The effects of simulated and natural rainfall on cochineal insects (Homoptera: Dactylopiidae): colony distribution and survival on cactus cladodes

ABSTRACT. 1. The distribution of Dactylopius opuntiae (Cockerell) colonies on Opuntia ficus-indica (L.) Miller cladodes collected during a prolonged drought, was recorded for the proximal and distal portions of the cladodes, for the exposed and sheltered surfaces, the edges, and in relation to the proximity of the colonies to areoles at the base of the thorns.
2. Most of the colonies were found on the more sheltered parts of the cladodes and a disproportionately large number of the colonies on exposed surfaces had settled below the areoles.
3. Individuals in colonies on the exposed surfaces were all protected by a relatively hard, compacted wax cover and were thus less vulnerable to simulated rainfall than those on sheltered surfaces. Thorns provided some protection from simulated rainfall for individuals in colonies on exposed surfaces, but not for individuals on sheltered surfaces.
4. Natural rainfall greatly reduced the numbers of colonies, but did not significantly change the distribution of the colonies on the cactus cladodes.
5. Rainfall does not explain the observed distribution of D. opuntiae colonies on the cladodes, but does explain the poor performance of cochineal insects as biological control agents of prickly pear cacti in high rainfall areas.     

Life history and natural enemy associations of calico scale (Homoptera: Coccidae) in Kentucky

Seasonal phenology of calico scale, Eulecanium cerasorum (Cockerell), was monitored for 3 yr on various deciduous tree species in central Kentucky. Infestations were found on 16 host species in six plant families. Calico scale is a univoltine parthenogenic species that overwinters as second instars on bark. Nymphs molted to adult females around mid-April and began producing eggs in late April. Mean fecundity ranged from 3,728 to 4,654 eggs per female, depending on host plant species. Date of first crawler hatch in 2001-2003 ranged from 21 to 26 May, corresponding to a mean accumulation of 818 +/- 2 Celsius degree-days (DDC), calculated from 1 January and a base of 4.4 degrees C. This value predicted crawler hatch within 2 d in Lexington, KY, in 2004. Crawler dispersal lasted 2 to 3 wk. Upon hatching, crawlers move to leaves where they feed during summer. Crawlers primarily settled on the abaxial side of leaves and their within-leaf distribution varied between different tree species. Settled crawlers molted in mid-July and second instars remained on leaves until late September through mid-October, when they returned to bark to overwinter. On hackberry, Celtis occidentalis L., they were concentrated toward the basal end of shoots, primarily because leaf flush continued beyond the end of the crawler dispersal period. Crawler distribution did not differ between upper and lower canopy zones. Fourteen species of parasitoids and a coccinellid beetle were reared from individual scales. Monitoring with sticky traps in tree canopies confirmed that targeting crawlers with insecticides during late May or June would not coincide with peak flight activity of the scale's primary parasitoids.     

Selection of large host plants for oviposition by a monophagous leaf beetle: nutritional quality or enemy‐free space?

Abstract.  1. Oviposition site selection is crucial for the reproductive success of herbivorous insects. According to the preference–performance hypothesis, females should oviposit on host plants that enhance the performance of their offspring. More specifically, the plant vigour hypothesis predicts that females should prefer large and vigorously growing host plants for oviposition and that larvae should perform best on these plants.
2. The present study examined whether females of the monophagous leaf beetle Cassida canaliculata Laich. (Coleoptera: Chrysomelidae) prefer to oviposit on large host plant individuals of the meadow clary and whether large host plants are of higher nutritional quality than small host plants. Subsequently, it was tested whether the female preference correlates with offspring performance and survival.
3. In the field, females preferred large host plant individuals for oviposition and host plant quality, i.e. leaf nitrogen content, was significantly higher in leaves of large than of small host plants.
4. In the laboratory, larval development time was shorter on leaves of large host plant individuals than on small host plant individuals, but this could not be shown in the field.
5. However, a predator-exclusion experiment in the field resulted in a higher survival of larvae on large host plants than on small host plants when all predators had free access to the plants. On caged host plants there was no difference in survival of larvae between plant size categories.
6. It is concluded that females of C. canaliculata select oviposition sites that enhance both performance and survival of their offspring, which meets the predictions of the plant vigour hypothesis.     

Aerial distribution, flight behaviour and eggload: their inter-relationship during dispersal by the sweetpotato whitefly

1. The aerial distribution of Bemisia tabaci Gennadius (the sweetpotato whitefly) was studied during the early ascent phase of flight, to test the degree to which dispersal patterns reflect the flight behaviour of individuals.
2. Marked whiteflies were trapped at four heights between 0 and 7·2 m above fallow ground, and at six distances between 0 and 100 m from the insect source. Insects were trapped during a 2–3 h period after the initiation of flight activity during the summers of 1995 and 1996.
3. Analysis of trap catch data revealed a clear negative exponential relationship between height and aerial distribution, and a slightly weaker negative power relationship between distance and aerial distribution. Marked insects were caught in the uppermost traps adjacent to the source, indicating that a portion of the population had a strong capacity for ascent out of the flight boundary layer.
4. Eggload decreased with the height, but not the distance, at which whiteflies were trapped. Mean eggload close to the ground was significantly greater than that for those trapped at 4·8 and 7·2 m, supporting the hypothesis that there is a trade-off between flight and oogenesis in weak-flying insects.
5. Air temperatures during the trapping periods were positively correlated with the proportion of male and female B. tabaci caught in the highest traps, but not in the most distant traps.
6. The significance of these results for accurate prediction of whitefly dispersal is discussed, and the importance of individual's behaviour in determining dispersal patterns of small insects is emphasized.     

Dispersal ability and host-plant characteristics influence spatial population structure of monophagous beetles

Abstract.  1. Dispersal plays an integral role in determining spatial population structure and, consequently, the long-term survival of many species. Theoretical studies indicate that dispersal increases with population density and decreasing habitat stability. In the case of monophagous insect herbivores, the stability of host-plant populations may influence their spatial population structure.
2. The tallgrass prairie in Iowa, U.S.A. is highly fragmented and most prairie insects face a landscape with fewer habitat patches and smaller host-plant populations than 150 years ago, potentially making dispersal between patches difficult. Some herbivores, however, use native plant species with weedy characteristics that have increased in abundance because of disturbances.
3. Mark–recapture data and presence–absence surveys were used to examine dispersal and spatial population structure of two monophagous beetles with host plants that exhibit different population stability and have responded differently to fragmentation of tallgrass prairie.
4. Chrysochus auratus Fabricius exhibits a patchy population structure and has relatively large dispersal distances and frequencies. Its host plant is variable locally in time and space, but is more abundant than 150 years ago. The other species, Anomoea laticlavia Forster, exhibits a metapopulation or non-equilibrium population structure and has relatively small dispersal distances and frequencies. Its host-plant populations are stable in time and space.
5. The results indicate that dispersal ability of monophagous beetles reflects the life-history dynamics of their host plants, but the spatial population structure exhibited today is strongly influenced by how the host plants have responded to the fragmentation process over both time and space.     

Polyphenic wax production in Abacarus hystrix(Acari: Eriophyidae), and implications for migratory fitness

Abstract. . Wax production in the grass-feeding eriophyid Abacarus hystrix (Nalepa) is seasonally variable; enlarged lateral and dorsal bands of wax filaments develop in adults of summer generations. Wax filaments increase the total surface area of summer-generation mites, without a concomitant increase in cuticular surface-to-volume ratio. Genetic and environmental interactions in changes in the rate of wax production were examined. High temperatures, but not changes in photoperiod, were found to stimulate increased rates of wax production. The degree of wax filamentation was independent of the genetic origin of mites, hence filaments are polyphenic, rather than polymorphic, in origin. The role of wax filaments in the regulation of water-loss and in drag maximization were experimentally investigated. The relative survival of waxed and unwaxed mites was compared at various temperature and humidity combinations, and the influence of wax filaments on surface drag was assessed by comparing the slopes of regression lines of cuticular surface area relative to terminal velocity of waxed and unwaxed mites. Significantly greater survival times in waxed compared to unwaxed mites at low relative humidities suggest mat wax filaments have a function in the regulation of water-loss. Significantly lower mean terminal velocities in waxed mites indicated that total drag relative to cuticular surface area was greater in waxed than unwaxed mites, probably as a result of increased skin friction generated by wax filaments. The increased non-cuticular surface area created by wax filaments may be advantageous during summer-migrations of A.hystrix by increasing wind shear stress on the mites at the leaf margin, enhancing buoyancy while air-borne and reducing the rate of desiccation-induced mortality.     

Progressive restriction of host plant exploitation along a climatic gradient: the willow psyllid Cacopsylla groenlandica in Greenland

1. Host plant selection by the endemic willow psyllid Cacopsylla groenlandica was studied at eight sites in three locations along a N–S climatic gradient in west Greenland.
2. C. groenlandica oviposited and developed on four willow species: Salix glauca, S. arctophila, S. uva-ursi and S. herbacea. Development took place on female catkins of all species and on leaf shoots of S. glauca .
3. At sites in southern Greenland, C. groenlandica fully utilized all four willow species. However, there was progressive reduction of host plant species and tissues exploited as latitude increased, such that towards the northern limit of its range the psyllid became highly specialized, feeding only on female catkins of S. glauca , despite the presence of alternative hosts. Where plant growth was tightly constrained within a restricted growing season, the demands of synchronizing with more than one species may have become too great, resulting in specialization on a single host.
4. The probable mechanisms producing specialization were between-species differences in host plant phenology, linked to variation in the growing season, set against the partially non-overlapping geographical distribution patterns of the host willows.
5. A small degree of phenological asynchrony between host plants can be advantageous for the insect. Under favourable conditions the reproductive season can be extended by using a series of host plants with staggered phenologies. Some 'early' hosts also create predator-free space. However, as asynchrony increases, the advantages disappear as the insect becomes unable to exploit the full range of host options.
6. The importance of host plant synchrony in the life histories of arctic insects is discussed.     

Trade-off in oviposition strategy: choosing poor quality host plants reduces mortality from natural enemies for a salt marsh planthopper

Abstract.  1. Both host plant nutrition and mortality from natural enemies have been predicted to significantly impact host plant selection and oviposition behaviour of phytophagous insects. It is unclear, however, if oviposition decisions maximise fitness.
2. This study examined whether the salt marsh planthopper Pissonotus quadripustulatus prefers higher quality host plants for oviposition, and if oviposition decisions are made so as to minimise mortality at the egg stage.
3. A controlled laboratory experiment and 4 years of field data were used to assess the rates of planthopper oviposition on higher quality 'green' and lower quality 'woody' stems of the host plant Borrichia frutescens . The numbers and percentages of healthy eggs and eggs that were killed by parasitoids or the host plant were recorded.
4. In all years, including the laboratory experiment, Pissonotus planthoppers laid more eggs on lower quality woody stems than on higher quality green stems. While host plant related egg mortality was higher in woody stems, the percentage of eggs parasitised was much greater in green stems. This resulted in a lower total mortality of eggs on woody stems.
5. The results of this study demonstrate that, although Pissonotus prefers lower quality host plants for oviposition, this actually increases fitness. These data seem to support the enemy free space hypothesis, and suggest that for phytophagous insects that experience the majority of mortality in the egg stage, oviposition choices may be made such that mortality is minimised.     

Kin-biased dispersal behaviour in the mango shield scale, Milviscutulus mangiferae

When fitness decreases with increasing density in a habitat, dispersal behaviour is expected to evolve. To avoid competition between kin, dispersal behaviour based on kin recognition should be more likely to occur when the individuals in a habitat are closely related. I tested this prediction with first-instar larvae (crawlers) of the mango shield scale, Milviscutulus mangiferae. The body size of adult females, a measure of fecundity, was larger when only one female was present on a leaf than when two were present. When I placed two crawlers on a leaf, they emigrated more frequently when they were siblings than when they were not related. I discuss the implication of the results for kin recognition in thelytokous parthenogenetic animals. Copyright 2000 The Association for the Study of Animal Behaviour.     

Leaf shape and the host-finding behaviour of two ovipositing monophagous butterfly species

Abstract. 1. Ovipositing Eurema brigitta (Wallace) and Eurema herla (W. S. Macleay) butterflies were followed in the field to determine the nature and extent of the pre-alighting discrimination shown by these insects towards their only host plant, Cassia mimosoides (L).
2. Both species tended to search in areas where plants with long, thin leaves (primarily grasses) were less common, on average, than in randomly placed quadrats. Both species were also more likely to alight on non-host plants with leaves of a similar size and shape to those of the host plant than on non-hosts with leaf shapes dissimilar to that of the host.
3. The search behaviour of these monophagous insects was not so specialized that the butterflies never alighted on non-hosts; in fact the majority of alightings were on non-host plants and the pre-alighting discrimination shown by these insects is clearly not the prime behavioural determinant of their monophagy.     

Colonization of growing fruit by the armored scales Lepidosaphes beckii, Parlatoria pergandii and Aonidiella aurantii (Hom., Diaspididae)

Abstract:  The process of invasion of the fruit surface by three species of armored scale insects [ Lepidosaphes beckii (Newman), Parlatoria pergandii Comstock and Aonidiella aurantii (Maskell)], was studied in citrus orchards in València, Spain. Three periods of intense crawler migration were detected annually for L. beckii and P. pergandii . Abundance of crawlers increased throughout the year and they moved outwards in the tree canopy. The fruit was continuously invaded during the period of fruit growth, starting at a very early stage of fruit development, by crawlers of the first generation of all the three species. Differences in fruit invasion and settling behaviour were found between the three species. First-generation crawlers tended to settle under the fruit calyx in L. beckii and P. pergandii , and on the exposed fruit surface in A. aurantii . During the second generation, most A. aurantii scales originated from reproducing females already settled on the fruit, whereas in L. beckii and P. pergandii new scales came preferentially from other tree substrates or from females settled under the fruit calyx. These findings suggest changes to optimize the timing of chemical sprays for controlling the scales.     

The influence of host plant diversity and food quality on larval survival of plant feeding heteropteran bugs

Abstract. 1. In a laboratory experiment, the influence of host plant diversity and food quality, in terms of nitrogen content, on the larval survival of two oligophagous bug species (Heteroptera, Miridae: Leptopterna dolobrata L., Notostira erratica L.) was investigated. Both species are strictly phytophagous and capable of feeding on a wide range of grass species. Moreover, they typically change their host plants during ontogenesis; it has been suggested that this behaviour is a response to the changing protein content of the hosts.
2. To investigate the importance of host plant diversity for these insects, the development of insects reared on grass monocultures was compared with that on mixtures of four grass species. In addition, the host grasses were grown under two nitrogen regimes to test whether nitrogen content is the key factor determining host plant switching.
3. Both species had a significantly higher survival rate when feeding on several host plants but only L. dolobrata showed a significant response to food nitrogen content. Furthermore, there was no correlation between the nitrogen content of the host plants and the survival rate of N. erratica larvae.
4. The study suggests that at least some Stenodemini need a variety of host plants during larval development but that the level of host plant nitrogen is not the main factor responsible for the observed diversity effect.     

Conserved oviposition preferences in alpine leaf beetle populations despite host shifts and isolation

Abstract.  1. Choosing the plant on which to lay their eggs is the last act of care that most female herbivorous insects bestow upon their offspring. These decisions play a pivotal role in insect–plant interactions, placing host preference under strong selection and contributing to the diversity of phytophagous insects as one of the first traits to adapt to new hosts.
2. This study presents a test of whether extreme isolation and exposure to different host plants can produce intra-specific divergence in oviposition preference in alpine insects. Geographic variation should impose selection to fine-tune host plant ranking and specificity to the plants normally encountered, to avoid wasting time during the very limited reproductive season experienced at high altitudes.
3. Beetles from five populations of Oreina elongata differing in host availability were offered three natural hosts: Cirsium spinosissimum , Adenostyles alliariae , and Adenostyles glabra . A novel application of a continuation ratio model (logistic regression) was made to sequential no-choice experiments, combined with quasi-likelihood analysis of multiple-choice experiments.
4. The results show little geographic variation in host plant choice: all populations strongly preferred Cirsium in multiple-choice trials, and in no-choice experiments laid around 47% of their remaining eggs during each stage, almost regardless of the host present.
5. Enemy-free space seems to explain the preference for Cirsium , but isolation and exposure to different plants has clearly not caused local adaptation in host plant ranking or specificity. Reasons for this conservatism despite divergence in other characteristics are discussed.     

Oviposition site selection in Cactoblastis cactorum (Lepidoptera): constraints and compromises

Summary Oviposition by Cactoblastis cactorum on Opuntia ficus-indica and O. aurantiaca was assessed from the positioning of egg sticks on plants in the field. The number of egg sticks laid on O. ficus-indica plants was affected by: (1) plant size; (2) moth emergence near the plant; (3) cladode condition; and (4) plant conspicuousness. These factors contributed towards the clumping of egg sticks on plants. There was no apparent oviposition preference for one of the two host plant species despite the fact that egg predation was higher and fecundity lower on O. aurantiaca. The selection of a site for oviposition on the host plants was influenced by: (1) cladode condition; (2) height above ground; and (3) shelter from wind during oviposition. Succulent cladodes were the favoured sites for oviposition. The evidence suggests that in C. cactorum, oviposition site selection is largely the net result of a compromise between oviposition behaviour selected for increasing the probability of juvenile survival and oviposition behaviour selected for increasing the probability of laying the full complement of eggs. In addition, environmental and physiological factors such as wind and wing-loading, are thought to place constraints on the range of sites available for oviposition.     

Host-plant use in the range expansion of the pine processionary moth, Thaumetopoea pityocampa

Abstract.  1. During range expansions of phytophagous insects, secondary or novel hosts may allow colonisation of areas without primary hosts. Because plant species often differ in their relative attractiveness and suitability for insects, insect preference for, and performance on, these hosts can determine recruitment potential in the current and future expansion areas.
2. This study explores the relative roles of female preference and larval performance in an important pine defoliator, Thaumetopoea pityocampa (Denis & Schiffermüller) (Lepidoptera, Notodontidae), which colonises three Pinus species at its current range margin in the Italian Alps: P. nigra (primary host), P. sylvestris (secondary host), and P. mugo (novel host).
3. Host use patterns in multiple insect populations were studied through choice and no-choice oviposition experiments in cages, field surveys of mixed stands, and laboratory and field monitoring of larval growth and mortality. It was predicted that a specific life-history trait – time limitation of short-lived females to lay a single batch of eggs – would act as a component of female performance, and lead to similar rates of host acceptance in no-choice settings.
4. In the choice experiment, P. nigra was accepted the most frequently while P. sylvestris was accepted the least frequently, confirming nest density patterns in the field. Contrary to prediction, females remained discriminating in no-choice settings in spite of time limitation. In contrast, relative growth rate (RGR) and mortality of larvae did not differ significantly among the three hosts, highlighting a discrepancy between female preference and larval performance.
5. Recruitment potential of T. pityocampa in future expansion into stands of P. sylvestris and P. mugo is evaluated by combining host quality, conservatism in oviposition behaviour, habitat suitability, and the opportunity for local adaptation.     

Trade-off between mobility and fitness in Cydia pomonella L. (Lepidoptera: Tortricidae)

Abstract.  1. The codling moth Cydia pomonella , one of the most important pest insects of apple fruit orchards worldwide, is regarded as a 'sedentary' insect, but the natural populations consist of both mobile and sedentary genotypes that display different dispersal capacity in the field. The current study investigated whether there is a fitness consequence of being mobile in this species by comparing two strains with contrasting levels of mobility obtained through bi-directional selection.
2. Female moths of the sedentary strain were significantly larger, laid more eggs during their lifespan, and both male and female moths of this strain lived longer, when compared with the mobile strain.
3. The age-specific fecundity of female moths from the sedentary strain was significantly greater, in particular between 3 and 5 days after emergence when the peaks of both oviposition and flight capacity occur, relative to the mobile moths, thereby leading to a greater reproductive function.
4. These differences resulted in different demographic parameters between the two strains. The sedentary strain had a significantly greater net reproductive rate ( R _o ) and a higher intrinsic rate of natural increase ( r _m ), although the generation time ( T ) was similar, hence requiring a shorter time to double the population size, in comparison to the mobile strain.
5. This study provides empirical evidence for the cost of being mobile in C. pomonella . Trade-offs between mobility and fitness might have contributed to the existence of genetic variance for dispersal behaviour in the natural populations and the maintenance of relative sedentariness in the species.     

胭脂虫寄生对仙人掌生长的影响

从印榕仙人掌(Opuntia ficus-indica Mill)植株高度、新增茎片数量和新增茎片面积3个方面研究不同密度胭脂虫Dactylopius coccus Costa寄生对仙人掌生长量的影响。结果表明:1)胭脂虫寄生对寄主仙人掌有明显的抑制作用,未被寄生的仙人掌株高、茎片数量和茎片面积的增加均明显大于被寄生的仙人掌;2)随着胭脂虫寄生密度的增加,仙人掌植株生长量的增加相应减少,生长高峰期的出现也会相应推迟;3)胭脂虫的寄生密度应控制在1000头/株以下,过大会导致仙人掌的死亡。     

Learning pays off: influence of experience on host finding and parasitism in Lariophagus distinguendus

Abstract. 1. The influence of experience on egg maturation, parasitism rate, and behaviour during host searching was investigated for Lariophagus distinguendus (Först.) (Hymenoptera: Pteromalidae) parasitizing larvae of the granary weevil Sitophilus granarius (L.) in grains of wheat Triticum aestivum L.
2. Dissection of female parasitoids and parasitism bioassays at high host density revealed that experience with hosts (e.g. by oviposition or by host feeding) is not required either for triggering oogenesis or for oviposition.
3. In parasitism experiments at low host density, when single host-infested grains were offered within a bulk of healthy grains, host finding and parasitism rate were increased by experience.
4. Behavioural observations revealed that searching time required for finding an infested grain was shorter for experienced parasitoids than for naive parasitoids, because travel time from grain to grain is shorter for experienced parasitoids, and because experienced parasitoids spend less time than naive parasitoids on non-infested grains.
5. In conclusion, experience due to host exposure increases parasitism and thereby the fitness of the parasitoids. It is discussed that this increase is more likely due to learning than to different egg load dynamics of experienced parasitoids.     

Evidence of active or passive downwind dispersal in mark–release–recapture of moths

Modelling moth dispersal in relation to wind direction and strength could greatly enhance the role of pheromone traps in biosecurity and pest management applications. Anemotaxis theory, which describes moth behaviour in the presence of a pheromone plume and is used as a framework for such models. Currently, however, that theory includes only three components: upwind, zigzagging, and sideways casting behaviour. We test anemotaxis theory by analysing the data from a series of mark–release–recapture experiments where the wind direction was known and the insects were trapped using an irregular grid of pheromone traps. The trapping results provide evidence of a downwind component to the flight patterns of the released insects. This active or passive downwind dispersal is likely to be an appetitive behaviour, occurring prior to the elicitation of pheromone‐oriented flight patterns (pheromone anemotaxis). Given the potential for significant displacement during downwind dispersal, this component will have impact on final trap captures and should be considered when constructing moth dispersal models.