Temporal variation in arthropod sampling effectiveness: the case for using the beat sheet method in cotton

Predatory insects and spiders are key elements of integrated pest management (IPM) programmes in agricultural crops such as cotton. Management decisions in IPM programmes should to be based on a reliable and efficient method for counting both predators and pests. Knowledge of the temporal constraints that influence sampling is required because arthropod abundance estimates are likely to vary over a growing season and within a day. Few studies have adequately quantified this effect using the beat sheet, a potentially important sampling method. We compared the commonly used methods of suction and visual sampling to the beat sheet, with reference to an absolute cage clamp method for determining the abundance of various arthropod taxa over 5 weeks. There were significantly more entomophagous arthropods recorded using the beat sheet and cage clamp methods than by using suction or visual sampling, and these differences were more pronounced as the plants grew. In a second trial, relative estimates of entomophagous and phytophagous arthropod abundance were made using beat sheet samples collected over a day. Beat sheet estimates of the abundance of only eight of the 43 taxa examined were found to vary significantly over a day. Beat sheet sampling is recommended in further studies of arthropod abundance in cotton, but researchers and pest management advisors should bear in mind the time of season and time of day effects.

Life tables of the processionary caterpillar Ochrogaster lunifer Herrich-Schaffer (Lepidoptera : Thaumetopoeidae) at local and regional scales

A major ongoing debate in population ecology has surrounded the causative factors underlying the abundance of phytophagous insects and whether or not these factors limit or regulate herbivore populations. However, it is often difficult to identify mortality agents in census data, and their distribution and relative importance across large spatial scales are rarely understood. Were, we present life tables for egg batches and larval cohorts of the processionary caterpillar Ochrogaster lunifer Herrich-Schaffer, using intensive local sampling combined with extensive regional monitoring to ascertain the relative importance of different mortality factors at different localities. Extinction of entire cohorts (representing the entire reproductive output of one female) at natural localities was high, with 82% of the initial 492 cohorts going extinct. Mortality was highest in the egg and early instar stages due to predation from dermestid beetles, and while different mortality factors (e.g. hatching failure, egg parasitism and failure to establish on the host) were present at many localities, dermestid predation, either directly observed or inferred from indirect evidence, was the dominant mortality factor at 89% of localities surveyed. Predation was significantly higher in plantations than in natural habitats. The second most important mortality factor was resource depletion, with 14 cohorts defoliating their hosts. Egg and larval parasitism were not major mortality agents. A combination of predation and resource depletion consistently accounted for the majority of mortality across localities, suggesting that both factors are important in limiting population abundance. This evidence shows that O. lunifer is not regulated by natural enemies alone, but that resource patches (Acacia trees) ultimately, and frequently, act together to limit population growth.

Host selection in phytophagous insects: a new explanation for learning in adults

Insect learning can change the preferences an egg laying female displays towards different host plant species. Current hypotheses propose that learning may be advantageous in adult host selection behaviour through improved recognition, accuracy or selectivity in foraging. In this paper, we present a hypothesis for when learning can be advantageous without such improvements in adult host foraging. Specifically, that learning can be an advantageous strategy for egg laying females when larvae must feed on more than one plant in order to complete development, if the fitness of larvae is reduced when they switch to a different host species. Here, larvae benefit from developing on the most abundant host species, which is the most likely choice of host for an adult insect which increases its preference for a host species through learning. The hypothesis is formalised with a mathematical model and we provide evidence from studies on the behavioural ecology, of a number of insect species which demonstrate that the assumptions of this hypothesis may frequently be fulfilled in nature. We discuss how multiple mechanisms may convey advantages in insect learning and that benefits to larval development, which have so far been overlooked, should be considered in explanations for the widespread occurrence of learning.

Habitat complexity, spatial interference, and "minimum risk distribution": a framework for population stability

In the past century, the debate over whether or not density-dependent factors regulate populations has generally focused on changes in mean population density, ignoring the spatial variance around the mean as unimportant noise. In an attempt to provide a different framework for understanding population dynamics based on individual fitness, this paper discusses the crucial role of spatial variability itself on the stability of insect populations. The advantages of this method are the following: (1) it is founded on evolutionary principles rather than post hoc assumptions; (2) it erects hypotheses that can be tested; and (3) it links disparate ecological schools, including spatial dynamics, behavioral ecology, preference-performance, and plant apparency into an overall framework. At the core of this framework, habitat complexity governs insect spatial variance. which in turn determines population stability. First, the minimum risk distribution (MRD) is defined as the spatial distribution of individuals that results in the minimum number of premature deaths in a population given the distribution of mortality risk in the habitat (and, therefore, leading to maximized population growth). The greater the divergence of actual spatial patterns of individuals from the MRD, the greater the reduction of population growth and size from high, unstable levels. Then, based on extensive data from 29 populations of the processionary caterpillar, Ochrogaster lunifer, four steps are used to test the effect of habitat interference on population growth rates. (1) The costs (increasing the risk of scramble competition) and benefits (decreasing the risk of inverse density-dependent predation) of egg and larval aggregation are quantified. (2) These costs and benefits, along with the distribution of resources, are used to construct the MRD for each habitat. (3) The MRD is used as a benchmark against which the actual spatial pattern of individuals is compared. The degree of divergence of the actual spatial pattern from the MRD is quantified for each of the 29 habitats. (4) Finally, indices of habitat complexity are used to provide highly accurate predictions of spatial divergence from the MRD, showing that habitat interference reduces population growth rates from high, unstable levels. The reason for the divergence appears to be that high levels of background vegetation (vegetation other than host plants) interfere with female host-searching behavior. This leads to a spatial distribution of egg batches with high mortality risk, and therefore lower population growth. Knowledge of the MRD in other species should be a highly effective means of predicting trends in population dynamics. Species with high divergence between their actual spatial distribution and their MRD may display relatively stable dynamics at low population levels. In contrast, species with low divergence should experience high levels of intragenerational population growth leading to frequent habitat-wide outbreaks and unstable dynamics in the long term. Six hypotheses, erected under the framework of spatial interference, are discussed, and future tests are suggested.

A general model for host plant selection in phytophagous insects

We develop a general theoretical framework for exploring the host plant selection behaviour of herbivorous insects. This model can be used to address a number of questions, including the evolution of specialists, generalists, preference hierarchies, and learning. We use our model to: (i) demonstrate the consequences of the extent to which the reproductive success of a foraging female is limited by the rate at which they find host plants (host limitation) or the number of eggs they carry (egg limitation); (ii) emphasize the different consequences of variation in behaviour before and after landing on (locating) a host (termed pre- and post-alighting, respectively); (iii) show that, in contrast to previous predictions, learning can be favoured in post-alighting behaviour-in particular, individuals can be selected to concentrate oviposition on an abundant low-quality host, whilst ignoring a rare higher-quality host; (iv) emphasize the importance of interactions between mechanisms in favouring specialization or learning. (C) 2002 Elsevier Science Ltd.

Relationship between oviposition preference and offspring performance in Australian Helicoverpa armigera (Hubner) (Lepidoptera : Noctuidae)

We investigated the oviposition preference and larval performance of Helicoverpa armigera under laboratory conditions to determine if the oviposition preference of individual females on maize, cowpea and cotton correlates with offspring performance on the leaves of the same host plants. The host-plant preference hierarchy of females did not correlate with their offspring performance. Female moths chose host plants that contributed less to their offspring fitness. Plant effects accounted for the largest amount of variation in offspring performance, while the effects of female (family) was low. The offspring of most females (80%, n = 10) were broadly similar, but 20% (two out of 10), showed marked difference in their offspring performance across the host-plant species. Similarly, there was no relation between larval feeding preference and performance. However, like most laboratory experiments, our experi-mental design does not allow the evaluation of ecological factors (for example, natural enemies, host abundance, etc.) that can play an important role in larval performance in the field. Overall, the results highlight the importance of carrying out preference performance analysis on the individual or family level, rather than pooling individuals to obtain average population data.

Life history parameters and biocontrol potential of the mealybug parasitoid Coccidoxenoides peregrinus (Timberlake) (Hymenoptera: Encyrtidae): Asexuality, fecundity and ovipositional patterns

Properties relevant to the ovipositional activity and lifetime productivity of Coccidoxenoides peregrinus (Timberlake) were assessed in the laboratory, to determine the potential of this species as a biocontrol agent against the citrus mealybug, Planococcus citri (Risso). In general, this species has not performed well in orchards, except for a few localities on different continents. The mode of reproduction of C peregrinus is almost entirely thelytokous, with males produced sporadically and at low frequency. The females have both pro-ovigenic and synovigenic traits, which raises questions of the utility of this distinction. The females have a high reproductive potential with 10-20 eggs per day available within the first two days (after a short (12 h) pre-oviposition period), and 80-150 eggs per day thereafter until death at about eight days. Mean lifetime fecundity was 239.2 +/- 34.3 eggs. C peregrinus oviposits across a range of P. citri instars, but productivity relies predominantly on second instar hosts. Second stage (N2) hosts received most eggs in choice (about 52%) and no-choice (about 50%) tests. Most eggs deposited into N2 hosts (82%) reached adult stage whereas only a few of those deposited into N1 and N3 (about 5% each) developed successfully. The haemolymph of parasitised reproductive mealybugs contained granular structures and no parasitoid eggs were found 24 h after exposure to ovipositing wasps. Also, no wasps emerged from parasitised adult hosts that were kept alive. Parasitoid eggs deposited into adult hosts were presumed encapsulated and destroyed, as control mealybugs (not exposed to female wasps) had no granular structures in their haemolymph. Wasps exposed to an abundance of hosts soon started ovipositing, but only for a relatively short time each day (about 2.5 h out of a 7 h exposure). They stopped ovipositing despite eggs judged to be mature in their ovaries. The reproductive output of C peregrinus is discussed in relation to the ecological factors that could influence this output, and the implications for biocontrol are discussed. (C) 2003 Elsevier Inc. All rights reserved.

Learning, odour preference and flower foraging in moths

Floral volatiles play a major role in plant-insect communication. We examined the influence of two volatiles, phenylacetaldehyde and a-pinene, on the innate and learnt foraging behaviour of the moth Helicoverpa armigera. In dual-choice wind tunnel tests, adult moths flew upwind towards both volatiles, with a preference for phenylacetaldehyde. When exposure to either of these volatiles was paired with a feeding stimulus (sucrose), all moths preferred the learnt odour in the preference test. This change in preference was not seen when moths were exposed to the odour without a feeding stimulus. The learnt preference for the odour was reduced when moths were left unfed for 24 h before the preference test. We tested whether moths could discriminate between flowers that differed in a single volatile component. Moths were trained to feed on flowers that were odour-enhanced using either phenylacetaldehyde or a-pinene. Choice tests were then carried out in an outdoor flight cage, using flowers enhanced with either volatile. Moths showed a significant preference for the flower type on which they were trained. Moths that were conditioned on flowers that were not odour-enhanced showed no preference for either of the odour-enhanced flower types. The results imply that moths may be discriminating among odour profiles of individual flowers from the same species. We discuss this behaviour within the context of nectar foraging in moths and odour signalling by flowering plants.

And the beak shall inherit - evolution in response to invasion

The increased demographic performance of biological invaders may often depend on their escape from specifically adapted enemies. Here we report that native taxa in colonized regions may swiftly evolve to exploit such emancipated exotic species because of selection caused by invaders. A native Australian true bug has expanded it host range to include a vine imported from tropical America that has become a serious environmental weed. Based on field comparisons and historical museum specimens, we show that over the past 30-40 years, seed feeding soapberry bugs have evolved 5-10% longer mouthparts, better suited to attack the forest-invading balloon vines, which have large fruits. Laboratory experiments show that these differences are genetically based, and result in a near-doubling of the rate at which seeds are attacked. Thus a native biota that initially permits invasion may rapidly respond in ways that ultimately facilitate control.