Pheromone trapping models for insect pest control

Pheromones have recently been suggested for use in insect pest control. We formulate a discrete version of a model with virgin females (or female equivalent of pheromone) in traps, and explore conditions under which this mechanism can eradicate the pest. Our model is density independent, thus when eradication is not possible, the pest increases without bound, in reality density dependent effects limit population growth. Successive refinements incorporating various features, show that delayed mating together with monogamy has a strong influence on the outcome, whereas other aspects of mating behaviour are relatively unimportant. All our models show that when males are in excess control is impossible but when some virgins are left unmated each day the outcome depends on the parameters. As the birth rate or survivorship increase, control by this method becomes more difficult.     

Potential for suppressingDiprion similis (Hymenoptera: Diprionidae) with pheromone trapping: A population model

The virgin female introduced pine sawfly, Diprion similis (Hartig ), produces a powerful sex pheromone capable of attracting males. Pending chemical identification of the substance, a pilot study in male annihilation is under consideration. A mathematical model was developed to assess the feasibility of population suppression using pheromone-baited traps. Departure from preexisting population models resulted chiefly from the arrhenotokous nature of the sawfly. The model predicts alternating sawfly generations which undergo first a large shift in the sex ratio but no population reduction, followed by a large decrease in population size with a moderating sex ratio. Four generations of intensive trapping would theoretically be sufficient to eliminate the sawfly from an isolated area.     

Simple mathematical models to describe the rate of mating in insect populations

Simple models are constructed to describe the rate of mating in insect populations. The models are based on the assumption of random mate-searching in a closed habitat, including four parameters, i.e., population size, sex ratio, searching efficiency and male's capacity on mating frequency. The modes of effects of these parameters on the rate of mating are analyzed and some principles deduced are discussed in relation to the mating process in natural populations.     

Mating patterns of a sphragis-bearing butterfly,Luehdorfia japonica leech (Lepidoptera: Papilionidae), with descriptions of mating behavior

Mating frequency of both sexes in a natural population of the papilionid butterfly Luehdorfia japonica was studied with special attention to the role of sphragis in preventing multiple matings by females. Males patrolled continuously within a patchy habitat throughout the warm daylight period in search for females. Mating took place without specialized courtship behavior. Males also attempted to copulate forcibly with previously mated females, but the presence of sphragis and/or the escape reaction of females prevented copulation. There was no specialized mate rejection behavior. Females mated early in their adult life, mainly on the day of emergence, and the frequency of mated females reached 100% within the first two or three weeks of their flight period. Spermatophore counts based on dissections of wild females possessing a sphragis indicated that they had never remated. Males were sexually active throughout their adult life. Male mating frequency was estimated from an index of scale-loss from the claspers and frequencies of males which had not mated, and those which had mated once, twice or three or more times were respectively estimated to be 33.7%, 40.3%, 18.2% and 7.8%.     

The sterile release method for population control with interspecific competition

A differential equations model of competing species with the release of sterile individuals of one of the species is examined. The system is found to have two positive steady states for certain parameter values; one of these is stable and the other is unstable. The system is quite resilient around the stable steady state. The release of steriles causes the nontarget species to increase in numbers. There exists a value of the release rate above which the pest species collapses to extinction. The existence of the competitor species assists the sterile release program since the pest equilibrium at any release rate is lower with the competitor species present than without it; in addition the release rate required to cause collapse of the pest species is lower with the competitor species present than without it. The effects of the parameters on the ease of eradication were examined. It was found that the ideal competitor species should have a high rate of increase, a large carrying capacity and exert strong competitive depression on the pests. The ideal pest would have a low rate of increase, a low carrying capacity and be a poor competitor.     

NM+: software implementing parentage-based models for estimating gene dispersal and mating patterns in plants

NM+ is computer software designed for making inferences on plant gene dispersal and mating patterns by modelling parentage probabilities of offspring based on the spatially explicit neighbourhood model. NM+ requires a sample of mapped and genotyped candidate parents and offspring; however, offspring may optionally be assigned to single maternal parents (forming so-called half-sib progeny arrays). Using maximum likelihood, NM+ estimates a number of parameters, including proportions of offspring due to self-fertilization, pollen immigration from outside of a defined study site, parameters of pollen (and/or seed) dispersal kernels (exponential-power, Weibull, geometric or 2Dt) and selection gradients relating covariates (phenotypic traits) with male (and/or female) reproductive success. NM+ allows for missing data both in parents and in offspring. It accounts for null alleles and their frequencies can optionally be considered as estimable parameters. Data files are formatted in a table-like structure so they can be easily prepared in a spreadsheet software. By default NM+ is for studying plant populations, however, it can be used for any organism as long as data requirements and model assumptions are met. NM+ runs under Windows, but it can be launched under Linux using WINE emulator. NM+ can be downloaded free of charge from http://www.genetyka.ukw.edu.pl/index_pliki/software.htm.     

Combining pheromone-baited and food-baited traps for insect pest control: Effects of additional control by parasitoids

Two age-structured population dynamic models are analyzed in which pheromone-baited trapping and food-baited trapping are used simultaneously to eradicate an insect pest. The pest species is assumed to be under partial control by a host-specific parasitoid species. The two models assume that density-dependent population regulation is accomplished either by host larval competition or by means of oviposition interference among the parasitoids. The two trap types interact in a positive synergistic manner and this combination appears to be very promising as a useful combination of pest control methods. Several features of the system are examined; the feature which appears to cause the greatest problem is the possibility of the parasitoids being attracted to the pheromone or the food traps. In either case, the degree of attraction does not have to be very great to undermine the control effort. It is seen that food trapping becomes indispensible if host pheromone is used by the parasitoids as a host-locating kairomone. If odor in the food traps is used by the parasitoids as kairomone, then the situation appears more optimistic, as the reduction in efficiency of the food traps appears much less than with the pheromone traps when pheromone acts as kairomone.     

Combining pheromone-baited and food-baited traps for insect pest control: Effects of developmental period

An age-structured population dynamics model is presented that incorporates pheromone-trapping and food-trapping as control methods for an insect pest. The model yields the following results. Low rates of pest survivorship allow lower trapping rates for control. Species with long developmental periods are easier to control than those with shorter developmental periods (other factors being equal) due to lower net survival. The rates of pheromone trapping alone for effective control are usually very high. The combination of pheromone and food trapping allows control with much lower trapping rates than either method alone. Even small amounts of immigration of adult pests into the control area renders pheromone control ineffective, whereas food traps suppress both the immigrants and the resident population. Food- (or odor-) baited traps which attract both males and females are only somewhat more efficient than those which attract females alone. The existence of density-dependent population regulation assists the control program substantially, but this assistance declines as food trapping becomes a more important part of the control program. Larval competition strongly affects the required trapping rates for eradication; species in which all larvae exert strong competition are much easier to control than those in whic the younger larvae contribute little to the total competitive depression.     

Aggregation pattern of individuals and the outcomes of competition within and between species: Differential equation models

The influence of spatial distribution pattern on the outcomes of intra- and interspecific competition is studied theoretically. The models developed are the generalized logistic andVolterra equations, whereLloyd 's indices of intra- and interspecies mean crowding were incorporated with their assumed linear relationship to mean density in order to express the intensity of crowding which is really effective to the existing individuals. It is shown that while the increasing patchiness of distribution has a pronounced effect of promoting the intraspecific competition and lowering the equilibrium density for individual populations, it generally relaxes the interspecific competition, making it easy for different species sharing the same niche, which would otherwise be incompatible, to coexist stably. These models thus provide a simplest theoretical basis to explain why many insect populations in nature are kept relatively rare in number and why a number of allied species often coexist freely sharing the same resource, against the “competitive exclusion principle” deduced from the originalVolterra equations.     

STORM: software for testing hypotheses of relatedness and mating patterns

Storm is a software package that allows users to test a variety of hypotheses regarding patterns of relatedness and patterns of mate choice and/or mate compatibility within a population. These functions are based on four main calculations that can be conducted either independently or in the hypothesis-testing framework: internal relatedness; homozygosity by loci; pairwise relatedness; and a new metric called allele inheritance, which calculates the proportion of loci at which an offspring inherits a paternal allele different from that inherited from its mother. STORM allows users to test four hypotheses based on these calculations and Monte Carlo simulations: (i) are individuals within observed associations or groupings more/less related than expected; (ii) do observed offspring have more/less genetic variability (based on internal relatedness or homozygosity by loci) than expected from the gene pool; (iii) are observed mating pairs more/less related than expected if mating is random with respect to relatedness; and (iv) do observed offspring inherit paternal alleles different from those inherited from the mother more/less often than expected based on Mendelian inheritance.     

Analysis of genetic control of mating behavior in screwworm (Diptera: Calliphoridae) males through diallel crosses and artificial selection

Summary The mode of genetic control of male screw-worm (Diptera: Calliphoridae) mating behavior was examined using diallel cross and artificial selection. Diallel crosses showed strong dominance effects, with hybrids being uniformly more successful in copulation than their more inbred parental strains. Weaker additive and reciprocal effects were also noted. Environmental (replicate) effects were highly significant. Regression of array variances and covariances indicated that epistatic interactions or unequal allele distribution during gametogenesis may have occurred and that high courtship propensity polygenes show dominance over low propensity genes. Artificial selection on males from outbred strains from Guatemala and Belize resulted in a decreased number of mating attempts for lines selected for reduced activity, but mating attempts in lines selected for high mating activity did not increase. A combination of inbreeding during the selection cycles as well as selection for recessive traits would explain this response. The two types of experiments were in general agreement, indicating significant dominance and environmental influence on male mating behavior with weaker additive and possible maternal effects.     

Individual optimization efforts and population dynamics: a mathematical model for the evolution of resource allocation strategies, with applications to reproductive and mating systems

We develop a formal framework for the optimal allocation of limited resources that includes and clarifies the interplay between individual optimization and the resulting effects at the population level. As an example, in regard to the evolution of sexual recombination, the paradox of the twofold cost of sex is avoided by distinguishing between the evolution of recombination and the subsequent emergence and stability of different mating types as a result of individual optimization within a population that benefits from recombination.

Effects of the juvenile hormone analogue methoprene and dietary protein on male melon fly Bactrocera cucurbitae (Diptera: Tephritidae) mating success

The effect of access to dietary protein (P) and the topical application of a juvenile hormone analogue (methoprene (M)) on mating behaviour of male melon fly Bactrocera cucurbitae was assessed in the laboratory and in field cages. Age, dietary protein and methoprene application increased the mating success and influenced the mating behaviour. Treatment with methoprene (M+) to protein-deprived (P−) males had only a modest effect on the acceleration of sexual maturity, but application of methoprene (M+) to protein-fed (P+) males greatly accelerated sexual maturity. Protein diet (P+) increased mating success of males in comparison to protein-deprived (P−) males. Protein and methoprene have a synergistic effect on mating behaviour, since M + P+ treated males exhibit reduced mating latency and achieved higher mating in younger ages than methoprene and/or protein-deprived males. Copulation duration was correlated with nutritional status and M + P+ males copulated longer at the age of advanced sexual maturity than M − P+ males. Our results suggest that in this species with a lek mating system, females discriminate between the males based on their sexual signals, which were influenced by protein in the adult diet, methoprene application and age. The results are discussed in the light of mating competitiveness of precocious treated young males and their relevance to Sterile Insect Technique application against this pest species.     

Seeing the forest through the trees: comprehensive inference on individual mating patterns in a mixed stand of Quercus robur and Q. petraea

Background and Aims

Sexual reproduction is one of the most important moments in a life cycle, determining the genetic composition of individual offspring. Controlled pollination experiments often show high variation in the mating system at the individual level, suggesting a persistence of individual variation in natural populations. Individual variation in mating patterns may have significant adaptive implications for a population and for the entire species. Nevertheless, field data rarely address individual differences in mating patterns, focusing rather on averages. This study aimed to quantify individual variation in the different components of mating patterns.

Methods

Microsatellite data were used from 421 adult trees and 1911 seeds, structured in 72 half-sib families collected in a single mixed stand of Quercus robur and Q. petraea in northern Poland. Using a Bayesian approach, mating patterns were investigated, taking into account pollen dispersal, male fecundity, possible hybridization and heterogeneity in immigrant pollen pools.

Key Results

Pollen dispersal followed a heavy-tailed distribution (283 m on average). In spite of high pollen mobility, immigrant pollen pools showed strong genetic structuring among mothers. At the individual level, immigrant pollen pools showed highly variable divergence rates, revealing that sources of immigrant pollen can vary greatly among particular trees. Within the stand, the distribution of male fecundity appeared highly skewed, with a small number of dominant males, resulting in a ratio of census to effective density of pollen donors of 5·3. Male fecundity was not correlated with tree diameter but showed strong cline-like spatial variation. This pattern can be attributed to environmental variation. Quercus petraea revealed a greater preference (74 %) towards intraspecific mating than Q. robur (36 %), although mating preferences varied among trees.

Conclusions

Mating patterns can reveal great variation among individuals, even within a single even-age stand. The results show that trees can mate assortatively, with little respect for spatial proximity. Such selective mating may be a result of variable combining compatibility among trees due to genetic and/or environmental factors.     

Effects of agricultural diversification on the abundance, distribution, and pest control potential of spiders: a review

A review of the literature showed that spider abundance was increased by diversification in 63% of studies. A comparison of diversification modes showed that spider abundance in the crop was increased in 33% of studies by `aggregated diversification' (e.g. intercropping and non-crop strips) and in 80% of studies by `interspersed diversification' (e.g., undersowing, partial weediness, mulching and reduced tillage). It is suggested that spiders tend to remain in diversified patches and that extending the diversification throughout the whole crop (as in interspersed diversification) offers the best prospects for improving pest control. There is little evidence that spiders walk in significant numbers into fields from uncultivated field edges, but diversification at the landscape level serves to foster large multi-species regional populations of spiders which are valuable as a source of aerial immigrants into newly planted crops. There are very few manipulative field studies where the impact of spiders on pests has been measured in diversified crops compared with undiversified controls. It is encouraging, however, that in those few studies an increased spider density resulted in improved pest control. Future work needs are identified.     

Mechanisms of dsRNA uptake in insects and potential of RNAi for pest control: A review

RNA interference already proved its usefulness in functional genomic research on insects, but it also has considerable potential for the control of pest insects. For this purpose, the insect should be able to autonomously take up the dsRNA, for example through feeding and digestion in its midgut. In this review we bring together current knowledge on the uptake mechanisms of dsRNA in insects and the potential of RNAi to affect pest insects. At least two pathways for dsRNA uptake in insects are described: the transmembrane channel-mediated uptake mechanism based on Caenorhabditis elegans’ SID-1 protein and an ‘alternative’ endocytosis-mediated uptake mechanism. In the second part of the review dsRNA feeding experiments on insects are brought together for the first time, highlighting the achievement of implementing RNAi in insect control with the first successful experiments in transgenic plants and the diversity of successfully tested insect orders/species and target genes. We conclude with points of discussion and concerns regarding further research on dsRNA uptake mechanisms and the promising application possibilities for RNAi in insect control.     

Variance in reproductive success and the opportunity for selection in a serially monogamous species: simulations of the mating system of <Emphasis Type="Italic">Tropheus</Emphasis> (Teleostei: Cichlidae)

Sexual selection is believed to play a major role in speciation processes of cichlid fishes in the East African Great Lakes, in particular Lakes Malawi and Victoria, by driving diversification in male nuptial coloration. In Lake Tanganyika, a high rate of color pattern diversification is found in the genus Tropheus, where more than 100, mostly allopatric, color morphs have been described. Whether color pattern differentiation in Tropheus could follow from sexual selection as well, has not yet been conclusively shown. Unlike typical sexually selected species, Tropheus are sexually monomorphic, establish temporary pair bonds and spawn monogamously. Variance in mating success among individuals would still allow for sexual selection, but due to their high population census sizes, parentage data from natural populations are difficult to obtain. Simulations designed according to existing data on mating behavior in Tropheus suggested that variance in male reproductive success can be substantial and lead to levels of opportunity for selection, which are similar to values estimated, for example, from natural populations of sexually dimorphic songbirds. Variance in male success was mostly affected by the way females perceived and reacted to differences between males, and to a lesser extent by the duration of components of the reproductive cycles of males and females. These results indicate that future work on the importance of sexual selection in Tropheus should concentrate on mate choice cues and female mate choice behavior, but does not depend critically on the acquisition of more detailed life history data.     

Assembly rules and community models for unicellular organisms: patterns in diatoms of boreal streams

1. Many studies have addressed either community models (e.g. Clementsian versus Gleasonian gradients) or assembly rules (e.g. nestedness, checkerboards) for higher plant and animal communities, but very few studies have examined different non‐random distribution patterns simultaneously with the same data set. Even fewer studies have addressed generalities in the distribution patterns of unicellular organisms, such as diatoms. 2. We studied non‐randomness in the spatial distribution and community composition of stream diatoms. Our data consisted of diatom surveys from 47 boreal headwater streams and small rivers in northern Finland. Our analytical approaches included ordinations, cluster analysis, null model analyses, and associated randomisation tests. 3. Stream diatom communities did not follow discrete Clementsian community types, where multiple species occur exclusively in a single community type. Rather, diatom species showed rather individualistic responses, leading to continuous Gleasonian variability in community composition. 4. Although continuous variability was the dominating pattern in the data, diatoms also showed significant nestedness and less overlap in species distribution than expected by chance. However, these patterns were probably only secondary signals from species’ individualistic responses to the environment. 5. Although unicellular organisms, such as diatoms, differ from multicellular organisms in several biological characteristics, they nevertheless appear to show largely similar non‐random distribution patterns previously found for higher plants and metazoans.