Dispersal of the sevenspotted lady beetle, Coccinella septempunctata, was measured in a series of mark-release-recapture experiments in Utah alfalfa. In three experiments, samples were taken in a radial pattern around the release point. Released beetles for the most part left the 0.36 ha (68 m diameter) sample area within 24 hours, and their average residence time in the sample area was calculated as 12, 6 and 1.6 h in the three experiments, respectively. The spatial distribution of beetles around the point of release could be described with normal distributions whose variance increased linearly in time with 3.8, 1.1 and 0.34 m2 per hour. In three additional field experiments the departure of marked beetles was compared between sugar-sprayed plots and control plots. Residence time was 20-30% longer in sugar-sprayed plots than in control plots, with mean residences of 5.3, 3.6, and 2.9 h in the sugar-sprayed plots in the three experiments, respectively, and means of 4.4, 2.7, and 2.4 h in the control plots. The density of unmarked beetles rose by a factor of 10-20 in the sugar sprayed plots during the first 4 to 6 hours following early morning spraying of sugar. This rapid and substantial increase in density cannot be explained by the slightly longer residence time in sugar-sprayed plots. We hypothesize that the aggregation in sugar-sprayed plots is mostly due to greatly increased immigration into those plots, in response to volatiles produced by the plant-pest-predator assembly.
The Old World ladybird Coccinella septempunctata has rapidly established itself as an abundant, widespread species throughout North America. Overwintering individuals of this species, and of native ladybirds, were collected from early season alfalfa in northern Utah during the period of initial establishment of the invader (1989 to 1999), and were measured for body size. Adult body size can vary widely within insect species, often reflecting differential success of individuals as immatures in obtaining food. Here I examine patterns of ladybird body size to address two questions associated with the establishment of C. septempunctata: (1) is there evidence for adverse impact on native species?, and (2) why has the invader has been so successful in establishment? As an indirect test of adverse competitive effect of the invader on native species, I determined whether mean body size of adults of the five most common native species (Coccinella tranversoguttata, Hippodamia convergens, H. quinquesignata, H. sinuata, and H. tredecimpunctata) declined over the period 1991-1997 as the invader increased rapidly in abundance. No such decline was observed for any of these species, thus providing no evidence that the invader's establishment has significantly increased scramble competition for food among immature ladybirds. I also compared body size distribution of the invading species with that of native species. The invader was distinctive in having particularly large variation in body size among individuals (i.e., in having relatively high proportions of both unusually large and small individuals). Such results are consistent with the hypothesis that the invader's success derives from being a generalist with much "ecological flexibility" in regard to the conditions under which it engages and succeeds in reproduction.
Given the generalist tendencies of most predatory arthropods, it is widely thought that their impact on a particular prey species in a given habitat (e.g., an insect pest in a crop) will depend frequently on the local availability of other prey (which for omnivorous predators, can include plant resources, such as fruit and pollen). Thus, from a slightly different perspective, aphids, other herbivorous insects, and plants often may interact indirectly by sharing natural enemies. Such interactions may be either negative or positive, as in the concepts of apparent competition and apparent mutualism, and may therefore have variable impact on the herbivores' host plants as well. I examine the different mechanisms for such indirect effects among herbivores as explored in the experimental literature. An impressive collective effort by numerous researchers recently has expanded considerably our empirical base of support for a variety of hypothesized mechanisms; aphids stand out as the most commonly studied subjects in research on these mechanisms. I therefore focus especially on the recent literature of cases involving aphids interacting indirectly with other prey for generalist predators. My remarks are organized by considering how the availability of alternate prey may alter functional and numerical (aggregative and reproductive) responses of predators to focal prey density. Although the distinctions among these different classes of predator responses and the associated indirect effects are often blurred and scale-dependent, this classification remains useful for organizing the diverse ways in which aphids have been found to participate in indirect interactions among prey as mediated by predators. Collectively, the results of the numerous studies reviewed here suggest that many such indirect interactions likely occur frequently in natural settings, with consequences ultimately for host plant performance.
In searching both for food to produce eggs and for suitable oviposition sites, females of aphidophagous ladybirds must be adapted to exploit prey that vary greatly in their occurrence and abundance over both space and time. A simple model of ladybird searching and oviposition behaviour emerged in the 1950s: adult ladybirds are highly mobile in traversing the landscape, but become less active and produce more eggs as their rate of aphid consumption increases. The net result is that most eggs tend to be laid at sites of high aphid density. Laboratory and field experiments and observations over the past several decades have generally supported this basic model, although the linkage between ladybird dispersal activity and local aphid density often appears to be relatively weak. Not all ladybird eggs are laid in patches of high aphid density. Females use resources from limited prey consumption to produce eggs in modest numbers. They may thus be prepared to lay some eggs quickly when they succeed in finding aphids in high numbers, but otherwise they may have little choice but to lay these eggs in suboptimal sites. Upon locating patches of high prey density, females are faced with the decision of how long to remain. The basic model raises the possibility that these females become passively trapped at such patches until local aphid density collapses. Recent studies, however, suggest that detection of oviposition-deterring pheromones may promote earlier departure from prey patches. Females may also have an innate tendency to disperse throughout their lives regardless of local conditions, as a bet-hedging strategy to spread their eggs widely over space. Additional studies are needed to evaluate further the degree to which females actively determine and vary the rhythms of dispersal and reproduction in response to the unpredictable and short-lived nature of populations of their aphid prey