Masahiro Osakabe

Deleterious Effects of Uv-B Radiation on Herbivorous Spider Mites: They Can Avoid It by Remaining on Lower Leaf Surfaces

ABSTRACT The herbivorous spider mite Tetranychus urticae usually remains on the lower leaf surfaces of its host plants. Although terrestrial animals are generally thought to be well protected from damage because of UV radiation, insect herbivory frequently increases when solar UV-B (280–315 nm) radiation is attenuated. As UV transmission through leaves is generally low because of the accumulation of compounds that act as selective sunscreens (e.g., phenolics), we hypothesized that T. urticae avoids solar UV-B radiation by staying on lower leaf surfaces. We examined whether artificial UV irradiation and solar UV affected the survival and reproduction of T. urticae and whether staying on lower leaf surfaces was beneficial to their performance under ambient UV radiation. We found that T. urticae was not well protected from UV-B radiation, because artificial UV-B irradiation strongly decreased survivorship and egg production. More importantly; compulsory solar UV irradiation treatments also had lethal effects on T. urticae, whereas the mites could avoid them if they remained on the lower leaf surfaces of their host plants. These results showed that access to habitats protected from sunlight, such as lower leaf surfaces, is likely essential for T. urticae survival under ambient UV-B radiation. The lethal effects of solar UV radiation may also affect the population dynamics of spider mites, and habitat (resource) limitation may increase the probability of interspecific interactions, such as competition and predation. In turn, the occurrence of these interactions in sheltered areas may be associated with observed increases in herbivory under conditions of solar UV-B-attenuation.

Spectrum‐specific Damage and Solar Ultraviolet Radiation Avoidance in the Two‐spotted Spider Mite

The spatial distribution of the two‐spotted spider mite Tetranychus urticae Koch is biased toward the lower surfaces of leaves as compared with the upper leaf surfaces on their host plants. Because of the deleterious effects of solar ultraviolet (UV) irradiation, we hypothesized T. urticae remains on lower leaf surfaces as an adaptation to avoid solar UV radiation (UVR). We examined the effects of solar UVR components on females and tested whether spatial distribution was associated with solar UVR avoidance. Attenuation of solar UVR using UV opaque film increased fecundity and reduced the movement of females from the upper to the lower leaf surfaces. In contrast, diverting solar UVR to the lower leaf surface using a light reflection sheet caused the mites to move from the lower to the upper leaf surfaces; however, attenuated UV reflection did not, suggesting that they occupy the lower leaf surface to avoid solar UVR. In monochromatic UVR tests, no eggs hatched when placed under 280–300u2003nm radiation, whereas almost all eggs hatched at 320–360u2003nm. Adult females, however, did not avoid wavelengths of 280 and 300u2003nm, but avoided 320–340u2003nm. We conclude that T. urticae exploit UVA information to avoid ambient UVB radiation.

Do plant mites commonly prefer the underside of leaves

The adaxial (upper) and abaxial (lower) surfaces of a plant leaf provide heterogeneous habitats for small arthropods with different environmental conditions, such as light, humidity, and surface morphology. As for plant mites, some agricultural pest species and their natural enemies have been observed to favor the abaxial leaf surface, which is considered an adaptation to avoid rain or solar ultraviolet radiation. However, whether such a preference for the leaf underside is a common behavioral trait in mites on wild vegetation remains unknown. The authors conducted a 2-year survey on the foliar mite assemblage found on Viburnum erosum var. punctatum, a deciduous shrub on which several mite taxa occur throughout the seasons, and 14 sympatric tree or shrub species in secondary broadleaf-forest sites in Kyoto, west–central Japan. We compared adaxial–abaxial surface distributions of mites among mite taxa, seasons, and morphology of host leaves (presence/absence of hairs and domatia). On V. erosum var. punctatum, seven of 11 distinguished mite taxa were significantly distributed in favor of abaxial leaf surfaces and the trend was seasonally stable, except for Eriophyoidea. Mite assemblages on 15 plant species were significantly biased towards the abaxial leaf surfaces, regardless of surface morphology. Our data suggest that many mite taxa commonly prefer to stay on abaxial leaf surfaces in wild vegetation. Oribatida displayed a relatively neutral distribution, and in Tenuipalpidae, the ratio of eggs collected from the adaxial versus the abaxial side was significantly higher than the ratio of the motile individuals, implying that some mite taxa exploit adaxial leaf surfaces as habitat.

Tolerance to Solar Ultraviolet-B Radiation in the Citrus Red Mite, An Upper Surface User of Host Plant Leaves

Plant‐dwelling mites are potentially exposed to solar ultraviolet‐B (UVB) radiation that causes deleterious and often lethal effects, leading most mites to inhabit the lower (underside) leaf surfaces. However, in species of spider mite belonging to the Genus Panonychus, a substantial portion of individuals occur on upper leaf surfaces. We investigated whether the upper leaf surfaces of citrus trees are favorable for P. citri, and to what extent they are tolerant to UVB radiation. If eggs are not adequately protected from UVB damage, females may avoid ovipositing on the upper surfaces of sunny leaves. To test this, we conducted laboratory experiments using a UVB lamp, and semioutdoor manipulative experiments. As a result, P. citri eggs are tolerant to UVB. Field studies revealed that the ratio of eggs and adult females on upper leaf surfaces were larger for shaded than for sunny leaves. However, 64–89% of eggs hatched successfully even on sunny upper leaf surfaces. Nutritional evaluation revealed that whether on sunny or shaded leaves, in fecundity and juvenile development P. citri reaped the fitness benefits of upper leaf surfaces. Consequently, P. citri is tolerant to UVB damage, and inhabiting the upper surfaces of shaded leaves is advantageous to this mite.

Seasonal changes in the deleterious effects of solar ultraviolet-B radiation on eggs of the twospotted spider mite, Tetranychus urticae (Acari: Tetranychidae)

Solar ultraviolet-B (UVB) radiation has deleterious effects on plant-dwelling mites. We assessed the biological effects of UVB radiation on the eggs of the twospotted spider mite, Tetranychus urticae Koch, under both near ambient (UV+) and UV-attenuated (UV−) conditions from spring to autumn and compared them to the effects of temperature and humidity. The ambient daily UVB irradiance increased from January to August and then decreased rapidly until December, whereas egg hatchability under UV+ was lowest in April (10.7%) and increased almost linearly until October (74.9–92.3%). In contrast, hatchability under UV− was consistently high (96.2–99.8%) through all seasons. For UV+, the stepwise multiple linear regression analysis supported the negative correlation of hatchability with cumulative UVB irradiance during egg periods (cumulative dose), but did not support that with the mean daily UVB irradiance (dose rate), suggesting that UVB-induced mortality in T. urticae eggs is cumulative dose dependent rather than dose rate dependent. The high mortality in April may have reflected the slower development caused by the relatively lower temperature and higher UVB radiation, increasing the cumulative dose, while the low mortality in October may have reflected the faster development caused by the relatively higher temperature and lower UVB radiation, decreasing the cumulative dose.

Web-Mediated Interspecific Competition Among Spider Mites

Some spider mites, such as Tetranychus spp. and Amphitetranychus spp., create complicated webs (CWs), whereas others, such as Panonychus spp., produce little webs (LWs). We verified whether interspecific competition occurred between CW and LW mites via habitat arrangement under laboratory conditions. The complicated webs produced by CW mites clearly inhibited juvenile development in LW mites, whereas there was no effect of LW mites on CW mites. In oviposition site choice tests, both CW and LW females preferred the lower surface of leaves to the upper surface. The preference of LW mites for the lower leaf surface, even in the presence of CW mite webs, suggests that the costs of amensalism are outweighed by the possible benefits, such as avoiding rain. These findings show that the shift in mite species composition from LW to CW mites can occur as a consequence of the interspecific association between spider mites via their webs, without pesticide applications or the presence of natural enemies.

Amensalism via webs causes unidirectional shifts of dominance in spider mite communities

Competitive displacement is considered the most severe consequence of interspecific competition; if a superior competitor invades the habitat of an inferior species, the inferior species will be displaced. Most displacements previously reported among arthropods were caused by exotic species. The lack of investigation of displacement among native species may be due to their apparently harmonious coexistence, even if it is equivalent to an outcome of interspecific association. A seasonal change in the species composition of spider mites, from Panonychus ulmi to Tetranychus urticae, is observed in apple trees worldwide. Previous laboratory experiments have revealed amensal effects of T. urticae on P. ulmi via their webs. Using manipulation experiments in an orchard, we tested whether this seasonal change in species composition occurred as the result of interspecific competition between these spider mites. Invasion by T. urticae prevented an increase in P. ulmi densities throughout the experimental periods. Degree of overlap relative to the independent distribution on a leaf-surface basis (ωS) changed from positive to negative with increasing density of T. urticae. T. urticae invasion drove P. ulmi toward upper leaf surfaces (competitor-free space). The niche adjustment by P. ulmi occurred between leaf surfaces but not among leaves. Our findings show that asymmetrical competition between T. urticae and P. ulmi plays an important role in this unidirectional displacement and that the existence of refuges within a leaf produces the apparently harmonious coexistence of the mites and obscures their negative association.

Vulnerability and behavioral response to ultraviolet radiation in the components of a foliar mite prey–predator system

Ambient ultraviolet-B (UVB) radiation impacts plant-dwelling arthropods including herbivorous and predatory mites. However, the effects of UVB on prey–predator systems, such as that between the herbivorous spider mite and predatory phytoseiid mite, are poorly understood. A comparative study was conducted to determine the vulnerability and behavioral responses of these mites to ultraviolet (UV) radiation. First, we analyzed dose–response (cumulative irradiance-mortality) curves for the eggs of phytoseiid mites (Neoseiulus californicus, Neoseiulus womersleyi, and Phytoseiulus persimilis) and the spider mite (Tetranychus urticae) to UVB radiation from a UV lamp. This indicated that the phytoseiid mites were more vulnerable than the spider mite, although P. persimilis was slightly more tolerant than the other two phytoseiid mites. Second, we compared the avoidance behavior of adult female N. californicus and two spider mite species (T. urticae, a lower leaf surface user; Panonychus citri, an upper leaf surface user) in response to solar UV and visible light. N. californicus actively avoided both types of radiation, whereas P. citri showed only minimal avoidance behavior. T. urticae actively avoided UV as well as N. californicus but exhibited a slow response to visible light as well as P. citri. Such variation in vulnerability and avoidance behavior accounts for differences in the species adaptations to solar UVB radiation. This may be the primary factor determining habitat use among these mites on host plant leaves, subsequently affecting accessibility by predators and also intraguild competition.

Evolutionary Aspects of Acaricide-Resistance Development in Spider Mites

Although the development of acaricide resistance in spider mites is a long-standing issue in agricultural fields, recent problems with acaricide resistance may be characterized by the development of complex- and/or multiresistance to acaricides in distinct classes. Such complexity of resistance is not likely to be a single mechanism. Pesticide resistance involves the microevolution of arthropod pests, and population genetics underlies the evolution. In this review, we address the genetic mechanisms of acaricide resistance evolution. We discuss genetic diversity and linkage of resistance genes, relationships between mite habitat and dispersal, and the effect of dispersal on population genetic structure and the dynamics of resistance genes. Finally, we attempt to present a comprehensive view of acaricide resistance evolution and suggest risks under globalization as well as possible approaches to managing acaricide resistance evolution or emergence.

The Bunsen-Roscoe reciprocity law in ultraviolet-B-induced mortality of the two-spotted spider mite Tetranychus urticae.

To determine whether the Bunsen-Roscoe reciprocity law (i.e., the extent of photochemical effects is determined by cumulative irradiance) is applicable to ultraviolet-B (UVB) damage in the twospotted spider mite Tetranychus urticae, egg hatchability and survival of individuals were assessed after irradiation with a UVB lamp using various combinations of intensity and time length. A positive linear correlation between probit mortality and cumulative UVB irradiance was detected in eggs, larvae, teleiochrysalis females, and adult females, regardless of UVB intensity (0.19-0.58 Wm(-2)). LD50 values were clearly higher in adult females, followed by teleiochrysalis females, larvae, and eggs. In eggs, reciprocity was obeyed not only at the UVB intensities listed above, but also at very low UVB intensity (0.014-0.023 Wm(-2)). Such reciprocity in the negative effects of UVB radiation was also observed for the developmental rate of juveniles and egg production of adult females. However, the LD50 value of eggs obtained using the UVB lamp (0.58 kJm(-2)) was lower than that elicited by solar UVB radiation in a previous outdoor experiment (about 50 kJm(-2)). These results suggest that a photoreactivation mechanism plays an important role in the survival of this mite under solar radiation.