Hotter droughts alter resource allocation to chemical defenses in piñon pine.

Heat and drought affect plant chemical defenses and thereby plant susceptibility to pests and pathogens. Monoterpenes are of particular importance for conifers as they play critical roles in defense against bark beetles. To date, work seeking to understand the impacts of heat and drought on monoterpenes has primarily focused on young potted seedlings, leaving it unclear how older age classes that are more vulnerable to bark beetles might respond to stress. Furthermore, we lack a clear picture of what carbon resources might be prioritized to support monoterpene synthesis under drought stress. To address this, we measured needle and woody tissue monoterpene concentrations and physiological variables simultaneously from mature piñon pines (Pinus edulis) from a unique temperature and drought manipulation field experiment. While heat had no effect on total monoterpene concentrations, trees under combined heat and drought stress exhibited ~ 85% and 35% increases in needle and woody tissue, respectively, over multiple years. Plant physiological variables like maximum photosynthesis each explained less than 10% of the variation in total monoterpenes for both tissue types while starch and glucose + fructose measured 1-month prior explained ~ 45% and 60% of the variation in woody tissue total monoterpene concentrations. Although total monoterpenes increased under combined stress, some key monoterpenes with known roles in bark beetle ecology decreased. These shifts may make trees more favorable for bark beetle attack rather than well defended, which one might conclude if only considering total monoterpene concentrations. Our results point to cumulative and synergistic effects of heat and drought that may reprioritize carbon allocation of specific non-structural carbohydrates toward defense.

Impacts of Dryland Cropping Systems on Ground Beetle Communities (Coleoptera: Carabidae) in the Northern Great Plains.

Ground beetles are natural predators of insect pests and small seeds in agroecosystems. In semiarid cropping systems of the Northern Great Plains, there is a lack of knowledge to how ground beetles are affected by diversified cover crop rotations. In a 2-yr study (2018 and 2019), our experiment was a restricted-randomization strip-plot design, comprising summer fallow, an early-season cover crop mixture (five species), and a mid-season cover crop mixture (seven species), with three cover crop termination methods (i.e., herbicide, grazing, and haying). Using pitfall traps, we sampled ground beetles in five 48-h intervals throughout the growing season (n = 135 per year) using growing degree day (GDD) accumulations to better understand changes to ground beetle communities. Data analysis included the use of linear mixed-effects models, perMANOVA, and non-metric multidimensional scaling ordinations. We did not observe differences among cover crop termination methods; however, activity density in the early-season cover crop mixture decreased and in summer fallow increased throughout the growing season, whereas the mid-season cover crop mixture peaked in the middle of the summer. Ground beetle richness and evenness showed a nonlinear tendency, peaking in the middle of the growing season, with marginal differences between cover crops or fallow after the termination events. Also, differences in ground beetle composition were greatest in the early- and mid-season cover crop mixtures earlier in the growing season. Our study supports the use of cover crop mixtures to enhance ground beetle communities, with potential implications for pest management in dryland cropping systems.

Establishing Diorhabda carinulata: Impact of Release Disturbances on Pheromone Emission and Influence of Pheromone Lures on Establishment.

Before weed biocontrol insects are transported and released in a new area, they are commonly collected into small paper containers, chilled, and kept under dark conditions. This process can be termed a pre-release protocol. The influence of a pre-release protocol on establishment success of a gregarious biological control agent was assessed using the northern tamarisk beetle, Diorhabda carinulata (Desbrochers), and its exotic, invasive host plant saltcedar (Tamarix spp.). Pre-release protocol impacts on aggregation pheromone production by D. carinulata were characterized under controlled conditions. Additional experiments were undertaken to determine if deployment of aggregation pheromone lures might enhance the agent's persistence at release sites. Adults that experienced the pre-release protocol produced less aggregation pheromone compared to undisturbed adults. Olfactometer bioassays indicated that a cohort of adults subjected to the pre-release protocol were less attractive to other adults than a control cohort. Efficacy of aggregation pheromone-based lures to retain adults at release sites was evaluated by comparing capture numbers of adult beetles at paired treatment and control release sites, 10-14 days after the release of 300, 500, or 1000 individuals. A greater number of adult D. carinulata were captured where the pheromone lures had been deployed compared to control release sites. Application of aggregation pheromone when a new release of D. carinulata is planned should allow biological control practitioners to increase retention of beetles at a release site.

Antixenosis, Antibiosis, and Potential Yield Compensatory Response in Barley Cultivars Exposed to Wheat Stem Sawfly (Hymenoptera: Cephidae) Under Field Conditions.

Wheat stem sawfly, Cephus cinctus Norton, is an economically serious pest of cereals grown in North America. Barley cultivars were previously planted as resistant crops in rotations to manage C. cinctus, but due to increasing levels of injury to this crop, this is no longer a valid management tactic in Montana. Therefore, we aimed to understand antixenosis (behavioral preference), antibiosis (mortality), and potential yield compensation (increased productivity in response to stem injuries) in barley exposed to C. cinctus. We examined these traits in eight barley cultivars. Antixenosis was assessed by counting number of eggs per stem and antibiosis was assessed by counting infested stems, dead larvae, and stems cut by mature larvae. Potential yield compensation was evaluated by comparing grain yield from three categories of stem infestation: 1) uninfested, 2) infested with dead larva, and 3) infested cut by mature larva at crop maturity. We found the greatest number of eggs per infested stem (1.80 ± 0.04), the highest proportion of infested stems (0.63 ± 0.01), and the highest proportion of cut stems (0.33 ± 0.01) in 'Hockett'. Seven out of eight cultivars had greater grain weight for infested stems than for uninfested stems. These cultivars may have compensatory responses to larval feeding injury. Overall, these barley cultivars contain varying levels of antixenosis, antibiosis, and differing levels of yield compensation. Our results provide foundational knowledge on barley traits that will provide a framework to further develop C. cinctus resistant or tolerant barley cultivars.

Analysis of recombinant inbred line populations derived from wheat landraces to identify new genes for wheat stem sawfly resistance.

Wheat landrace accessions were chosen from areas of the world with historical European wheat stem sawfly (Cephus pygmaeus L.) selection pressure to develop six recombinant inbred line (RIL) populations. Molecular maps were constructed, and resistance due to antibiosis and antixenosis was assessed at sites in Montana naturally infested by Cephus cinctus Norton, the wheat stem sawfly (WSS). Novel QTLs were identified along with QTL previously identified in elite germplasm. A newly identified QTL on chromosome 1B provided a new source for pith-filled solid stems. An allele for resistance on chromosome 4A unrelated to solid stems was identified in four of the six RIL populations. A landrace from Turkey, PI 166471, contained alleles at three QTLs causing high levels of larval mortality. None of the QTLs were related to stem solidness, but their combined effect provided resistance similar to that observed in a solid-stemmed check cultivar. These results show the utility of genetic populations derived from geographically targeted landrace accessions to identify new alleles for insect resistance. New PCR-based molecular markers were developed for introgression of novel alleles for WSS resistance into elite lines. Comparison of results with previous analysis of elite cultivars addresses changes in allele frequencies during the wheat breeding process.

A large-scale multiomics analysis of wheat stem solidness and the wheat stem sawfly feeding response, and syntenic associations in barley, Brachypodium, and rice.

The wheat stem sawfly (WSS), Cephus cinctus Norton (Hymenoptera: Cephidae), is an important pest of wheat and other cereals, threatening the quality and quantity of grain production. WSS larvae feed and develop inside the stem where they are protected from the external environment; therefore, pest management strategies primarily rely on host plant resistance. A major locus on the long arm of wheat chromosome 3B underlies most of the variation in stem solidness; however, the impact of stem solidness on WSS feeding has not been completely characterized. Here, we used a multiomics approach to examine the response to WSS in both solid- and semi-solid-stemmed wheat varieties. The combined transcriptomic, proteomic, and metabolomic data revealed that two important molecular pathways, phenylpropanoid and phosphate pentose, are involved in plant defense against WSS. We also detected a general downregulation of several key defense transcripts, including those encoding secondary metabolites such as DIMBOA, tricetin, and lignin, which suggested that the WSS larva might interfere with plant defense. We comparatively analyzed the stem solidness genomic region known to be associated with WSS tolerance in wild emmer, durum, and bread wheats, and described syntenic regions in the close relatives barley, Brachypodium, and rice. Additionally, microRNAs identified from the same genomic region revealed potential regulatory pathways associated with the WSS response. We propose a model outlining the molecular responses of the WSS-wheat interactions. These findings provide insight into the link between stem solidness and WSS feeding at the molecular level.

Correction to: A large-scale multiomics analysis of wheat stem solidness and the wheat stem sawfly feeding response, and syntenic associations in barley, Brachypodium, and rice.

The original version of this article contained a mistake. The word "RefSeq v.1" was incorrectly inserted on page 7. The correct sentence should be: To identify the differentially regulated transcripts, clean RNA-Seq reads were mapped onto the T. aestivum Chinese Spring chromosome 3B pseudomolecule.

Host plant quantitative trait loci affect specific behavioral sequences in oviposition by a stem-mining insect.

KEY MESSAGE: Genetic diversity in quantitative loci associated with plant traits used by insects as cues for host selection can influence oviposition behavior and maternal choice. Host plant selection for oviposition is an important determinant of progeny performance and survival for phytophagous insects. Specific cues from the plant influence insect oviposition behavior; but, to date, no set of host plant quantitative trait loci (QTLs) have been shown to have an effect on behavioral sequences leading to oviposition. Three QTLs in wheat (Triticum aestivum L.) have been identified as influencing resistance to the wheat stem sawfly (WSS) (Cephus cinctus Norton). Wheat near-isogenic lines (NILs) for each of the three QTLs were used to test whether foraging WSS were able to discriminate variation in plant cues resulting from allelic changes. A QTL on chromosome 3B (Qss-msub-3BL) previously associated with stem solidness and larval antibiosis was shown to affect WSS oviposition behavior, host preference, and field infestation. Decreased preference for oviposition was also related to a QTL allele on chromosome 2D (Qwss.msub-2D). A QTL on chromosome 4A (Qwss.msub-4A.1) affected host plant attractiveness to foraging females, but did not change oviposition preference after females landed on the stem. These findings show that oviposition decisions regarding potential plant hosts require WSS females to discriminate signals from the plant associated with allelic variation at host plant quantitative loci. Allele types in a host plant QTL associated with differential survival of immature progeny can affect maternal choices for oviposition. The multidisciplinary approach used here may lead to the identification of plant genes with important community consequences, and may complement the use of antibiosis due to solid stems to control the wheat stem sawfly in agroecosystems.

Untargeted metabolomics profiling of oat (Avena sativa L.) and wheat (Triticum aestivum L.) infested with wheat stem sawfly (Cephus cinctus Norton) reveals differences associated with plant defense and insect nutrition.

Introduction: Wheat stem sawfly (WSS), Cephus cinctus Norton, is a major pest of common bread wheat (Triticum aestivum L.) and other cultivated cereals in North America. Planting of cultivars with solid stems has been the primary management strategy to prevent yield loss due to WSS infestation, however expression of this phenotype can vary depending on environmental conditions and solid stems hinder biological control of WSS via braconid parasitoids Bracon cephi (Gahan) and Bracon lissogaster Muesebeck. In the hollow stems of oat (Avena sativa L.), WSS larvae experience 100% mortality before they reach late instars, but the mechanisms for this observed resistance have not been characterized. Objective: The objective of this study was to explore additional sources of resistance outside of the historic solid stem phenotype. Methods: Here, we use an untargeted metabolomics approach to examine the response of the metabolome of two cultivars of oat and four cultivars of spring wheat to infestation by WSS. Using liquid chromatography-mass spectrometry (LC-MS), differentially expressed metabolites were identified between oat and wheat which were associated with the phenylpropanoid pathway, phospholipid biosynthesis and signaling, the salicylic acid signaling pathway, indole-3-acetic acid (IAA) degradation, and biosynthesis of 1,4-benzoxazin-3-ones (Bxs). Several phospho- and galacto- lipids were found in higher abundance in oat, and with the exception of early stem solidness cultivar Conan, both species experienced a decrease in abundance once infested. In all wheat cultivars except Conan, an increase in abundance was observed for Bxs HMDBOA-glc and DIBOA-ß-D-glucoside after infestation, indicating that this pathway is involved in wheat response to infestation in both solid and hollow stemmed cultivars. Differences between species in compounds involved in IAA biosynthesis, degradation and inactivation suggest that wheat may respond to infestation by inactivating IAA or altering the IAA pool in stem tissue. Conclusion: We propose that the species differences found here likely affect the survival of WSS larvae and may also be associated with differences in stem architecture at the molecular level. Our findings suggest pathways to focus on for future studies in elucidating plant response to WSS infestation.

Cowpea extrafloral nectar has potential to provide ecosystem services lost in agricultural intensification and support native parasitoids that suppress the wheat stem sawfly.

The native parasitoids Bracon cephi (Gahan) and B. lissogaster Muesebeck (Hymenoptera: Braconidae) reduce populations of Cephus cinctus Norton (Hymenoptera: Cephidae), a native grassland species, and major wheat pest on the Northern Great Plains of North America. Non-host feeding adults of these braconids increase longevity, egg load, and egg volume when provisioned carbohydrate-rich diets. Nutrition from nectar can enhance the success of natural enemies in pest management programs. Cowpea, Vigna unguiculata (L.) Walpers, is a potential cover-crop that could add resilient features to the landscape and has extrafloral nectaries (EFN), easy-access nectar sources for beneficial insects. If more cowpea was grown on the Northern Great Plains, would B. cephi and B. lissogaster benefit from foraging on putatively beneficial EFN? We investigated cowpea inflorescence stalk extrafloral nectars (IS-EFN) and leaf stipel extrafloral nectars (LS-EFN) as potential food sources for these parasitoids. Females were caged on EFN sources on living cowpea plants to assess longevity. Egg load and volume were measured at 2, 5, and 10 days after placement. Bracon cephi survived 10 days on water, 38 days on IS-EFN; B. lissogaster 6 days on water, 28 days on IS-EFN. Bracon lissogaster maintained a constant egg load and volume across treatments while B. cephi produced 2.1-fold more eggs that were 1.6-fold larger on IS-EFN. Y-tube olfactometry indicated adult females were attracted to airstreams containing cowpea volatiles. These results demonstrate that non-native, warm-season cowpea benefits these native parasitoids and may improve conservation biocontrol of C. cinctus.

Using Stable Isotopes to Determine Natal Origin and Feeding Habits of the Army Cutworm Moth, Euxoa auxiliaris (Lepidoptera: Noctuidae).

The army cutworm, Euxoa auxiliaris (Grote), is a migratory noctuid that is both an agricultural pest and an important late-season food source for grizzly bears, Ursus arctos horribilis (Linnaeus, Carnivora: Ursidae), within the Greater Yellowstone Ecosystem. Beyond the confirmation of the moths' seasonal, elevational migration in the mid-1900s, little else has been documented about their migratory patterns. To address this missing ecological component, we examined (1) migratory routes during their spring and fall migratory periods throughout their natal range, the Great Plains, and (2) natal origin at two of their summering ranges using stable hydrogen (δ2H) analyses of wings from samples collected within the areas of interest. Stable carbon (δ13C) and stable nitrogen (δ15N) analyses of wings were used to evaluate larval feeding habits of the migrants and agricultural intensity of natal origin sites, respectively. Results suggest that, rather than migrating exclusively east to west, army cutworm moths are also migrating north to south during their spring migration. Moths did not exhibit natal origin site fidelity when returning to the Great Plains. Migrants collected from the Absaroka Range had the highest probability of natal origin in Alberta, British Columbia, Saskatchewan, the most southern region of the Northwest Territories, and second highest probability of origin in Montana, Wyoming, and Idaho. Migrants collected in the Lewis Range had the highest probability of origin in the same provinces of Canada. Results suggest that migrants of the Absaroka Range fed exclusively on C3 plants as larvae and rarely fed in heavily fertilized agroecosystems.

Plant Volatiles and Oviposition Behavior in the Selection of Barley Cultivars by Wheat Stem Sawfly (Hymenoptera: Cephidae).

Wheat stem sawfly, [Cephus cinctus (Hymenoptera: Cephidae)], females display complex behaviors for host selection and oviposition. Susceptible hollow stem wheat (Triticum aestivum L.) cultivars release a greater amount of attractive compound, (Z)-3-hexenyl acetate and receive a greater number of eggs compared to resistant solid stem wheat cultivars. However, barley (Hordeum vulgare L.) is becoming a more common host for C. cinctus in Montana. Therefore, how do host selection and oviposition behaviors on barley cultivars compare to what happens when encountering wheat cultivars? To answer this question, we carried out greenhouse experiments using two barley cultivars: 'Hockett' and 'Craft'. Between these cultivars at Zadoks stages 34 and 49, we compared host selection decisions using a Y-tube olfactometer, compared oviposition behaviors on stems, and counted the number of eggs inside individual stems. In Y-tube bioassays, we found a greater number of C. cinctus females were attracted to the airstream passing over 'Hockett' than 'Craft' barley cultivars. Although the frequencies of oviposition behaviors were similar between these cultivars, the number of eggs was greater in 'Hockett'. Volatile profiles indicated that the amount of linalool was greater in the airstream from 'Craft' than in 'Hockett' at Zadoks 34 while the amount of (Z)-3-hexenyl acetate was greater in airstream from 'Hockett' at both Zadoks 34 and 49. These results suggest that volatiles of barley plants influenced host selection behavior of ovipositing C. cinctus females, while other discriminating behaviors do not differ between cultivars.

Photosynthesis and yield reductions from wheat stem sawfly (Hymenoptera: Cephidae): interactions with wheat solidness, water stress, and phosphorus deficiency.

The impact of herbivory on plants is variable and influenced by several factors. The current study examined causes of variation in the impact of larval stem mining by the wheat stem sawfly, Cephus cinctus Norton (Hymenoptera: Cephidae), on spring wheat, Triticum aestivum L. We performed greenhouse experiments over 2 yr to (1) study whether biotic (hollow versus solid stemmed host wheat) and abiotic (water, phosphorus stress) factors interact with C. cinctus stem mining to influence degree of mined stem physiological (photosynthesis) and yield (grain weight) reductions; and (2) determine whether whole plant yield compensatory responses occur to offset stem-mining reductions. Flag leaf photosynthetic reduction was not detected 16-20 d after infestation, but were detected at 40-42 d and doubled from water or phosphorus stresses. Main stem grain weight decreased from 10 to 25% from stem mining, largely due to reductions in grain size, with greater reductions under low phosphorus and/or water levels. Phosphorus-deficient plants without water stress were most susceptible to C. cinctus, more than doubling the grain weight reduction due to larval feeding relative to other water and phosphorus treatments. Two solid stemmed varieties with stem mining had less grain weight loss than a hollow stemmed variety, so greater internal mechanical resistance may reduce larval stem mining and plant yield reductions. Our results emphasize the importance of sufficient water and macronutrients for plants grown in regions impacted by C. cinctus. Also, solid stemmed varieties not only reduce wheat lodging from C. cinctus, they may reduce harvested grain losses from infested stems.

Multiple decrement life tables of Cephus cinctus Norton (Hymenoptera: Cephidae) across a set of barley cultivars: The importance of plant defense versus cannibalism.

Accurately estimating cause-specific mortality for immature insect herbivores is usually difficult. The insects are exposed to abiotic and biotic mortality factors, causing cadavers to simply disappear before cause of mortality can be recorded. Also, insect herbivores are often highly mobile on hosts, making it difficult to follow patterns for individuals through time. In contrast, the wheat stem sawfly, Cephus cinctus Norton, spends its entire egg, larval, and pupal period inside a host stem. Therefore, with periodic sampling stage-specific causes of mortality can be ascertained. Consequently, we examined C. cinctus mortality in eight barley, Hordeum vulgare L., cultivars in two locations in Montana from 2016 to 2018 by collecting stem samples from stem elongation to crop maturity at weekly intervals, and collecting overwintered barley stubs the following spring and summer from the same plots. If larvae were present, we examined larval status-dead or alive-and categorized dead individuals into one of 5 mortality categories: plant defense, cannibalism, parasitism, pathogens, and unknown factors. We used multiple decrement life tables to estimate cause-specific mortality and irreplaceable mortality (the proportion of mortality from a given cause that cannot be replaced by other causes of mortality). Plant defense (antibiosis) caused 85.7 ± 3.6%, cannibalism (governed by antixenosis) caused 70.1 ± 7.6%, parasitism caused 13.8 ± 5.9%, unknown factors caused 38.5 ± 7.6%, and pathogens caused 14.7 ± 8.5% mortality in the presence of all causes of mortality. Similarly, irreplaceable mortality due to plant defense was 22.3 ± 6.4%, cannibalism was 29.1± 4.2%, unknown factors was 6.2 ± 1.8%, pathogens was 0.9 ± 0.5%, and parasitism was 1. 5 ± 0. 6%. Antibiosis traits primarily killed newly emerged larvae, while other traits supported more favorable oviposition decisions by females, increasing mortality by obligate cannibalism. Our results suggest that breeding barley for resistance to C. cinctus targeting both categories of traits (antibiosis and antixenosis) is a highly valuable tactic for management of this important pest.

Effect of Precipitation and Temperature on Larval Survival of Cephus cinctus (Hymenoptera: Cephidae) in Barley Cultivars.

Host plant traits strongly affect survivorship of insect herbivores, and host suitability is especially important for the wheat stem sawfly, Cephus cinctus Norton, which spends its entire egg, larval, and pupal periods in a single stem. Measuring larval survival inside stems from egg hatch through diapause-mediated dormancy is a potential measure of population size for the next year but is also useful in assessing effects of growing season precipitation and temperature. Larval growth is synchronized with host plant growth, and the larva cannot switch hosts. Thus, incorporating plant physiological time, as growing degree days (GDD), may yield a better prediction of larval survival. Therefore, we assessed wheat stem sawfly survival from early larval growth to the beginning of autumnal diapause in barley cultivars selected from across feed, forage, and two- or six-row malt groups. Field experiments were conducted in Gallatin and Chouteau counties, Montana, in 2016 and 2017. We used Kaplan-Meier estimation to assess larval survival among cultivars. We found that the survival of pre-diapause larvae was greatest in 'Hockett' (36.5%) and lowest in 'Celebration' (15.4%). Precipitation and temperature during the growing season affected temporal patterns for larval survival across study sites. Adjusting survivorship curves using site-specific GDD accumulation allowed cultivar-specific survivorship to be estimated more precisely for each site, despite differing environmental influences. Our findings suggest that measuring wheat stem sawfly survival across barley cultivars and standardizing by site-specific GDDs may provide better recommendations on barley cultivars that impede wheat stem sawfly population growth and reduce economic losses.

An Herbivore-Induced Plant Volatile From Saltcedar (Tamarix spp.) Is Repellent to Diorhabda carinulata (Coleoptera: Chrysomelidae).

The leaf beetle Diorhabda carinulata Desbrochers (Coleoptera: Chrysomelidae) was introduced into the United States in 1999 for classical biological control of the exotic woody invader saltcedar (Tamarix spp. L. [Caryophyllales: Tamaricaceae]). The recent southern expansion of the range of D. carinulata in the United States has precipitated conflict between proponents of biological control of Tamarix and those with concerns over habitat conservation for avian species. Several semiochemicals that mediate aggregations by this species have been reported, but no repellent compounds have been recorded thus far. We now report a repellent compound, 4-oxo-(E)-2-hexenal, induced by adult D. carinulata feeding on saltcedar foliage. Collection of headspace volatiles, gas chromatography mass spectrometry, and electroantennographic analyses identified 4-oxo-(E)-2-hexenal as an insect-induced compound that is antennally active. Behavioral and exposure assays were conducted to test for repellency and toxicity in adults and larvae. Headspace volatiles were also collected from adult males exposed to 4-oxo-(E)-2-hexenal to determine the impact exposure might have on the emission of the aggregation pheromone. 4-Oxo-(E)-2-hexenal elicited electrophysiological responses in adults of both sexes. Behavioral responses indicated repellency across multiple doses for reproductive D. carinulata adults but not in nonreproductive adults. Exposure assays indicated altered behaviors in first instar larvae and adults, but not in third instar larvae. Collection of headspace volatiles indicated that exposure to 4-oxo-(E)-2-hexenal did not alter emission of the D. carinulata aggregation pheromone by adult males. The continued development and field deployment of this repellent compound may provide a new tool for the management of D. carinulata.

Mycoses of wheat stem sawfly (Hymenoptera: Cephidae) larvae by Fusarium spp. isolates.

A complex of Fusarium spp., including F. pseudograminearum, F. culmorum, F. avenaceum, F. equiseti, and F. acuminatum, was isolated from field-collected larval cadavers of wheat stem sawfly at two locations for 2 yr. The Fusarium spp. isolates caused mortality in both diapausing larvae in a topical bioassay and in developing larvae feeding in infected stems in a greenhouse experiment. Larval mortality was >90% in both experiments at the highest dose. The pattern of correlation between integument discoloration, hyphal growth, and larval mortality showed that the Fusarium spp. isolates actively infect larvae and kill them, rather than colonizing larval tissue as secondary postmortem invaders. The versatility of Fusarium spp. as plant and insect pathogens enables colonization that results in disease in wheat plants and subsequent mortality of the wheat stem sawfly larvae developing within the same tissue.

Impact of Diuraphis noxia and Rhopalosiphum padi (Hemiptera: Aphididae) on primary physiology of four near-isogenic wheat lines.

The impact of feeding injury by the Russian wheat aphid, Diuraphis noxia (Mordvilko) (Hemiptera: Aphididae), and bird cherry-oat aphid, Rhopalosiphum padi (L.) (Hemiptera: Aphididae) on susceptible and resistant wheat, Triticum aestivum L., near-isogenic lines 'Tugela' (susceptible), Tugela-Dn1 (antibiotic), Tugela-Dn2 (tolerant), and Tugela-Dn5 (antixenotic) was evaluated by assessing photosynthetic parameters. Photosynthesis and closely related parameters, pigment composition, and nonstructural carbohydrates were measured at 1, 3, and 9 d after aphids were introduced on plants maintained under greenhouse conditions. Overall, R. padi had a higher reproductive capacity within a period of 9 d compared with D. noxia on all lines except Tugela-Dn2. Although the visible injury symptoms associated with aphid injury can be highly species specific, the data indicate that photosynthetic reduction is a common physiological pattern of wheat response to aphid feeding, irrespective of chlorosis elicitation. Although both aphids negatively affected net photosynthesis, D. noxia had a greater impact than R. padi, even when aphid numbers were considerably fewer for D. noxia (100-150 aphids per plant) compared with R. padi (> 200 aphids per plant). The photosynthetic pigment and carbohydrate data suggest that the initial net photosynthesis reduction elicited by aphid feeding may not be directly related to the light reaction portion of the photosynthetic pathway via pigment losses. It is also unlikely that source-sink manipulation is the primary cause for the observed short-term inhibition of photosynthesis.

Cultivar preferences of ovipositing wheat stem sawflies as influenced by the amount of volatile attractant.

The wheat stem sawfly, Cephus cinctus Norton, causes severe losses in wheat grown in the northern Great Plains. Much of the affected area is planted in monoculture with wheat, Triticum aestivum L., grown in large fields alternating yearly between crop and no-till fallow. The crop and fallow fields are adjacent. This cropping landscape creates pronounced edge effects of sawfly infestations and may be amenable to trap cropping using existing agricultural practices. The behavioral preference for two wheat varieties was assessed in the context of developing trap crops for this insect. In field nurseries, stem lodging assessments indicated that the cultivar 'Conan' was infrequently damaged, whereas 'Reeder' was often heavily damaged. In laboratory choice and no-choice tests, 'Reeder' was significantly preferred by ovipositing wheat stem sawfly females. These two cultivars did not differ significantly in height or developmental stage, factors known to impact sawfly preference. Although Conan received fewer eggs than Reeder in no-choice tests, oviposition was further reduced in choice tests, indicating that females clearly preferred Reeder. In field trials where the overall dimensions of the spatial structure in choice tests was varied, females always selected Reeder over Conan in alternating block, row, and interseeded planting scenarios. Reeder releases greater amounts of the attractive compound, (Z)-3-hexenyl acetate than Conan but is similar to Conan for three other known, behaviorally active volatile compounds. The results are discussed in terms of cultivar selection for large scale trap crop experiments for the wheat stem sawfly.

Dryland Organic Farming Partially Offsets Negative Effects of Highly Simplified Agricultural Landscapes on Forbs, Bees, and Bee-Flower Networks.

Industrialized farming practices result in simplified agricultural landscapes, reduced biodiversity, and degraded species-interaction networks. Thus far, most research assessing the combined effects of farming systems and landscape complexity on beneficial insects has been conducted in relatively diversified and mesic systems and may not represent the large-scale, monoculture-based dryland agriculture that dominates many regions worldwide. Specifically, the effects of farming systems on forbs, bees, and their interactions are poorly understood in highly simplified dryland landscapes such as those in the Northern Great Plains, United States, an area globally important for conventional and organic small grain, pulse, forage, and oilseed production. During a 3-yr (2013-2015) study, we assessed 1) the effects of dryland no-till conventional and tilled organic farming on forbs, bees, and bee-flower networks and 2) the relationship between natural habitat and bee abundance. Flower density and richness were greater in tilled organic fields than in no-till conventional fields, and forb community composition differed between farming systems. We observed high bee diversity (109 taxa) in this highly simplified landscape, and bee abundance, richness, and community composition were similar between systems. Compared with tilled organic fields, bee-flower interactions in no-till conventional fields were poorly connected, suggesting these systems maintain relatively impoverished plant-pollinator networks. Natural habitat (11% of the landscape) did not affect small-bodied bee abundance in either farming system but positively affected large-bodied bees within 2,000 m of crop-field centers. In highly simplified agricultural landscapes, dryland organic farming and no-till conventional farming together support relatively high bee diversity, presumably because dryland organic farming enhances floral resources and bee-flower networks, and no-till management in conventional farming provides undisturbed ground-nesting habitats for wild bees (Hymenoptera: Apoidea).