Mutualism between ants and honeydew-producing hemipterans is a highly successful evolutionary innovation that attains the status of ecological keystone across many terrestrial ecosystems, involving a multitude of actors through direct or cascading effects. In these relationships, ants often protect their hemipteran partners against their arthropod natural enemies, sometimes interfering with the biological control of pest species. However, the dynamics of these interactions are highly variable based on the specific identity of all the actors involved, and baseline data remain scarce. We performed a field experiment exposing colonies of the walnut aphid Panaphis juglandis attended by five European ant species (Camponotus piceus, Ca. vagus, Crematogaster scutellaris, Dolichoderus quadripunctatus, Lasius emarginatus) to a native and an exotic lady beetle (Adalia bipunctata and Harmonia axyridis), documenting the behavioural interactions between these insects and the performance of ants in the protection of the aphids. Our results reveal a significant behavioural diversity among the ant species involved, with D. quadripunctatus and L. emarginatus being the most aggressive and having the best performance as aphid defenders, and Ca. piceus being least effective and often fleeing away. Cr. scutellaris displayed a rare rescue behaviour attempting to pull away the aphids that the lady beetles grabbed. On the other hand, behavioural responses to A. bipunctata and H. axyridis were similar. Further investigations are needed to understand the eco-ethological implications of these differences, while a better understanding of ant behavioural diversity may help refine biological control strategies.

Since its arrival in 2004, the non-native Harlequin ladybird Harmonia axyridis (Coleoptera: Coccinellidae) has rapidly spread throughout Britain, and it is now the most common coccinellid in England. There have since been concerns about the detrimental effects it may have on native coccinellids because there is a strong correlation between the arrival of H. axyridis and the decline in native species, including the two-spot ladybird, Adalia bipunctata. However, there have been few studies of the behavioural interactions between these two species, which occupy a high-niche overlap. This study investigated if the presence of H. axyridis impacts the feeding behaviour of A. bipunctata through direct competition for aphid prey. Foraging and interactive behaviour of A. bipunctata and H. axyridis were investigated within microcosms. Adalia bipuncata exhibited a similar consumption rate and time in the presence of H. axridis, yet H. axyridis consumed 3.5 times more prey items and were seven times faster compared to A. bipuncata. Observations showed that H. axyridis does not directly disrupt the feeding behaviour of A. bipunctata, but rather indirectly excludes the native species through being a superior competitor for prey items. Results indicate that the decline in native coccinellid species may be a consequence of H. axyridis competitive advantage, but that the concept of coexistence should not be dismissed.

Developmental rates are highly variable, both within and between genotypes and populations. But the rationale for two differential (slow and fast) developmental rates within same cohort under varying prey supply has yet not been explored. For this purpose, we investigated the effect of scarce and abundant prey supply on slow and fast development at 27°C in two aphidophagous ladybirds, Menochilus sexmaculatus (Fabricius) and Propylea dissecta (Mulsant) and its effect on their body mass and reproductive attributes. The ladybirds were provided with scarce and abundant supply of Aphis craccivora Koch under standardized abiotic conditions in the laboratory. A clear bimodal (two peaks, where the first peak represented the fast developing individuals and the second peak slow developing individuals) pattern of distribution for both prey supplies was obtained, which got skewed with change in prey supply. On abundant prey supply, more fast developing individuals (139 M. sexmaculatus and 123 P. dissecta) were found and less (46 M. sexmaculatus and 36 P. dissecta) on scarce prey supply. Slow developing individuals had female biased sex ratio, higher longevity and lower body mass. Fast developing females laid higher number of eggs with higher egg viability. Results of the study are indicative of occurrence and constancy of the slow and fast developing individuals in the egg batch.

The ladybird beetle, Eriopis connexa (Germar) (Coleoptera: Coccinellidae), is one of the commonest predators of aphids (Hemiptera: Aphididae) in the cotton agroecosystem and in many other row and fruit crops in Brazil, and has been introduced into other countries such as the USA for purposes of aphid control. In addition, the boll weevil, Anthonomus grandis Boheman (Coleoptera: Curculionidae) is the most serious cotton pest where it occurs, including Brazil. Controlling boll weevils and other pests such as cotton defoliators still tends to involve the intense application of insecticides to secure cotton production. The pyrethroid insecticide lambda-cyhalothrin (LCT) is commonly used, but this compound is not effective against aphids; hence, a desirable strategy would be to maintain E. connexa populations in cotton fields where LCT is applied. Using populations of E. connexa resistant (Res) and susceptible (Sus) to LCT, we compared behavioural responses on treated cotton plants and under confinement on partially and fully treated surfaces, and assessed the insects' survival on treated plants compared with that of the boll weevil. The E. connexa resistant population caged on treated plants with 15 and 75 g a.i. ha−1 exhibited ≫82% survival for both insecticide concentrations compared with ≪3% and ≪17% survival for susceptible E. connexa populations and boll weevils, respectively. The response of E. connexa Res and Sus populations when released, either on the soil or on the plant canopy, indicated avoidance towards treated plants, as measured by elapsed time to assess the plant. When compared with susceptible individuals, resistant ones took longer time to suffer insecticide knockdown, had a higher recovery rate after suffering knockdown, and spent more time in the plant canopy. Based on behavioural parameters evaluated in treated arenas, no ladybird beetles exhibited repellency. However, irritability was evident, with the susceptible population exhibiting greater irritability compared with the resistant population and a subgroup comprising resistant individuals that had recovered from knockdown. The outcomes for the E. connexa Res population indicate a promising strategy for its maintenance when using the insecticide LCT in integrated pest management schemes to control boll weevil or other non-target pest of ladybird beetles in cotton fields.

Functional response of all instars, adult males and females of the ladybird Coccinella septempunctata L. was studied against two species of aphids, namely Lipaphis erysimi (Kalt.) and Brevicoryne brassicae L., infesting mustard and cabbage crops, respectively. The predatory stages of C. septempunctata exhibited a type II response on both the prey species. The attack rate was highest (0.0192) with the lowest handling time (7.9 min) when the adult female consumed L. erysimi. There was a significant difference in the predicted values of consumption of L. erysimi by fourth instars, adult males and females compared with those of B. brassicae, indicating that the former prey is preferred over the latter one. The adult females prey upon significantly more numbers of aphids than the adult males.

Larvae of aphidophagous ladybirds leave a cue in their tracks that deters oviposition. The influence of relatedness on this behaviour is for the first time explored in this paper. Two-spot ladybird females (Adalia bipunctata L.) under different conditions (young and naive, young and experienced, and old and naive) were exposed either to (i) clean filter paper, (ii) filter paper contaminated with unrelated larval tracks or (iii) filter paper contaminated with related larval tracks from their offspring. Oviposition time was recorded during nine hours. Oviposition was inhibited by larval tracks, and was more so by related than unrelated ones. Experienced females showed the same behaviour but with lower intensity. With old females, response was not significant with any type of larval tracks. This is the first report of the effect of relatedness on a ladybird's response to larval tracks.

Activity of 12 beneficial invertebrate groups was assessed in 2005 and 2006 on a farm in northern England split into conventional and organic management halves, using pitfall and pan traps set in both crops and field boundaries. Management, crop and boundary structure influences on invertebrate activity were assessed, as was the relationship between crop and boundary type. Classification of crop and boundary assemblages produced three and two groups, respectively, in both years. Organic arable crops had well-defined assemblages in both years; and, while grass and grass/clover fields were separated from conventional arable fields in 2005, there was mixing in 2006. One boundary group, in both years, was dominated by conventional arable fields with tall herbaceous boundary vegetation. The other group had more organic arable and grassy fields with shorter boundary vegetation. Redundancy analyses showed that a number of groups (Cantharidae, Coccinellidae, Syrphidae, Ichneumonidae, Braconidae, Proctotrupoidea, Lycosidae) were more active in organic arable fields with more Staphylinidae in conventional arable crops and no obvious trend with Carabidae, Hemiptera, Neuroptera and Linyphiidae. Activity of some groups, especially Coccinellidae, Syrphidae and parasitic wasps, was strongly related to weed cover. Staphylinidae were most active in tall herbaceous boundaries by conventional arable crops with more of a number of groups (Cantharidae, Coccinellidae, parasitic wasps) in short herbaceous boundaries by organic arable crops. Organic management produced most differences in aerially-dispersed invertebrates, and management had a profound effect on activity in field boundaries. Possible management prescriptions to increase invertebrate activity include changing sowing times, weed cover manipulation and field boundary and margin management.

The age-specific mating incidence and mating behaviour of an aphidophagous ladybird beetle Coelophora saucia Mulsant were studied. Males started mating at the age of 4 days. Newly emerged females mated with older males albeit forcibly. The incidence of mating was 100% at the age of 10 days. Courtship involved five steps, viz. approach, watch, examine, mount and attempt, and was male dominated under laboratory conditions. Results indicate that physical cues probably play a role in the initiation of courtship, while chemical signals are probably involved in the continuation of courtship, culminating in intromission. The mating duration, latent period and number of bouts gradually decreased with increase in the number of matings; however, the interval between bouts slightly increased. Pseudo-mating with dead mates was observed in C. saucia. The oviposition period, fecundity and % egg viability were highest in females mated multiple times, while lowest in those that mated once.

The success or not of ladybirds as biological control agents is dependent on both their foraging behaviour and their individual survival rates. The former is a function of the habitats they utilise; the latter, a consequence of their reproductive strategy. Egg clustering was investigated in two ladybird species, Aphidecta obliterata, a conifer specialist, and Adalia bipunctata, an arboreal woodland generalist. The effect of oviposition substrate (filter papers vs. spruce needles) on clutch size and oviposition preference was also tested. Adalia bipunctata laid significantly more eggs than A. obliterata. The size of egg clusters laid by the two coccinellids varied between species and substrate types. Adalia bipunctata laid larger egg clusters than A. obliterata, with batches reaching a maximum size of 32 eggs on spruce and 41 eggs on paper, while those of A. obliterata contained a maximum of 5 eggs on spruce and 9 eggs on paper. Of the clusters laid by A. obliterata, 18.6% of those on paper and 21.4% of those on spruce contained only a single egg, whereas a minimum of two eggs per cluster were laid by A. bipunctata. Smaller clusters were laid on the spruce cuttings by both species when compared with those laid on the filter paper, but A. obliterata laid significantly more eggs on spruce than on the filter paper (77% vs. 23%), whilst A. bipunctata laid significantly more eggs on the filter paper (91%). It is suggested that coccinellid eggs are more likely to be washed off spruce needles than broad leaves and that, by laying smaller egg clusters on spruce, A. obliterata reduces this risk. Adalia bipunctata usually lays its eggs on the underside of broad leaved trees and thus does not face this risk and thus can lay larger egg clusters. No differences in cannibalism rates were found between the two species. These findings have implications for the use of ladybirds as biological control agents in spruce forests.

The time during which prey remains are detectable in the gut of a predator is an important consideration in the interpretation of molecular gut-content data, because predators with longer detectability times may appear on the basis of unweighted data to be disproportionately important agents of prey population suppression. The rate of decay in detectability, typically expressed as the half-life, depends on many variables; one that has not been explicitly examined is the manner in which the predator processes prey items. The influence of differences in feeding mode and digestive physiology on the half-life of DNA for a single prey species, the Colorado potato beetle Leptinotarsa decemlineata (Say), is examined in two predators that differ dramatically in these attributes: the pink ladybeetle, Coleomegilla maculata (DeGeer), which feeds by chewing and then ingesting the macerated material into the gut for digestion; and the spined soldier bug, Podisus maculiventris (Say), which physically and enzymatically processes the prey extra-orally before ingestion and further digestion in the gut. In order to standardize the amount of DNA consumed per predator, a single L. decemlineata egg was used as the prey item; all predators were third instars. The PCR assay yields estimated prey DNA half-lives, for animals maintained under field temperatures, of 7.0 h in C. maculata and 50.9 h in P. maculiventris. The difference in the prey DNA half-lives from these two predators underscores the need to determine detectabilities from assemblages of predators differing in feeding mode and digestive physiology, in order to weight positives properly, and hence determine the predators' relative impacts on prey population suppression.

Whilst most animals invest equally in males and females when they reproduce, a variety of vertically transmitted parasites has evolved the ability to distort the offspring sex ratios of their hosts. One such group of parasites are male-killing bacteria. Here we report the discovery of females of the ladybird Anisosticta novemdecimpunctata that produced highly female-biased offspring sex ratios associated with a 50% reduction in egg hatch rate. This trait was maternally transmitted with high efficiency, was antibiotic sensitive and was infectious following experimental haemolymph injection. We identified the cause as a male-killing Spiroplasma bacterium and phylogenetic analysis of rDNA revealed that it belongs to the Spiroplasma ixodetis clade in which other sex ratio distorters lie. We tested the potential for interspecific horizontal transfer by injection from an infected A. novemdecimpunctata line into uninfected individuals of the two-spot ladybird Adalia bipunctata. In this novel host, the bacterium was able to establish infection, transmit vertically and kill male embryos.

Adult pale morphs of an aphidophagous lady beetle, Propylea dissecta (Mulsant) (Coleoptera: Coccinellidae), were reared on a non-insect diet for periods ranging from 0 to 30 days after eclosion. The prey-deprived females, when provided with the cowpea aphid, Aphis craccivora Koch (Homoptera: Aphididae), laid fertile eggs, indicating that they had retained their reproductive potential. The oviposition period, post-oviposition period, fecundity, viability of eggs and longevity of females decreased significantly, whereas the pre-oviposition period and developmental period of progeny increased significantly with an increase in the duration of prey deprivation. There was no significant effect on the mean reproductive rate (16.14 ± 0.53 to 18.03 ± 0.67 eggs per day) at various prey deprivation levels, although the reproductive output and oviposition duration decreased. These findings indicate that lady beetle females can withstand the stress of prey non-availability, by switching to alternative food sources and reproducing when essential prey becomes available again.

The demographic statistics of the ladybird Exochomus flaviventris Mader (Coleoptera: Coccinellidae) were investigated under laboratory conditions on the cassava mealybug, Phenacoccus manihoti Matile-Ferrero (Homoptera: Pseudococcidae), reared on four host plants with different levels of antibiotic resistance to the mealybug: two cassava (Manihot esculenta Crantz, Euphorbiaceae) varieties, Incoza and Zanaga, Faux-Caoutchouc (FC) (hybrid of M. esculenta × M. glaziovii Muel. Arg.) and water weed (Talinum triangulare Jack, Portulacacae). Preimaginal mortality, mean oviposition time and total fecundity of female ladybirds were strongly affected by the host plant even though there was no linkage with antibiotic resistance. The net reproduction rates R0 of the predator were 1.5,1.7 and 2.4 times higher on Zanaga than on water weed, Incoza and FC respectively, and the generation time (T) was significantly longer on Zanaga than on the other three plants. The intrinsic rate of increase rm was not modified by the antibiotic resistance of the host plants. With the two cassava varieties, Incoza and Zanaga, it was found that antibiosis significantly affected the life history parameters of the ladybird without modifying its fitness, and provided better cassava mealybug control.

Inherited bacteria that parasitically distort the pattern of sex allocation of their host, biasing allocation towards female progeny, are found in many arthropods. One such manipulation is male-killing, where male progeny of infected females die during embryogenesis. We here provide evidence for a male-killing bacterium in the coccinellid beetle, Adonia variegata. We then address 3 questions. First, is this male-killing bacterium one that is found in other hosts, or does it represent a new transition to male-killing within the eubacteria? Using the sequence of the 16S rDNA of the bacterium, we found that the male-killing bacterium is a member of the Flavobacteria–Bacteroides group, most closely related to the male-killing bacterium in another ladybird beetle, Coleomegilla maculata. Secondly, is there any evidence that this bacterium affects female host physiology? In a paired test under nutritional stress, we found no evidence for a physiological benefit to infection, and weak evidence of a physiological cost, in terms of reduced fecundity. Thirdly, is there any evidence of host involvement in the transmission of the bacterium to the germ line? We found no evidence of host involvement. Rather, bacteria migrated to the ovariole independently of host cells. We conclude that the bacterium is a parasite, and discuss how 2 different species of ladybird come to be infected with 1 lineage of bacterium, and why case studies of male-killing bacteria have generally found little evidence of any symbiont contribution to host physiological functioning.

An annual legume cover crop was evaluated in pecan orchards to increase coccinellids that would reduce pecan aphids. Treatments were a ‘Dixie’ crimson clover and hairy vetch mixture and a grass sod. Coccinellids were abundant on the legume ground covers, but coccinellid density in the tree canopy was rarely affected by ground cover treatment. There were substantial differences between the coccinellid species collected from the legumes and from the trees. In Oklahoma, Olla v-nigrum (Mulsant) and Cycloneda munda (Say) were the main species in the trees, while Hippodamia convergens Guerin, Coccinella septempunctata L, and Coleomegilla maculata lengi Timberlake were dominant species in the legumes. In Georgia, Harmonia axyridis (Pallas) was the dominant species in the trees, and C. septempunctata in the legumes. Pecan aphids at two Oklahoma sites usually were not affected by cover crop treatment, but at the Georgia site, early-season aphids frequently were lower with a legume cover crop than with a grass sod. The accumulated effect of all aphid predators and parasitoids attracted to the legume aphids may have reduced the density of the pecan aphid.

We evaluated selected cool-season annual and perennial legumes as potential ground covers to supply nitrogen and to increase beneficial arthropod populations in a pecan orchard. Densities of aphids (Homoptera: Aphididae), lady beetles (Coleoptera: Coccinellidae), damsel bug (Hemiptera: Nabidae), green lacewings (Neuroptera: Chrysopidae), brown lacewings (Neuroptera: Hemerobiidae), hover flies (Diptera: Syrphidae), spined soldier bug and other stink bugs (Hemiptera: Pentatomidae), and spiders (Araneida) were monitored at 7–14 day intervals during the growing season for three years. Aboveground biomass production and nitrogen content of the legumes was measured for two years. Aphids peaked during early spring each year, with the highest density usually on ‘Dixie’ crimson clover and ‘Kenland’ red clover. Density of lady beetles was positively correlated with that of aphids, but spider densities were not. Other arthropods usually were not abundant. Nitrogen in the tops of the annual legumes ranged from 20 kg/ha to 89 kg/ha when assessed after a single harvest at anthesis; for the perennial legumes it was from 108 kg/ha to 179 kg/ha following two harvests in June and September. We chose two annual legumes (‘Dixie’ crimson clover and hairy vetch) and two perennial legumes (‘Louisiana S-1’ white clover and ‘Kenland’ red clover) for further evaluation.

The diversity and abundance of insect predators of the bean aphid, Aphis fabae Scop. (Homoptera: Aphididae) on common beans, Phaseolus vulgaris L., were evaluated in Central Kenya. Predators included species in the Orders Coleoptera, Diptera, Hemiptera and Neuroptera. Coccinellidae were the most abundant and diverse group and several species, such as Hippodamia variegata Goeze and Platynaspis capicola Crotch, had hitherto not been recorded on common beans in Kenya. Higher numbers of predators developed on beans during the short rains (up to 540/100 m2 plot) as compared to the long rain season (182/plot) of 1985. In both seasons, numbers of coccinellid larvae were high after the fourth week of bean growth. The study provides evidence of an important build-up of insect predators on beans.

Annual legumes and mixtures of annual legumes and grasses can perform several functions as understory cover crops in pecan orchards, such as providing nitrogenrich organic matter to improve soil fertility, or by sustaining lady beetles and other arthropods that may aid the biological control of pecan pests. Remaining questions concern selection of appropriate plant materials; whether to use cover crops singly or in mixtures; how to ensure reseeding as well as a substantial N contribution; whether, when, and how to use mowing and tillage; and fertilization options. Different considerations apply when dealing with cool- vs. warm-season cover crops. With minor adjustments, growers could adapt present cultural practices to include cool-season cover crops. These could be used throughout the orchard, by establishing appropriate self-reseeding species and avoiding both excessive mowing and indiscriminate placement of N-rich fertilizers. Within alleys, alternating 2-m strips of cool-season cover crops could be tilled in mid to late April or allowed to mature. The tilled strips would supply N to pecan trees immediately, whereas the adjoining untilled (remnant) strips could be mowed after seed is mature, to ensure dispersal of seed and reestablishment of cover crops over the entire alley. Cool-season annual legumes that die or are killed in late spring will probably furnish N and other nutrients at a suitable time, particularly in orchards with sprinkler irrigation. Warm-season cover crops, if desired, should be restricted to alleys to reduce possible competition with pecan. Alleys provide better illumination than do tree rows during periods when pecan trees are in leaf, and the tillage mentioned above will encourage emergence of warm-season cover crops. If these die or are killed in late summer or early fall, timing of N release may not be optimal, in the absence of adequate irrigation. Many options and tradeoffs need to be explored before choosing a cover-crop system. Attimes, several objectives may appear to conflict, and even delicately-managed mixtures of species may not fulfill all the desired functions.

Aleurolobus barodensis (Maskell) was more abundant than Neomaskellia andropogonis Corbett on sugar-cane in Pakistan. Azotus delhiensis Lal, Eretmocerus sp., Prospaltella flava Shafee, Amitus sp. and Tetrastichus sp. parasitized nymphs and pupae of A. barodensis and E. serius Silvestri those of N. andropogonis. Chrysopa carnea Steph., Brumoides suturalis (F.), Catana parcesetosa (Sic.), Oenopia sauzeti Muls., Scymnus sp. and S. nubilus Muls. preyed on the immatures of the aleurodids.

In Peshawar valley where the crop is aerially sprayed twice a year to control Chilo infuscatellus Sn., Tryporyza nivella F., A. barodensis and Pyrilla spp., the rate of parasitism on A. barodensis was very low and its density was very high, being 330 immatures/leaf at Charsadda and 425 at Mardan in November 1979. At Sialkot (Punjab) where the crop is not sprayed, the rate of parasitism, mainly by A. delhiensis and Amitus sp., at this time was 22% and the density of A. barodensis was 70.4 immatures/leaf. The maximum rate of parasitism was 71% in January. The abundance of A. barodensis in Peshawar valley seems to be due to the mortality of natural enemies because of aerial spraying, therefore, judicious use of insecticides and redistribution of natural enemies in these areas is recommended.