Exotic Lepidopteran Leafminers in North American Apple Orchards: Rise to Prominence, Management, and Future Threats

The European tentiform leafminers, Phyllonorycter blancardella (Fabricius) and P. mespilella (Hübner) (Lepidoptera: Gracillariidae), have infested apple, Malus domestica Borkhausen, in North America for at least 60–70 years. Unreliable taxonomic methods and lack of voucher specimens, however, have contributed to poor understanding of precisely when, where, and how these leafminers were introduced. Tentiform leafminers developed into significant foliar pests at about the time when adult resistance to broad-spectrum insecticides was detected in the 1970s and 1980s. At present, growers manage leafminers with insecticides and, to a lesser extent, with biological control. Most management programs for orchard pests rely on insecticides that are highly toxic to parasitic wasps, which hampers biological control. New techniques promoted in the context of integrated pest management (IPM) may improve biological control by reducing insecticidal use, but their adoption will depend on cost, labor, need to control other pests, and demands of the marketplace. Management programs must be flexible enough to accommodate changes in the species composition of leafminers and their parasitoids. Procedures to exclude pests may slow the rate of new introductions of leafminers and other pests, but they will not entirely protect the North American apple industry. Once new apple pests reach North America, they face few obstacles to further dispersal. The abundance of potential host plants, the lack of rigorous inspections, and new retail practices may facilitate the dispersal of exotic pests.     

LOCAL ADAPTATION AND AGENTS OF SELECTION IN A MOBILE INSECT

The deme-formation hypothesis states that selection can produce adaptive genetic variation within and among phytophagous insect populations. We conducted three field experiments and tested this prediction by transferring eggs and measuring performance of a mobile leafmining insect, Stilbosis quadricustatella. In Experiment 1, we compared the rate of mine initiation of leafminers transferred to natal and novel sites. In Experiment 2, we compared mine-initiation rate of leafminers transferred to natal and novel host-plant species. In Experiment 3, we compared the mine-initiation rate, mine-completion rate, and sources of mortality of miners transferred to neighboring natal and novel Quercus geminata trees. In the first, second, and third experiments, leafminer larvae initiated significantly more mines at the natal site, on the natal plant species, and on the natal Q. geminata tree, evidence for adaptive differentiation. Furthermore, plant-mediated mortality was significantly lower among miners transferred to natal Q. geminata trees. This result supports a key assumption of the deme-formation hypothesis: insects adapt to the defensive phenotypes of individual trees. However, natural-enemy mortality was significantly higher among miners transferred to natal trees, essentially reversing the plant effect. Therefore, rates of successful mine completion were similar on natal (19%) and novel (17%) trees. This experiment suggests that host plants and natural enemies may represent opposing forces of selection. Leafminers adapted to individual trees may realize a selective advantage only when natural-enemy densities are low.     

多食性斑潜蝇对寄主植物的选择

综述了有关斑潜蝇对寄主植物选择的研究进展。斑潜蝇对寄主植物的选择表现为对植物组织的选择、对植物不同部位叶片的选择、对同种植物不同品种的选择、对不同寄主植物的选择四个方面。斑潜蝇对寄主植物的选择主要受叶表面的物理性质、植物叶的化学物质成分、成长经历等因素的影响。阐述了多食性斑潜蝇寄主植物范围扩大的原因。     

The effect of host instar on the size and sex ratio of the endoparasitoidDacnusa sibirica

Older instars of the leafminer hostChromatomyia syngenesiae Hardy (Diptera: Agromyzidae) at parasitization resulted in larger female adults of the larval-pupal endoparasitoidDacnusa sibirica Telenga (Hymenoptera: Braconidae). The sex ratio in the parasitoid offspring became increasingly female biased and development period decreased with increasing age of the host at parasitization. Older host instars were therefore considered to produce a fitter parasitoid.     

Observations on the mating behavior of the alfalfa blotch leafminer,Agromyza frontella (Rondani) (Diptera: Agromyzidae), and evidence of a female sex pheromone

The mating behavior of Agromyza frontella was studied under laboratory conditions. Adults were able to mate on the day of emergence, with no evident periodicity throughout the photophase. The host plant was essential for mating to occur, its presence affecting female receptivity rather than male copulatory behavior. Males generally entered a stationary phase once in the proximity of a female, before undertaking a final approach. This stationary behavior frequently resulted in male aggregations around a female, and under such conditions males exhibited a characteristic wing vibrating behavior. As male wing vibration was not an essential behavior for successful mating, and rarely occurred during male encounters in the absence of females or when only one male was near a female, it was considered as being primarily a male-male signal. The majority of females that mated exhibited an ovipositor pumping behavior that stimulated the male approach. However, such behavior was not essential to attract mates, as dead females elicited the entire sequence of male mating behavior. This suggested the presence of a cuticular sex pheromone, as reported for other species of higher Diptera. Whole virgin female (<24- h or 3- day-old) hexane extracts applied to male cadavers increased the time males spent on the plant, the number of contacts with the treated cadavers, the incidence of attempted copulations, and the wing vibrating behavior between males compared with controls using untreated cadavers. The results obtained indicate that females control copulation in A. frontella and that both semiochemical and visual cues are important in eliciting male mating behavior.     

Towards using of new and safety material against tomato leafminer,Tuta absoluta (Mayrick)

New silica formulations offer expanded possibilities for use in horticultural crops. However, many crop pests are found on the leaf underside, and this is especially challenging when using silica because the substance must have direct contact with the insect to be effective. Silica a new and safety material was used against the second instar larvae of tomato leafminer, Tuta absoluta (Mayrick) alone and combined with other pesticides (chlorfenapyr, spinetoram and chlorantraniliprole). The results showed that when silica was used alone, the percents of mortalities were 51.7, 26.7 and 13.3%. The results showed also chlorantraniliprole was the most toxic compared with the other compounds. The per cents of mortalities were 90, 65 and 30% with the first, second and third concentrations, respectively. When silica was used in a combination with the tested pesticides, the mixture of silica and chlorfenapyr was the most toxic against the second instar larvae of T. absoluta, the percents of mortalities were 100, 76.7 and 60% with the first, second and third concentrations, respectively. The results recommended that silica was moderate toxic to the second instar larvae of T. absoluta when it used alone, so it can be used when this pest under the economic threshold or with integrated pest management programme. Silica has an activation action on the tested pesticides. Although silica was combined with one-fifth of the field rate of the tested pesticides, the toxicity of these pesticides was increased.     

Economic injury level and crop loss assessment for Tuta absoluta (Lepidoptera: Gelechiidae) on different tomato cultivars

Specifying the relationship between pests and their host plants in terms of damage is one of the basic concepts of integrated pest management (IPM) programmes. The crop loss and economic injury level (EIL) of Tuta absoluta (Lepidoptera: Gelechiidae) were determined on different tomato cultivars using gain threshold (field experiment) and regression (semi‐field experiment) methods. By assessing the crop loss in 2015 and 2016, four out of seven infested tomato cultivars had a significant yield loss in terms of weight of total fruits versus the control under the open‐field conditions. However, the total number of tomato cultivars had no significant difference to control treatment. The semi‐field experiment included six treatments (0, 2, 4, 6, 8 and 10 gravid females/plant) and examined two tomato cultivars. The tomato cages with between 6 and 10 gravid females/plant showed the highest number and weight of damaged fruits. Based on pesticide control costs, the market value of tomato and the control efficacy, we determined the EIL for T. absoluta. In the field experiment, the EIL value for T. absoluta on “Petomech,” “Cal JN3,” “Rio Grande” and “Early Urbana Y” cultivars was 4.15, 4.47, 4.70 and 5.04 larvae per plant, respectively. Based on regression equations, we calculated the EIL values for T. absoluta on “Cal JN3” and “Early Urbana Y” to be 4.75 and 5.44 larvae/plant, respectively. Crop loss assessment and EIL are necessary components of cost‐effective IPM programs and can be effective tools for making decisions about the application of pesticides against T. absoluta.     

Natural enemies of Tuta absoluta in the Mediterranean basin,Europe and South America

The tomato leafminer Tuta absoluta (Meyrick) (Lepidoptera: Gelechiidae) represents a global threat to commercial tomato (Solanum lycopersicum L.) production, both in open field and greenhouse. Native to South America, it spread over the Mediterranean Basin, Europe, Africa and part of Asia in only 12 years, and currently it is reported in over 80 countries. Biological control is one of the options for its control and a large number of natural enemies has been reported in association with the pest, both in the areas of origin and of introduction. The egg parasitoid Trichogramma pretiosum, in South America, and the mirid predators Macrolophus pygmaeus and Nesidiocoris tenuis, in Europe and the Mediterranean basin, are used as commercial biocontrol agents. Even if several natural enemies might be promising candidates for biocontrol, their potential role in quantitative pest reduction has been seldom established under practical tomato production conditions.

Since climatic suitability indices predict a high probability for continued invasion by T. absoluta, mainly in China and the USA, there is an urgent need for new control options. In order to minimise the use of broad spectrum insecticides, biocontrol techniques should be considered. As tomato is produced seasonally, augmentative biocontrol seems to be the most effective control option, but pest reduction might be optimised by adding conservation biocontrol, and by combining biocontrol within IPM programmes.

Here, an overview of predators and parasitoids of T. absoluta in South American and Euro-Mediterranean regions, and their biological control efficacy under laboratory, semi-field and field conditions is provided.     

Climate change scenarios and models yield conflicting predictions about the future risk of an invasive species in North America

• 1 The pea leafminer Liriomyza huidobrensis (Blanchard) (Diptera: Agromyzidae) is an invasive species in North America and a serious economic pest on a wide variety of crops. We developed a bioclimatic envelope model (BEM) for this species and examined the envelope's potential location in North America under various future climates.
• 2 We compared the future bioclimatic envelopes for L. huidobrensis using either simple scenarios comprising uniform changes in temperature/precipitation or climate projections from general circulation models (GCMs). Our simple scenarios were: (i) an increase of 0.1°C per degree in latitude with a 20% increase in summer precipitation and a 20% decrease in winter precipitation and (ii) an overall increase of 3°C everywhere, also with the same changes in precipitation. For GCM‐modelled climate change, we used the Canadian Centre for Climate Modelling and Analysis GCM (CGCM2) and the Hadley Centre climate model (HadCM3), each in combination with two scenarios from the Special Report on Emissions Scenarios (A2 and B2).
• 3 The BEM results using the simple scenarios were more similar to each other than to the results obtained using GCM projections. The results were also qualitatively different (i.e. spatially different and divergent) depending on which GCM‐scenario combination was used.
• 4 This modelling exercise illustrates that: (i) results using first approximation simple climate change scenarios can give predictions very different from those that use GCM‐modelled climate projections (comprising a result that has worrying implications for empirical impact research) and that (ii) different GCM‐models using the same scenario can give very different results (implying strong model dependency in projected biological impacts).

     

Visual, semi-quantitative assessments allow accurate estimates of leafminer population densities: an example comparing image processing and visual evaluation of damage by the horse chestnut leafminer Cameraria ohridella (Lep., Gracillariidae)

Abstract: Qualitative or semi-quantitative visual assessments are most often used for estimating population size of herbivorous insects. The precision of these estimates, however, is often difficult to establish. A 'simulation game' with the horse chestnut leafminer, Cameraria ohridella Deschka & Dimic (Lep., Gracillariidae) shows that visual, semi-quantitative assessments can provide accurate information. Damaged areas of 411 horse chestnut leaves collected in 100 sites were closely related to mine numbers despite some variability in mine and leaf size ( R ² = 0.915; n  = 411; P < 0.001). On the basis of this relationship, two methods of population assessment are compared: (i) digital image processing of leaf damage and (ii) visual assessment using a damage key reflecting the relative infested area on each leaf (0, 0%; 1, 0–2%; 2, 2–5%; 3, 5–10%; 4, 10–25%; 5, 25–50%; 6, 50–75%; 7, 75–100%). Both methods used to estimate damage presented a similar, close relationship to the 'real' numbers of mines ( R ² = 0.858; n  = 777; P < 0.001 for image processing and R ² = 0.905; n  = 777; P < 0.001 for visual assessment). The potential of using visual assessments as an accurate and fast method in situ at the tree scale is discussed.