Prey discrimination by a generalist coccinellid predator: effect of prey age or parasitism?

1. Patterns of prey discrimination by the generalist predatory coccinellid Coleomegilla maculata lengi Timb. (Coleoptera: Coccinellidae) were studied in relation to prey quality, under laboratory conditions. 2. Choice experiments were performed in which second‐ and fourth‐instar coccinellids had the choice between Trichoplusia ni (Lepidoptera: Noctuidae) eggs that were young or old, unparasitised or parasitised by Trichogramma evanescens Westwood (Hymenoptera: Trichogrammatidae). 3. Coleomegilla maculata larvae did not exhibit any preference for parasitised or unparasitised eggs at a similar stage of development but always preferred younger eggs, regardless of whether or not they were parasitised. Furthermore, the percentage of rejection was higher and handling time longer on the less preferred egg type. Pre‐imaginal development time, food intake to reach adulthood, and survival of coccinellid immatures were altered when coccinellid larvae were fed with parasitised and old unparasitised eggs. 4. These results indicate that C. maculata larvae select eggs based on their stage of development regardless of parasitism, and that prey quality of aged prey is lower. 5. Fourth‐instar coccinellid larvae spent less time in patches containing solely parasitised old eggs, and their level of exploitation was greatly reduced, compared with homogeneous patches containing unparasitised young eggs. This suggests that C. maculata larvae respond to variable patch quality by using flexible decision rules that reflect the payoff of the patch.     

Evidence for utilization of Diptera in the diet of field-collected coccinellid larvae from an antibody-based detection system

Aphidophagous coccinellid larvae have a wide range of potential prey in alfalfa and during times of low aphid abundance, larvae may supplement their diet with alternative prey. To understand the effects of the seasonal aphid availability on alternative prey use, an order-specific monoclonal antibody, DrosW-VI-B8, was used to examine the frequency of dipteran predation by these important natural enemies. Over 400 larvae were hand-collected from alfalfa and, in parallel, arthropod abundance was recorded. Harmonia axyridis and Coccinella septempunctata larvae were abundant early in the season when aphid populations were at their peak and Coleomegilla maculata larvae were collected later in the season when potato leafhoppers were abundant in the alfalfa. A relatively low proportion of field-collected H. axyridis, C. septempunctata, and C. maculata tested positive for dipteran proteins throughout the season. Similar to prior studies examining stage differences in coccinellid food breadth, older instars tested positive for dipteran proteins (3rd instar, 6% positive; 4th instar, 7% positive) but no early instars screened positive. This study provides a valuable insight into the trophic linkages that exist between coccinellid larvae and Diptera.     

Suitability of exotic and native lady beetle eggs (Coleoptera: Coccinellidae) for development of lady beetle larvae

Predation upon lady beetle (Coleoptera: Coccinellidae) eggs in the field is most often instances of egg cannibalism by larvae or adults while the majority of the remaining predation events upon coccinellid eggs is done by other species of Coccinellidae. Thus the recent introduction and establishment of Harmonia axyridis in the US could negatively affect native species of Coccinellidae via egg predation. However, little is known regarding the suitability of interspecific coccinellid eggs as a food source for larval development. In this study, it was found that native first or third instar Coleomegilla maculata and Olla v-nigrum larvae were incapable of surviving to the adult stage when provided solely exotic H. axyridis eggs. In stark contrast, H. axyridis larvae survived equally well when cannibalizing eggs or eating eggs of either native species. When C. maculata and O. v-nigrum were grouped as ‘native’ and compared with the exotic H. axyridis, more native eggs were attacked than exotic eggs and a higher percentage of eggs was attacked by H. axyridis larvae. Native and exotic larvae attacked a similar percentage of native eggs but native larvae attacked significantly fewer exotic eggs than did exotic larvae. These data suggest that H. axyridis may prey upon the eggs of these native species, when encountered in the field, compared with the likelihood of the native species preying upon H. axyridis eggs. Therefore, eggs of the native species C. maculata and O. v-nigrum will continue to be subjected to cannibalism and also to possible predation by other native species and the exotic H. axyridis.     

Relative toxicity of three wheat herbicides to two species of Coccinellidae

Abstract On the High Plains of the USA, herbicides specific for broad-leaf weeds are regularly applied to winter wheat in the early spring, sometimes late enough to coincide with the colonization of fields by cereal aphids and their natural enemies. We tested the toxicity of three such herbicides, Ally (Dupont), Rave (Syngenta) and 2,4-D ester (generic), to neonate larvae of two coccinellid species important in cereal aphid biocontrol, Coleomegilla maculata DeGeer and Hippodamia convergens Guérin-Méneville. Topical treatment of larvae with 2,4-D resulted in 25% and 60% mortality in the two species, respectively, with surviving C. maculata larvae experiencing a 5% increase in developmental time. No significant effects were noted for the other two materials, save for a 2.5% increase in developmental time for C. maculata larvae exposed to Rave. No material caused significant mortality in either species when larvae were fed on prey (Schizaphis graminum Rondani) treated with herbicide 24 h earlier, although 2,4-D reduced developmental time slightly in C. maculata. When herbicide applications are delayed enough in spring to coincide with aphid activity in wheat, farmers can reduce the risk of disrupting biological control by selecting an alternative to 2,4-D.     

Differential predation by Coleomegilla maculata on Colorado potato beetle strains that vary in growth on tomato

This study tests the hypothesis that the generalist predator Coleomegilla maculata DeGeer causes differential mortality of Colorado potato beetle, Leptinotarsa decemlineata (Say), larvae differing in their degree of genetic adaptation to tomato (Lycopersicon esculentum Mill.) as a host plant. Results of a series of laboratory experiments demonstrate that adult C. maculata can cause higher mortality to nonadapted than adapted Colorado potato beetle larvae. The extent of differential mortality caused by C. maculata depended on age of potato beetle larvae; presence of potato beetle eggs; whether or not the predator had a choice among prey items; and, in choice situations, the ratio of adapted to nonadapted potato beetle larvae. Although adult C. maculata have the potential to prey differentially on tomato-adapted and nonadapted Colorado potato beetle larvae in mixed populations, the magnitude of differential predation in a natural setting could be highly variable.     

Why eat extrafloral nectar? Understanding food selection by Coleomegilla maculata (Coleoptera: Coccinellidae)

Methods of increasing predator abundance within a habitat include the incorporation of non-prey food items, yet the influence of this on predation intensity toward herbivores remains unknown. In order to gain an understanding of nectar feeding in the predaceous beetle, Coleomegilla maculata (DeGeer), laboratory studies were conducted evaluating prey consumption in the presence of extrafloral nectaries. The physiology of beetles with access to prey only and a mixed diet were compared. To elucidate results of beetle physiology, Y-tube olfactometer studies were conducted and preferences between food types evaluated. Coleomegilla maculata females consumed 9 % fewer aphids when nectar was available. Lipid and glycogen content, as well as oocyte volume were not increased upon consumption of a mixed diet. Evaluation of predator behavior when offered both food resources together and separately demonstrated that extrafloral nectaries are attractive.     

Functional Response ofColeomegilla maculata(Coleoptera: Coccinellidae) to Colorado Potato Beetle Eggs (Coleoptera: Chrysomelidae)

A comparative study of the functional response ofColeomegilla maculataDeGeer (Coleoptera: Coccinellidae) fourth instars was conducted under laboratory, greenhouse, and field conditions. In the laboratory, individual larvae were placed in 9-cm petri dishes for 24 h, with 1, 3, 5, or 7 Colorado potato beetle (Leptinotarsa decemlineata[Say]) (Coleoptera: Chrysomelidae) egg masses. Each egg mass was standardized at 15 eggs. In the greenhouse and field,C. maculatalarvae were provided with an equivalent of 0.5 to 35L. decemlineataegg masses/m²of potato leaf. Fourth instars ofC. maculataexhibited a type II functional response toL. decemlineataeggs under laboratory, greenhouse, and field conditions. Predator search efficiency was inversely related with prey density. The maximum mean attack rate (8.7 eggs) byC. maculatalarvae in the field was about half the mean attack rate in the laboratory (17.6 eggs) and greenhouse (20.1 eggs). The difference in prey density between the laboratory and field seems to have been a major contributing factor in determining the rate of predation, whereas differences in environmental conditions (e.g., temperature and possible alternate food) may explain the differences observed in the predation rate in the greenhouse and field.     

Is Tetranychus urticae suitable prey for development and reproduction of nafve Coleomegilla maculata？

The lady beetle Coleomegilla maculata De Geer is an omnivorous predator that could help suppress aphid and spider mite populations on plants in greenhouses, plantscapes or interiorscapes. We are assessing the nutritional requirements and feeding behavior of C. maculata on target prey （spider mites） and factitious （unnatural） food. Our ultimate goal is to develop an efficacious diet to mass produce C. maculata. In this study, we tested the hypothesis that Tetranychus urticae Koch （two-spotted spider mite） is not suitable prey for development and reproduction of naive C. maculata （i.e., with no prior exposure to T. urticae）. Our objectives were to （i） provide baseline data on the effects of consuming T. urticae on C. maculata life history, （ii） to compare the effects of consuming all stages of T. urticae versus eggs ofMusca domestica L. （common housefly）, and （iii） to determine if the consumption of plant products was beneficial. We used C. maculata from a colony reared only on Ephestia kuehniella Zeller （Mediterranean flour moth） eggs. In experiments, C. maculata larvae were reared from the first instar to adult stage with prey/food in replicated arenas; adult females were paired with a single male with prey/food. The results showed that naive C. maculata readily attacked and consumed T. urticae. Nevertheless, T. urticae was less suitable than M. domestica eggs for C. maculata development and reproduction. Applying a synthetic pollen-Chlorella alga powder （SPCA） in arenas containing T. urticae appeared to boost C. maculata female development and reproduction.     

Instar-specific defense ofColeomegilla maculata lengi (Col.: Coccinellidae): Influence on attack success of the intraguild predatorChrysoperla rufilabris (Neur.: Chrysopidae)

We conducted a laboratory study of the behavioural interactions between larval stages of the lady beetleColeomegilla maculata lengi Timberlake and an intraguild predator, the lacewingChrysoperla rufilabris Burmeister. Ladybird susceptibility to predation by third instarC. rufilabris is age-specific and influenced by larval defensive strategies. First and second instars ofC. maculata were captured more consistently by chrysopid larvae than third and fourth instars. Young coccinellids generally exhibit escape reactions (dropping, fleeing, retreating) but, do not survive after being caught by lacewings. In addition to escape reactions, older larvae may actively defend themselves by wriggling or biting, even after being pierced. The proportion of physical interactions that resulted in coccinellid death was significantly lower for older instars (<2%) than for first (35%) and second (19%) instars. Capture efficiency byC. rufilabris was related to the coccinellid-chrysopid size ratio. Dropping off the leaves, the most common defensive behaviour displayed by coccinellid larvae, contributed to reduce overall mortality of first instars to a level similar to the one observed for older stages. However, lacewings were able to prevent coccinellid dropping by impaling and holding their prey in the air, or by falling with the ladybirds to the ground where they continued their attack.     

Competitive interactions between an exotic and a native ladybeetle: a field cage study

We conducted field‐cage studies on the direct interactions between a coccinellid species native to North America, Coleomegilla maculata De Geer, and a species introduced from Asia, Harmonia axyridis (Pallas) (Coleoptera: Coccinellidae). We compared the mortality and weight gain of larvae of both species in field cages that enclosed one or both species with corn plants containing high or low aphid numbers. We did not find a significant effect of the presence of H. axyridis on the survival or weight gain of C. maculata, but H. axyridis larvae weighed more when kept with C. maculata for 5 days than when kept with equal numbers of conspecifics. This suggests that intraspecific competition was stronger for H. axyridis than the interspecific competition with C. maculata. The spatial distribution of C. maculata over the plants differed between single‐species and two‐species treatments in a manner that suggested that this species avoided interactions with H. axyridis.     

The effects of diet on herbivory by a predaceous lady beetle

We hypothesize that herbivory by predators is affected by the complexity of their diet. Coleomegilla maculata larvae fed prey – Acyrthosiphon pisum or Ostrinia nubilalis eggs – consumed two to three times more plant tissue (pinto beans) than those fed a mixed diet (containing pollen substitute and prey-based components). Simple, prey-only diets appear to lack plant-based nutrients important to this predator.     

Soybean Aphid Predators in Québec and the Suitability of Aphis glycines as Prey for Three Coccinellidae

Since its invasion of North America in 2000, the soybean aphid, Aphis glycines Matsumura (Homoptera: Aphididae) has notably changed the arthropod community of the soybean agroecosystem. The objectives of this study were to characterize the foliar predatory fauna associated with A. glycines in commercial soybean fields in Québec, Canada, and to evaluate the suitability of A. glycines as prey for three coccinellid species: Harmonia axyridis Pallas, Propylea quatuordecimpunctata L. and Coleomegilla maculata lengi Timberlake (Coleoptera: Coccinellidae). Field surveys showed that several predators responded rapidly to expanding and increasing populations of the soybean aphid. Coccinellidae were the most important aphidophagous predators observed in 2002 (58.6%) and 2003 (44.8%), with mainly four native and naturalized species co-occurring with the soybean aphid throughout the growing season. Measurement of fitness parameters under laboratory conditions (survival, development time, longevity, fecundity) indicated that A. glycines is an excellent prey for the development and reproduction of all three of the coccinellid species studied. The intrinsic rate of natural increase (r_m) was highest for H. axyridis (0.238 d⁻¹), intermediate for P. quatuordecimpunctata (0.215 d⁻¹) and lowest for C. maculata (0.134 d⁻¹).     

Who is the puppet master? Replication of a parasitic wasp-associated virus correlates with host behaviour manipulation

Many parasites modify their host behaviour to improve their own transmission and survival, but the proximate mechanisms remain poorly understood. An original model consists of the parasitoid Dinocampus coccinellae and its coccinellid host, Coleomegilla maculata; during the behaviour manipulation, the parasitoid is not in contact with its host anymore. We report herein the discovery and characterization of a new RNA virus of the parasitoid (D. coccinellae paralysis virus, DcPV). Using a combination of RT-qPCR and transmission electron microscopy, we demonstrate that DcPV is stored in the oviduct of parasitoid females, replicates in parasitoid larvae and is transmitted to the host during larval development. Next, DcPV replication in the host''s nervous tissue induces a severe neuropathy and antiviral immune response that correlate with the paralytic symptoms characterizing the behaviour manipulation. Remarkably, virus clearance correlates with recovery of normal coccinellid behaviour. These results provide evidence that changes in ladybeetle behaviour most likely result from DcPV replication in the cerebral ganglia rather than by manipulation by the parasitoid. This offers stimulating prospects for research on parasitic manipulation by suggesting for the first time that behaviour manipulation could be symbiont-mediated.     

Interactions between an introduced and indigenous coccinellid species at different prey densities

Coccinella septempunctata L. (Coleoptera: Coccinellidae), a Palearctic coccinellid, has established and rapidly spread throughout the United States. This quantitative examination of larval interactions between C. septempunctata and Coleomegilla maculata (DeGeer) (Coleoptera: Coccinellidae), a Nearctic coccinellid, was conducted under controlled prey densities. Larvae of both coccinellid species are affected by a limited diet [one pea aphid per day Acyrthosiphon pisum (Harris) (Homoptera: Aphididae)] compared with an excess diet (>20 aphids per day). Larval survival decreased from 86 to 63% in C. maculata and from 84 to 33% in C. septempunctata; mean preimaginal developmental time increased from 20.6 to 26.7 days in C. maculata, and from 18.1 to 32.0 days in C. septempunctata. Additionally, on one aphid per day, mean adult weight was reduced from 12.39 to 9.79 mg in C. maculata, and from 39.57 to 14.44 mg in C. septempunctata. Interspecific interactions, favoring C. septempunctata over C. maculata at a␣low prey density (one aphid per day), take the form of␣reduced survival of C. maculata compared with C.␣septempunctata (14 versus 66%). Reduced survival of␣C. maculata may be the result of competition for aphids or intraguild predation by C. septempunctata on C.␣maculata. No interspecific interactions (measured as effects on larval survival, preimaginal developmental time, and adult weight) were observed between larvae of these two species at a high prey density (>20 aphids per predator per day). Received: 15 September 1997 / Accepted: 20 May 1998     

Changes in digestive rate of a predatory beetle over its larval stage: Implications for dietary breadth

Prey and non-prey foods differ substantially in their suitability for zoophytophagous omnivores, but the relative quality of these foods depends on the stage-specific digestive capabilities of the organism in question. Quantitative (or real-time) PCR was used to amplify food-specific DNA and measure consumption rates and digestion efficiencies of four foods - two prey (Aphis glycines and Leptinotarsa decemlineata eggs) and two non-prey (Zea mays pollen and the yeast Saccharomyces cerevisiae) species - over different larval stages of Coleomegilla maculata. The amount of Z. mays pollen consumed increased as larvae aged, but not proportionately with larval size, such that consumption rates decreased uniformly with insect age. While aging larvae fed A. glycines had a similar pattern in their diminishing consumption rates, they consumed similar amounts of A. glycines regardless of age, suggesting a negative feedback mechanism for consumption of this species of aphids. Older larvae digested three of the four foods significantly more efficiently than younger larvae, the exception being larvae fed A. glycines which was digested at a similar rate throughout the larval stage. There was a significant effect of time on food quantity detected for all four species of food. We conclude that C. maculata expands its physiological capacity for digesting prey and non-prey foods as they age in order to better accommodate the increased nutritional needs of the older larvae. This strategy has important implications for the life history strategies of zoophytophagous insects and how they function within foods webs.     

Importance of predation by Coleomegilla maculata larvae in the natural control of the corn earworm in sweet corn

The corn earworm, Helicoverpa zea (Boddie) (Lepidoptera: Noctuidae) is a serious pest of many crops in North America, particularly sweet corn. The larvae of Coleomegilla maculata (DeGeer) (Coleoptera: Coccinellidae) are the most frequently observed predators of H. zea eggs in sweet corn. Companion cropping strategies have been developed to augment densities of C. maculata in sweet corn plantings. In this study, the importance of predation by C. maculata larvae was evaluated in experiments when larval abundance was manipulated using hand removal and physical exclusion. In 2003 and 2004, sentinel H. zea eggs were exposed on corn for 48 h. Egg mortality was 13.8% greater in the treatment where C. maculata larvae were present. In 2004 and 2005 eggs were exposed on corn under the same hand removal and physical exclusion techniques and monitored at 3-h intervals for 24 h to directly observe predation on sentinel eggs. Coleomegilla maculata larvae were the most frequently observed predators of the eggs, accounting for 45.9% of 85 total observed predation events in the control. Whereas in the treatment where C. maculata larvae were reduced, egg survival was 26.13% greater and only 37 total predation events were observed. All pest eggs were eliminated from individual ears 22.1% more often when C. maculata larvae were present at natural densities indicating that predation by this predator stage lessens crop damage by H. zea. The results of this study suggest that C. maculata larvae are an irreplaceable source of natural mortality for H. zea eggs on sweet corn.     

Response of coccinellids to their aphid prey at different spatial scales

Predators that have an increasing numerical response for aggregation, attack and oviposition to increasing prey density are thought to be ideal for biological control. However density-dependent processes are infrequently detected and explanations include differences in the scales at which observations are made, behavioral differences among species, and habitat features. We examined the aggregation of four species of colonizing adult coccinellids to varying prey densities at two spatial scales in a maize system. Three of the species, Adalia bipunctata, Hippodamia tredecimpunctata, and Hippodamia convergens, responded to aphid abundance at the plant scale, and one species, Coleomegilla maculata, responded to the average aphid density at the plot (10×10 m) scale. In addition, H. convergens responded to individual plants with high aphid abundance in those plots with many plants of high aphid abundance. These results suggests that C. maculata (and possibly H. convergens) may be better able to colonize fields before aphid populations reach high levels, whereas A. bipunctata and H. tredecimpunctata may only be able to respond to high aphid abundance at the plant scale. This study suggests that spatial scale can affect predator–prey dynamics in a species-specific manner. However, the differences among coccinellid species in the community appear to be complementary, potentially contributing to greater aphid suppression.An erratum to this article can be found at     

Populations of Predaceous Natural Enemies Developing on Insect-Resistant and Susceptible Tomato in North Carolina

Naturally occurring populations of immature and adultGeocoris punctipes,adultColeomegilla maculataand immature coccinellids were monitored on field-grown tomato lines susceptible and resistant toManduca sextaandHelicoverpa zea. Helicoverpa zeaandHeliothis virescenseggs and small larvae that serve as prey for these predators also were monitored. MoreH. zeaandH. virescenseggs and small larvae were found on resistant than on susceptible plant lines. However, similar populations of largeH. zeaandH. virescenslarvae were found on resistant and susceptible plants. The number of adultGeocoris punctipes,adultColeomegilla maculataand immature coccinellids on resistant plants was always as high or higher than the number on susceptible plants. The data demonstrate no incompatibility of host-plant resistance with biological control provided by these predaceous insects, but indicate that the number ofG. punctipesand coccinellids required to provide effective biological control may develop too late in the season to be of practical value. Large populations of stilt bugs (Jalysus wickhami,Hemiptera: Berytidae) and spiders were observed to occur earlier in the growing season than eitherG. punctipesor coccinellids and may be a significant source of mortality forH. zeaeggs and small larvae.     

A Comprehensive Assessment of the Effects of Bt Cotton on Coleomegilla maculata Demonstrates No Detrimental Effects by Cry1Ac and Cry2Ab

The ladybird beetle, Coleomegilla maculata (DeGeer), is a common and abundant predator in many cropping systems. Its larvae and adults are predaceous, feeding on aphids, thrips, lepidopteran larvae and plant tissues, such as pollen. Therefore, this species is exposed to insecticidal proteins expressed in insect-resistant, genetically engineered cotton expressing Cry proteins derived from Bacillus thuringiensis (Bt). A tritrophic bioassay was conduced to evaluate the potential impact of Cry2Ab- and Cry1Ac-expressing cotton on fitness parameters of C. maculata using Bt-susceptible and -resistant larvae of Trichoplusia ni as prey. Coleomegilla maculata survival, development time, adult weight and fecundity were not different when they were fed with resistant T. ni larvae reared on either Bt or control cotton. To ensure that C. maculata were not sensitive to the tested Cry toxins independent from the plant background and to add certainty to the hazard assessment, C. maculata larvae were fed artificial diet incorporated with Cry2Ab, Cry1Ac or both at >10 times higher concentrations than in cotton tissue. Artificial diet containing E-64 was included as a positive control. No differences were detected in any life-table parameters between Cry protein-containing diet treatments and the control diet. In contrast, larvae of C. maculata fed the E-64 could not develop to the pupal stage and the 7-d larval weight was significantly negatively affected. In both feeding assays, the stability and bioactivity of Cry proteins in the food sources were confirmed by ELISA and sensitive-insect bioassays. Our results show that C. maculata is not affected by Bt cotton and is not sensitive to Cry2Ab and Cry1Ac at concentrations exceeding the levels in Bt cotton, thus demonstrating that Bt cotton will pose a negligible risk to C. maculata. More importantly, this study demonstrates a comprehensive system for assessing the risk of genetically modified plants on non-target organisms.     

Adult Coccinellid Activity and Predation on Aphids in Spring Cereals

Improved understanding of coccinellid activity and predation on aphids in the field could clarify their potential in aphid biological control. Our objective was to determine the influence of abiotic and biotic factors on activity and predation by adults of three coccinellid species (Hippodamia convergens Guerin-Meneville, H. tredecimpunctata tibialis (Say), and Coleomegilla maculata DeGeer) in spring cereal fields. The proportion of time coccinellids spent searching was correlated with air temperature, aphid density, and time of day. The relationship between searching and these variables differed among species. Mean walking speed ranged from 66.4 cm/min for C. maculata to 83.2 cm/min for H. tredecimpunctata and increased with temperature for all species. The frequency of short flights (<2 m) by beetles increased with temperature and decreased with increasing aphid density for all species, but the relationship of short flight frequency to these variables differed among species. The frequency of long flights (>2 m) was similar for all species and influenced by temperature and calendar date. The frequencies with which aphids were encountered and eaten were correlated with aphid density and temperature for H. convergens and H. tredecimpunctata. Despite over 250 h spent observing adult coccinellid behavior in the field, predation data were insufficient to develop a useful predation model.