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Sugar Feeding by Coccinellids Under Field Conditions BioControl (2011) 56:305–314 DOI 10.1007/s10526-010-9337-3 Sugar feeding by coccinellids under field conditions: the effects of sugar sprays in soybean Michael P. Seagraves • Yukie Kajita • Donald C. Weber • John J. Obrycki • Jonathan G. Lundgren Received: 26 July 2010 / Accepted: 18 November 2010 / Published online: 5 December 2010 Ó International Organization for Biological Control (outside the USA) 2010 Abstract Sucrose was applied weekly throughout economically threatening levels in SD. Coccinellids the growing season at three US locations (South digested sugar meals within 1 h of consumption Dakota [SD], Maryland [MD], and Kentucky [KY]), (measured using the cold anthrone test). Despite this and coccinellids and aphids (Aphis glycines Matsum- narrow window of detection, field-collected coccin- ura [Hemiptera: Aphididae]) were sampled 24 h later. ellids frequently tested positive for fructose. Under Total coccinellid densities were 50–77% greater in natural conditions, sugar is commonly ingested by sugar-sprayed soybean than in untreated plots. Coc- coccinellids and sugar sprays increase coccinellid cinella septempuncata L., Hippodamia convergens densities and their consumption of sugar. Sugar Gue´rin-Me´neville, and Harmonia axyridis Pallas sprays did not enhance biological control of aphids were more abundant where sugar was applied. in this experiment. Coleomegilla maculata (DeGeer) was found in equally low numbers in all treatments. Aphid densi- Keywords Anthrone Á Aphis glycines Á ties were similar in both treatments, and only reached Conservation biological control Á Food spray Á Omnivory Á Non-prey foods Handling Editor: Patrick De Clercq Introduction M. P. Seagraves Á J. G. Lundgren (&) USDA-ARS, North Central Agricultural Research Coccinellids feed on a wide range of sugar sources Laboratory, 2923 Medary Avenue, Brookings, under field conditions (Lundgren 2009a). In particu- SD 57006, USA lar, they are regular visitors to nectaries, both floral e-mail: [email protected] (Bugg 1987; Nalepa et al. 1992; Patt et al. 1997) and Present Address: extrafloral (Pemberton and Vandenberg 1993), and M. P. Seagraves honeydew coated plant surfaces. Honeydew functions Driscoll Strawberry Associates, Watsonville, as a contact kairomone for sternorrhynchan-feeding CA 95077, USA coccinellids, which promotes area-restricted search Y. Kajita Á J. J. Obrycki and oviposition (Carter and Dixon 1984; Evans and Department of Entomology, University of Kentucky, Dixon 1986; van den Meiracker et al. 1990), and it Lexington, KY 40546, USA also is a food for coccinellid larvae and adults (Putman 1955; Yinon 1969). D. C. Weber USDA-ARS, Invasive Insect Biocontrol and Behavior Sugar feeding improves the fitness and performance Laboratory, Beltsville, MD 20705, USA of coccinellids. The flight capacity of Coccinella 123 306 M. P. Seagraves et al. septempunctata L. is increased by sugar feeding in agroecosystems, and to determine if this feeding (Nedved et al. 2001). In the absence of prey, sugar behavior can be exploited to improve biological control consumption can increase the survival and longevity of of a serious crop pest. coccinellids (Matsuka et al. 1982;Dreyeretal.1997; Lundgren 2009a). Feeding on simple carbohydrates alone cannot support coccinellid reproduction, but diets Methods that contain both sugar and prey improve fecundity and other adult performance measurements compared to Field prey alone (Evans 2000; Lundgren 2009a). Specifically, sugar consumption reduces preoviposition period and Coordinated research was replicated at locations in increases survival, while decreasing egg resorption Kentucky (KY), Maryland (MD), and South Dakota during reproductive diapause (Smith and Krischik 1999; (SD). In SD, research was conducted at the Eastern Reznik and Vaghina 2006). South Dakota Soil and Water Research Farm near Supplementation of available sugar in cropping Brookings, SD (latitude, longitude: 44.348, -96.811). systems has long shown potential as a means of Soybean (var. Asgrow 1702) was planted on 15-May attracting, conserving and enhancing natural enemies 2009. In Kentucky at Spindle Top Farm (latitude, in cropland. Methods of sugar provisioning include longitude: 38.13, -84.52) soybean (Asgrow 4605) planting non-crop plants that produce nectar near was planted on 22-May 2009. In Maryland soybean crop fields, selection of nectaried crop varieties, and (var. Doeblers TS219OR) was planted at North Farm, applying food sprays (Lundgren 2009b). Techniques Beltsville Agricultural Research Center (latitude, lon- such as habitat manipulations that provide nectar- gitude: 39.034, -76.932), on 2-June 2009. At all bearing plants (Agnew et al. 1982; Bugg et al. 1991; locations row spacing was 76 cm and glyphosate was Patt et al. 1997), and application of food sprays applied for weed control. Plots (5 9 5 m each) were (Hagen et al. 1970; Wade et al. 2008; Lundgren arranged in a randomized complete block design with 2009b; Obrycki et al. 2009) increase coccinellid three replicates in KY and MD and four in SD. These densities. But no studies have attempted to quantify plots were embedded in larger soybean fields with at the degree to which these sugar-augmentation strat- least 50 m separating plots. egies affect the feeding behavior of coccinellids At all locations, weekly applications of 15% under natural conditions, or indeed whether coccin- sucrose solutions (790 l ha-1, total volume of 2 l) ellids consume sugar in the field at all. were evenly applied to the sugar-sprayed plots In this study we examined the numerical and feeding beginning on 22-June 2009 (SD and KY) and 6-July response of selected predaceous coccinellid species to 2009 (MD). Sugar was reapplied until soybean aphids weekly applications of sucrose in soybean at three exceeded the economic injury level of 690 aphids per locations across the United States. Soybean was selected plant (Ragsdale et al. 2007) in SD. Plots were as an experimental system because it is a major sampled on days without rain, after the morning dew component of the landscape in the United States, and had left the plants (i.e., in the afternoon). coccinellids are important predators of the invasive soybean aphid, Aphis glycines Matsumura (Hemiptera: Sampling and lady beetle collection Aphididae). Specific hypotheses that we tested are that (1) common agricultural coccinellids regularly consume All plots were sampled for insects 24 h after sugar sugar under natural conditions, and (2) early-season application using whole plant counts for soybean sugar sprays aggregate coccinellids to soybean fields, aphids and sweep samples for coccinellids. To max- and delay the onset of soybean aphid populations. To imize efficiency during aphid sampling, 20 randomly test these hypotheses, field sampling was paired with gut selected plants per plot were examined when 0–80% of content analysis using the cold anthrone test as applied plants were infested, ten plants when 81–99% of plants recently for other natural enemy groups (Heimpel et al. were infested, and five plants were sampled at 100% 2004; Lee et al. 2006; Taylor and Pfannenstiel 2008). aphid infestation. Fifty sweep samples (38 cm diam- Our intent was to provide information on how an eter sweep net) were collected in each plot and important predatorgroup responds toa commonnutrient coccinellids were counted and identified to species. 123 Sugar feeding by lady beetles 307 Aphids per plant and lady beetles per 50 sweeps absorbance of the subsample that received sulfuric acid were square-root transformed for each location, and only (i.e., background absorbance) was subtracted treatment means were compared using repeated- from the value of the subsample that received anthrone measures ANOVA (with treatment as the between- reagent. Lady beetles of each species (consolidated subject variable and sample date as the within- from at least 25 individuals) were starved for 24 h, and subjects variable). Additionally, seasonal sums of their guts were dissected and centrifuged as described total lady beetles captured per plot at each location, above to be used as negative controls (e.g., unfed and individual lady beetle species per plot at each controls). At least two wells of each negative control location, were compared between treatments and (of each lady beetle species) were included on each among study locations using two-factor ANOVA plate, and the standard deviation of this control series (with treatment and location as factors). was calculated for each beetle species on each plate. All adult lady beetles were placed in individual A sample was considered positive for fructose if the 1.5 ml centrifuge tubes and stored on dry ice as they difference in the optical density of the sample exceeded were transported to the laboratory. The samples were that of its no-anthrone control difference by a threshold then stored at –20°C until their guts were analyzed. In value (specifically, three times the standard deviation SD, each plot was visually examined for 4 min after of the starved beetle control series for each plate, e.g., sampling, and all lady beetles were hand-collected and Lee et al. 2006; Harwood et al. 2007). the guts of these beetles were also analyzed. Also in SD, The mean proportion of coccinellids testing coccinellids were collected from other habitats (e.g., positive for fructose was tested on arcsine-trans- wheat, field margins) and tested for sugar feeding. formed proportions using a two-factor ANOVA (location and treatment being the factors). This test Anthrone test for detection of sugar feeding was not conducted for KY as too few individuals were collected from the untreated plots to serve as a Gut content analyses of sugar contents were conducted comparison to those collected in the sugar-sprayed using methods of van Handel (1985). Thawed coc- soybean. cinellids were rinsed in water to remove any sugar contamination. The entire gut was then dissected in Sugar feeding laboratory assay Ringer’s solution (0.75 g NaCl2, 0.35 g KCl, 0.28 g CaCl2 in 1 l H2O) using heat-sterilized tools, and the Independent laboratory feeding assays were per- excised gut was placed directly into 50 llof2% formed to determine the relative rates of fructose sodium sulphate buffer.
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