving higher N fertilization. Almost all larvae reared on artificial diets underwent only five instars before pupation. However, most larvae reared on cotton plants, irrespective of N levels, experienced a supernumerary sixth larval instar. Furthermore, significantly more larvae reared on lower N cotton plants underwent supernumerary development compared to larvae reared on higher N cotton plants. Life‐time feeding damage per larva ranged from 55 to 65 cm2, depending on the nutritional quality of the food plant, although the differences were not statistically significant. Larvae distinguished between cotton plants with various nutritional qualities and fed preferentially on higher N plants. Female moth oviposition choice was also affected by host plant nutritional quality: cotton plants with higher N levels were preferentially chosen by S. exigua females for oviposition. The mechanisms of these effects are unclear, but they can have important implications for population dynamics and pest status of beet armyworms in the field.

Effects of nitrogen fertilization on tritrophic interactions

Tritrophic interactions (plant—herbivore—natural enemy) are basic components of nearly all ecosystems, and are often heavily shaped by bottom-up forces. Numerous factors influence plants’ growth, defense, reproduction, and survival. One critical factor in plant life histories and subsequent trophic levels is nitrogen (N). Because of its importance to plant productivity, N is one of the most frequently used anthropogenic fertilizers in agricultural production and can exert a variety of bottom-up effects and potentially significantly alter tritrophic interactions through various mechanisms. In this paper, the potential effects of N on tritrophic interactions are reviewed. First, in plant-herbivore interactions, N availability can alter quality of the plant (from the herbivore’s nutritional perspective) as food by various means. Second, nitrogen effects can extend directly to natural enemies through herbivores by changes in herbivore quality vis-à-vis the natural enemy, and may even provide herbivores with a defense against natural enemies. Nitrogen also may affect the plant’s indirect defenses, namely the efficacy of natural enemies that kill herbivores attacking the plant. The effects may be expressed via (1) quantitatively and/or qualitatively changing herbivore-induced plant volatiles or other plant features that are crucial for foraging and attack success of natural enemies, (2) modifying plant architecture that might affect natural enemy function, and (3) altering the quality of plant-associated food and shelter for natural enemies. These effects, and their interactive top–down and bottom-up influences, have received limited attention to date, but are of growing significance with the need for expanding global food production (with accompanying use of fertilizer amendments), the widening risks of fertilizer pollution, and the continued increase in atmospheric CO2.

Influence of Cover Crops on Insect Pests and Predators in Conservation Tillage Cotton

Abstract In fall 2000, an on-farm sustainable agricultural research project was established for cotton, Gossypium hirsutum L., in Tift County, Georgia. The objective of our 2-yr research project was to determine the impact of several cover crops on pest and predator insects in cotton. The five cover crop treatments included 1) cereal rye, Secale cereale L., a standard grass cover crop; 2) crimson clover, Trifolium incarnatum L., a standard legume cover crop; 3) a legume mixture of balansa clover, Trifolium michelianum Savi; crimson clover; and hairy vetch, Vicia villosa Roth; 4) a legume mixture + rye combination; and 5) no cover crop in conventionally tilled fields. Three main groups or species of pests were collected in cover crops and cotton: 1) the heliothines Heliothis virescens (F.) and Helicoverpa zea (Boddie); 2) the tarnished plant bug, Lygus lineolaris (Palisot de Beauvois); and 3) stink bugs. The main stink bugs collected were the southern green stink bug, Nezara viridula (L.); the brown stink bug, Euschistus servus (Say); and the green stink bug, Acrosternum hilare (Say). Cotton aphids, Aphis gossypii Glover, were collected only on cotton. For both years of the study, the heliothines were the only pests that exceeded their economic threshold in cotton, and the number of times this threshold was exceeded in cotton was higher in control cotton than in crimson clover and rye cotton. Heliothine predators and aphidophagous lady beetles occurred in cover crops and cotton during both years of the experiment. Geocoris punctipes (Say), Orius insidiosus (Say), and red imported fire ant, Solenopsis invicta Buren were relatively the most abundant heliothine predators observed. Lady beetles included the convergent lady beetle, Hippodamia convergens Guérin-Méneville; the sevenspotted lady beetle, Coccinella septempunctata L.; spotted lady beetle, Coleomegilla maculata (DeGeer); and the multicolored Asian lady beetle, Harmonia axyridis (Pallas). Density of G. punctipes was higher in cotton fields previously planted in crimson clover compared with control cotton fields for all combined sampling dates in 2001. Intercropping cotton in live strips of cover crop was probably responsible for the relay of G. punctipes onto cotton in these crimson clover fields. Density of O. insidiosus was not significantly different between cover crop and control cotton fields. Lady beetles seemed to relay from cover crops into cotton. Conservation of the habitat of fire ants during planting probably was responsible for the higher density of red imported fire ants observed in all conservation tillage cotton fields relative to control cotton fields. Reduction in the number of times in which economic thresholds for heliothines were exceeded in crimson clover and rye compared with control fields indicated that the buildup of predaceous fire ants and G. punctipes in these cover crops subsequently resulted in reduction in the level of heliothines in conservation tillage cotton with these cover crops compared with conventional tillage cotton without cover crops.

From Asian curiosity to eruptive American pest: Megacopta cribraria (Hemiptera: Plataspidae) and prospects for its biological control

The kudzu bug or bean plataspid, Megacopta cribraria (Fabricius), is native to Asia where it appears to be widely distributed (although the taxonomy is not entirely clear), but is infrequently a pest of legumes. This bug appeared in 2009 in the southeastern United States, where it is closely associated with kudzu, Pueraria montana Lour. [Merr.] variety lobata [Willd.] Maesen & S. Almeida. However, the insect has become a consistent economic pest of soybeans, Glycine max (L.) Merr., and some other leguminous crops in areas where large numbers can build in kudzu, in addition to being a considerable nuisance in urban landscapes where kudzu occurs. The insect has remarkable capacity for movement and has spread rapidly from nine Georgia counties in 2009 to seven states in 2012. Despite being a nuisance in urban areas and a crop pest, high populations of the bug also reduce the biomass of kudzu, which is itself a seriously problematic invasive weed, complicating the status of M. cribraria in its expanded range. Extant predators and a pathogen in the US have been observed attacking kudzu bugs in the laboratory and field, but no parasitism of eggs or nymphs has been observed to date. A single record of parasitism of an adult kudzu bug by a tachinid fly is known from the US, but no other adult parasitism has been observed in the US or elsewhere. Extant enemies may eventually significantly reduce the bug’s populations, but at present native enemies appear to be insufficient for the task, and exotic enemies from the kudzu bug’s native range may offer the best possibility for effective biological control in the US. Based on the available literature, the best option for an importation biological control program appears to be the platygastrid egg parasitoid Paratelenomus saccharalis (Dodd) because of its apparent host specificity, intimate biological linkages with M. cribraria, and wide geographic distribution in the Eastern Hemisphere. Other natural enemies may eventually emerge as good candidates for importation, but at present P. saccharalis appears to be the most promising.

Resistance mechanisms against arthropod herbivores in cotton and their interactions with natural enemies

Cotton plants (genus Gossypium) are grown on more than 30 million hectares worldwide and are a major source of fiber. The plants possess a wide range of direct and indirect resistance mechanisms against herbivorous arthropods. Direct resistance mechanisms include morphological traits such as trichomes and a range of secondary metabolites. The best known insecticidal compounds are the terpenoid gossypol and its precursors and related compounds. Indirect resistance mechanisms include herbivore-induced volatiles and extrafloral nectaries that allow plants to attract and sustain natural enemy populations. We discuss these resistance traits of cotton, their induction by herbivores, and their impact on herbivores and natural enemies. In addition, we discuss the use of genetically engineered cotton plants to control pest Lepidoptera and the influence of environmental factors on the resistance traits.

Pest trade-offs in technology: reduced damage by caterpillars in Bt cotton benefits aphids

The rapid adoption of genetically engineered (GE) plants that express insecticidal Cry proteins derived from Bacillus thuringiensis (Bt) has raised concerns about their potential impact on non-target organisms. This includes the possibility that non-target herbivores develop into pests. Although studies have now reported increased populations of non-target herbivores in Bt cotton, the underlying mechanisms are not fully understood. We propose that lack of herbivore-induced secondary metabolites in Bt cotton represents a mechanism that benefits non-target herbivores. We show that, because of effective suppression of Bt-sensitive lepidopteran herbivores, Bt cotton contains reduced levels of induced terpenoids. We also show that changes in the overall level of these defensive secondary metabolites are associated with improved performance of a Bt-insensitive herbivore, the cotton aphid, under glasshouse conditions. These effects, however, were not as clearly evident under field conditions as aphid populations were not correlated with the amount of terpenoids measured in the plants. Nevertheless, increased aphid numbers were visible in Bt cotton compared with non-Bt cotton on some sampling dates. Identification of this mechanism increases our understanding of how insect-resistant crops impact herbivore communities and helps underpin the sustainable use of GE varieties.

Impact of Brown Stink Bug (Heteroptera: Pentatomidae) Feeding on Corn Grain Yield Components and Quality

ABSTRACT Brown stink bug, Euschistus servus (Say) (Heteroptera: Pentatomidae), damage on developing corn, Zea mays L., ears was examined in 2005 and 2006 by using eight parameters related to its yield and kernel quality. Stink bug infestations were initiated when the corn plants were at tasseling (VT), mid-silking (R1), and blister (R2) stages by using zero, three, and six in 2005 or zero, one, two, and four bugs per ear in 2006, and maintained for 9 d. The percentage of discolored kernels was affected by stink bug number in both years, but not always affected by plant growth stage. The growth stage effect on the percentage of discolored kernels was significant in 2006, but not in 2005. The percentage of aborted kernels was affected by both stink bug number and plant growth stage in 2005 but not in 2006. Kernel weight was significantly reduced when three E. servus adults were confined on a corn ear at stage VT or R1 for 9 d in 2005, whereas one or two adults per ear resulted in no kernel weight loss, but four E. servus adults did cause significant kernel weight loss at stage VT in 2006. Stink bug feeding injury at stage R2 did not affect kernel damage, ear weight or grain weight in either year. The infestation duration (9 or 18 d) was positively correlated to the percentage of discolored kernels but did not affect kernel or ear weight. Based on the regression equations between the kernel weight and stink bug number, the gain threshold or economic injury level should be 0.5 bugs per ear for 9 d at stage VT and less for stage R1. This information will be useful in developing management guidelines for stink bugs in field corn during ear formation and early grain filling stages.

Colonization preference of Euschistus servus and Nezara viridula in transgenic cotton varieties, peanut, and soybean

Producers of Bt cotton, Gossypium hirsutum L. (Malvaceae), in the southeastern USA face significant losses from highly polyphagous stink bug species. These problems may be exacerbated by crop rotation practices that often result in cotton, peanut, Arachis hypogaea L., and soybean, Glycine max (L.) Merrill (both Fabaceae), growing in close proximity to one another. Because all of these crops are hosts for the major pest stink bug species in the region, we experimentally examined colonization preference of these species among the crops to clarify this aspect of their population dynamics. We planted peanut, soybean, Bt cotton, and glyphosate‐tolerant (RR) non‐Bt cotton at three sites over 3 years in replicated plots ranging from 192 to 1 323 m2 and calculated odds ratios for colonization of each crop for Nezara viridula (L.) and Euschistus servus (Say) (both Hemiptera: Pentatomidae). In four of five experiments, both E. servus and N. viridula preferred soybean significantly more often than Bt cotton, non‐Bt cotton, and peanut. Neither N. viridula nor E. servus showed any preference between non‐Bt and Bt cotton in any experiment. Both species had higher numbers in Bt and non‐Bt cotton relative to peanut; this was not significant for any single experiment, but analyses across all experiments indicated that N. viridula preferred Bt and non‐Bt cotton significantly more often than peanut. Our results suggest that soybean in the landscape may function as a sink for stink bug populations relative to nearby peanut and cotton when the soybean is in the reproductive stage of development. Stink bug preference for soybean may reduce pest pressure in near‐by crops, but population increases in soybean could lead to this crop functioning as a source for later‐season pest pressure in cotton.

Competition between stink bug and heliothine caterpillar pests on cotton at within‐plant spatial scales

Outbreaks of non‐target pests associated with transgenic Bt cotton threaten the economic and ecological benefits of the technology in cotton‐producing countries. In the southeastern USA, stink bug pests, namely Nezara viridula L. and Euschistus servus Say (both Heteroptera: Pentatomidae), have recently become severe problems associated with Bt cotton, requiring continued insecticide use. However, the causes of non‐target pest outbreaks remain unclear. Release from competition with Bt‐susceptible pest species, in addition to other mechanisms, may contribute to increased stink bug populations in Bt cotton. We investigated the competitive interactions between the two stink bug species and the Bt‐susceptible pests Helicoverpa zea (Boddie) and Heliothis virescens Fabricius (both Lepidoptera: Noctuidae) on non‐Bt cotton. We tested for competition effects on stink bug growth rates in no‐choice experiments at two spatial scales: a single cotton boll and a branch with multiple developing bolls. Although caterpillars of the two species had equivalent effects on resource availability, they had distinct effects on stink bug growth rates. Fourth instar H. zea reduced growth rates of both stink bug species by 60% when caged on a single cotton boll and reduced growth rates of only E. servus by 36% when caged on a cotton branch. In contrast, H. virescens had no effect on stink bug growth rates. Resource competition was apparent in the interactions between H. zea and E. servus, but interference competition may have contributed to the interactions as well. Competitive release of stink bug populations in Bt cotton is possible, and should be more likely for E. servus than for N. viridula. Understanding the causes of non‐target pest outbreaks in Bt cotton will contribute to improved environmental risk assessments of future releases of Bt cotton and related transgenic crops.

Influence of handling and conditioning protocol on learning and memory of Microplitis croceipes

Microplitis croceipes (Cresson) (Hymenoptera: Braconidae) learns odors in association with both hosts and food. The food‐associated ‘seeking’ behavior of M. croceipes was investigated under various training protocols utilizing the conditioning odor, 3‐octanone. We investigated the effects of odor training, or its lack, training duration, training frequency, time elapsed after training, wasp hunger state, and training reinforcement, on the food‐seeking responses of M. croceipes females. We found that odor‐trained females show strong food seeking responses, whereas non‐odor‐trained females do not respond to the odor, and that a single 10 s association with the odor whilst feeding on sugar water subsequently conditioned the wasps to exhibiting significant responses to it. Increases in training time to more than 10 s did not improve their responses. Repetition of the food–odor associations increased a wasps recall, as well as its response over time, compared to a single exposure. Repeated exposure to the learned odor in the absence of a food reward decreased the responses of less hungry individuals. However, the level of response increased significantly following a single reinforcement with the food–odor association. Understanding the factors that influence learning in parasitoids can enhance our ability to predict their foraging behavior, and opens up avenues for the development of effective biological detectors.