Push-pull strategies involve the behavioral manipulation of insect pests and their natural enemies via the integration of stimuli that act to make the protected resource unattractive or unsuitable to the pests (push) while luring them toward an attractive source (pull) from where the pests are subsequently removed. The push and pull components are generally nontoxic. Therefore, the strategies are usually integrated with methods for population reduction, preferably biological control. Push-pull strategies maximize efficacy of behavior-manipulating stimuli through the additive and synergistic effects of integrating their use. By orchestrating a predictable distribution of pests, efficiency of population-reducing components can also be increased. The strategy is a useful tool for integrated pest management programs reducing pesticide input. We describe the principles of the strategy, list the potential components, and present case studies reviewing work on the development and use of push-pull strategies in each of the major areas of pest control.

The Use of Push-Pull Strategies in Integrated Pest Management

A list of 1719 chemical compounds identified from headspace samples of floral scent is presented. The list has been compiled from some 270 published papers, including analyses of 991 species of flowering plants and a few gymnosperms, a sample including seed plants from 90 families and 38 orders. The compounds belong to seven major compound classes, of which the aliphatics, the benzenoids and phenylpropanoids, and, among the terpenes, the mono- and sesquiterpenes, occur in most orders of seeds plants. C5-branched compounds, irregular terpenes, nitrogen-containing compounds, and a class of miscellaneous cyclic compounds have been recorded in about two-thirds of the orders. Sulfur-containing compounds occur in a third of the orders, whereas diterpenes have been reported from three orders only. The most common single compounds in floral scent are the monoterpenes limonene, (E)-β-ocimene, myrcene, linalool, α- and β-pinene, and the benzenoids benzaldehyde, methyl 2-hydroxybenzoate (methyl salicylate), benzyl alcohol, and 2-phenyl ethanol, which occur in 54–71% of the families investigated so far. The sesquiterpene caryophyllene and the irregular terpene 6-methyl-5-hepten-2-one are also common and occur in more than 50% of the families. Orchidaceae are by far the best investigated family, followed by several families known to have many species with strongly scented flowers, such as Araceae, Arecaceae, Magnoliaceae, and Rosaceae. However, the majority of angiosperm families are still poorly investigated. Relationships between floral scent and pollination, chemistry, evolution, and phylogeny are briefly discussed. It is concluded that floral scent chemistry is of little use for phylogenetic estimates above the genus level, whereas the distribution and combinations of floral scent compounds at species and subspecific levels is a promising field of investigation for the understanding of adaptations and evolutionary processes in angiosperms.

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Diversity and Distribution of Floral Scent

The statistical analysis of compositional data

Bee pollinators are currently recorded with many different sampling methods. However, the relative performances of these methods have not been systematically evaluated and compared. In response to the strong need to record ongoing shifts in pollinator diversity and abundance, global and regional pollinator initiatives must adopt standardized sampling protocols when developing large-scale and long-term monitoring schemes. We systematically evaluated the performance of six sampling methods (observation plots, pan traps, standardized and variable transect walks, trap nests with reed internodes or paper tubes) that are commonly used across a wide range of geographical regions in Europe and in two habitat types (agricultural and seminatural). We focused on bees since they represent the most important pollinator group worldwide. Several characteristics of the methods were considered in order to evaluate their performance in assessing bee diversity: sample coverage, observed species richness, species richness estimators, collector biases (identified by subunit-based rarefaction curves), species composition of the samples, and the indication of overall bee species richness (estimated from combined total samples). The most efficient method in all geographical regions, in both the agricultural and seminatural habitats, was the pan trap method. It had the highest sample coverage, collected the highest number of species, showed negligible collector bias, detected similar species as the transect methods, and was the best indicator of overall bee species richness. The transect methods were also relatively efficient, but they had a significant collector bias. The observation plots showed poor performance. As trap nests are restricted to cavity-nesting bee species, they had a naturally low sample coverage. However, both trap nest types detected additional species that were not recorded by any of the other methods. For large-scale and long-term monitoring schemes with surveyors with different experience levels, we recommend pan traps as the most efficient, unbiased, and cost-effective method for sampling bee diversity. Trap nests with reed internodes could be used as a complementary sampling method to maximize the numbers of collected species. Transect walks are the principal method for detailed studies focusing on plant-pollinator associations. Moreover, they can be used in monitoring schemes after training the surveyors to standardize their collection skills.

Measuring bee diversity in different european habitats and biogeographical regions

This paper reviews the literature concerning digestion and nutrient content of pollen. Four topics are addressed in detail: 1) The mechanism of pollen digestion by animals; 2) The efficiency of mechanical and digestive removal of pollen content by various animals; 3) Range and taxonomic distribution of pollen nutrients, and 4) Adaptive hypotheses proposed to associate pollen chemistry with pollinator reward. Studies on the mechanism(s) of pollen digestion remain inconclusive, but suggest that differences in digestibility among pollen types may reflect differences in pollen wall porosity, thickness, and composition. Although hummingbirds reportedly digest pollen very poorly, most animals studied, including those that do not regularly consume pollen, can digest 50–100% of ingested grains. Overlooked and recent research of pollen protein content shows that pollen grains may contain over 60% protein, double the amount cited in some studies of pollen-feeding animals. Adaptive hypotheses that associate pollen starch and pollen caloric content with pollinator reward remain unsubstantiated when critically viewed through the lens of phylogeny.

Pollen nutritional content and digestibility for animals

1 Since the late 1970s, the western flower thrips has spread from its original distribution in western North America to become a major worldwide crop pest.

2 A wide range of data sources have been used to map the original distribution in the U.S.A. and Canada, and the progress of the spread in the U.S.A., Canada, Europe, northern Africa and Australia.

3 The possible reasons for the start of the spread are discussed. The most likely reason is that intensive insecticide use in horticulture in the 1970s and 1980s selected an insecticide resistant strain or strains. These then established in glasshouses across North America and spread from there to Europe, Asia, Africa and Australia.

4 The international spread of the western flower thrips occurred predominantly by the movement of horticultural material, such as cuttings, seedlings and potted plants. Within Europe, an outward spread from the original outbreak in the Netherlands is discernible. The speed of spread was 229 ± 20 km/year.

5 The spread has not been restricted to glasshouses. The western flower thrips has established outdoors in areas with milder winters; for example, across the southern U.S.A., southern Europe and Australia. It also overwinters in some regions with colder winters.

6 Polyphagous phytophagous thrips have many factors predisposing them to become worldwide crop pests, particularly in glasshouses. Some other species that might spread in a similar way to the western flower thrips are listed.

The spread of the western flower thrips Frankliniella occidentalis (Pergande)

 1 This paper analyses catches of flower thrips, grass thrips and predatory flies in water-traps of seven colours



2 A correlation is demonstrated between type of host-plant of thrips and the relative numbers caught by traps of different colours



3 The literature is reviewed and some general relationships with the effectiveness of different trap colours are hypothesized for: non-grass foliage insects and their predators and parasites; grass foliage insects; flower-dwelling insects; predators and parasites not associated with foliage; biting insects; and wood-borers



4 This may permit trap colours to be chosen, in particular circumstances, that are ecologically selective for different types of insect

Ecologically selective coloured traps.

Western flower thrips, Frankliniella occidentalis, first arose as an important invasive pest of many crops during the 1970s-1980s. The tremendous growth in international agricultural trade that developed then fostered the invasiveness of western flower thrips. We examine current knowledge regarding the biology of western flower thrips, with an emphasis on characteristics that contribute to its invasiveness and pest status. Efforts to control this pest and the tospoviruses that it vectors with intensive insecticide applications have been unsuccessful and have created significant problems because of the development of resistance to numerous insecticides and associated outbreaks of secondary pests. We synthesize information on effective integrated management approaches for western flower thrips that have developed through research on its biology, behavior, and ecology. We further highlight emerging topics regarding the species status of western flower thrips, as well as its genetics, biology, and ecology that facilitate its use as a model study organism and will guide development of appropriate management practices.

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Invasion Biology, Ecology, and Management of Western Flower Thrips.

Two major components have been detected in the headspace volatiles of adult male Frankliniella occidentalis (Pergande) (Thysanoptera: Thripidae) that are not present in the headspace volatiles of adult females. The compounds were identified as (R)-lavandulyl acetate and neryl (S)-2-methylbutanoate by comparison with synthetic standards using gas chromatography (GC), GC–mass spectrometry (MS), and chiral GC. Field trials were conducted with synthetic compounds in naturally infested crops of sweet pepper grown in large plastic greenhouses in Spain. The catch of adult females and males on blue sticky traps was increased by neryl (S)-2-methylbutanoate alone or by a 1:1 blend of (R)-lavandulyl acetate and neryl (S)-2-methylbutanoate, but (R)-lavandulyl acetate was not active alone. This is the first identification of an aggregation pheromone in the order Thysanoptera. The possible role of (R)-lavandulyl acetate is discussed.

Identification of a male-produced aggregation pheromone in the western flower thrips Frankliniella occidentalis

Eight species of thrips in three families fed readily on individual grains of pollen. Feeding behaviour is described. Thrips fed on the contents of the grains, but left the pollen wall. In Thrips fuscipennis Halid., the feeding time per grain increased in direct proportion to grain volume up to a grain volume of about 60,000 μm'. but then began to level off. Large grains and tetrads were usually abandoned before all the contents were removed. Mean feeding times ranged from 3 s to 120 s per grain.



Thrips appeared to have alternate periods of feeding and non-feeding. Drops of fluid were sometimes deposited from the end of the abdomen after non-feeding periods. Large amounts of pollen can be consumed; one thrips fed on more than 100 pollen grains in less than 15 min. Pollen-feeding appears to be widespread among flower thrips. The implications for pollination ecology are discussed.

William D. J. Kirk

Papers

The spread of the western flower thrips Frankliniella occidentalis (Pergande)

Ecologically selective coloured traps.

Invasion Biology, Ecology, and Management of Western Flower Thrips.

Identification of a male-produced aggregation pheromone in the western flower thrips Frankliniella occidentalis

Pollen-feeding in thrips (Insecta: Thysanoptera)