Mayura Takada

Indirect effects of herbivory by deer reduce abundance and species richness of web spiders

Abstract: We examined the abundance and species richness of web spiders in forests with and without deer to test the hypothesis that herbivory by deer simplifies habitat structure, thereby reducing the number of web spiders. The number of individuals, the number of species, and the availability of potential web sites were all lower where deer were present. The decrease in the abundance of spiders in the presence of deer was more prominent in large species. The species richness standardized by rarefaction was still lower where deer were present in one of two seasons, suggesting that the reduced number of species in the presence of deer was not simply a by-product of the decreased number of individuals. Web site availability was positively correlated with the number of individuals as well as with the number of species when both forest types were combined. However, prey availability, estimated by the number of insects captured with sticky traps, did not differ significantly between sites with and without deer. Thus, the decrease in spiders was most likely caused by indirect non-trophic effects of herbivory that were mediated by the simplification of habitat structure, not by a decrease in prey abundance.

HABITAT PROVISIONING FOR ABOVEGROUND PREDATORS DECREASES DETRITIVORES

Although provisioning of habitat by ecosystem engineers is one of the most common biological interactions, previous studies have mostly focused on facilitative or bottom-up processes. Here we show that engineering effects can indirectly strengthen top-down effects mediated by predator abundance. We conducted a small-scale manipulative field experiment and broad-scale field observations of the plant, web spider, and detrital insect system in forest understory habitats. In the field experiment, artificially increasing architectural elements enhanced the abundance of spiders by providing physical support for web building. Moreover, aerial insects derived from the detrital food web decreased in response to increased spider abundance. As artificial architecture per se did not affect aerial detritivores, these results indicate that ecosystem engineering indirectly strengthens top-down effects mediated by predators. In field observations conducted in 12 cedar forests, path analyses supported the importance of an indirect pathway originating from understory vegetation complexity to spider abundance and to aerial detritivores. The effect size of spiders on detrital insects was similar in the field experiment and in the observations. These results indicate that the engineering effects of plants cascade to detrital insects through web spiders across different scales.

Contrasting Responses of Web-Building Spiders to Deer Browsing Among Habitats and Feeding Guilds

Abstract We examined web-building spider species richness and abundance in forests across a deer density gradient to determine the effects of sika deer browsing on spiders among habitats and feeding guilds. Deer decreased the abundance of web-building spiders in understory vegetation but increased their abundance in the litter layer. Deer seemed to affect web-building spiders in the understory vegetation by reducing the number of sites for webs because vegetation complexity was positively correlated with spider density and negatively correlated with deer density. In contrast, the presence of vegetation just above the litter layer decreased the spider density, and deer exerted a negative effect on this vegetation, possibly resulting in an indirect positive effect on spider density. The vegetation just above the litter layer may be unsuitable as a scaffold for building webs if it is too flexible to serve as a reliable web support, and may even hinder spiders from building webs on litter. Alternatively, the negative effect of this vegetation on spiders in the litter may be as a result of reduced local prey availability under the leaves because of the reduced accessibility of aerial insects. The response to deer browsing on web-building spiders that inhabit the understory vegetation varied with feeding guild. Deer tended to affect web-invading spiders, which inhabit the webs of other spiders and steal prey, more heavily than other web-building spiders, probably because of the accumulated effects of habitat fragmentation through the trophic levels. Thus, the treatment of a particular higher-order taxon as a homogeneous group could result in misleading conclusions about the effects of mammalian herbivores.

Regional differences in the morphology of a shrub Damnacanthus indicus: An induced resistance to deer herbivory?

We report here the possibility of an induced resistance of a spiny shrub Damnacanthus indicus against deer herbivory. Six characters of D. indicus were compared between regions with and without deer herbivory on the Boso Peninsula. We found that D. indicus in browsed areas produced smaller leaves, thicker spines, and shorter internode distances between spines than those in unbrowsed areas whereas the length of spines and the angle of a pair of spines did not differ significantly. It is likely that D. indicus shows an induced resistance by producing smaller leaves, and by increasing stoutness of spines and spine density.

Facilitation of a Native Pest of Rice, Stenotus rubrovittatus (Hemiptera: Miridae), by the Non-Native Lolium multiflorum (Cyperales: Poaceae) in an Agricultural Landscape

ABSTRACT Source populations of polyphagous pests often occur on host plants other than the economically damaged crop. We evaluated the contribution of patches of a non-native meadow grass, Lolium multiflorum Lam. (Poaceae), and other weeds growing in fallow fields or meadows as source hosts of an important native pest of rice, Stenotus rubrovittatus (Matsumura) (Hemiptera: Miridae), in an agricultural landscape of northern Japan. Periodical censuses of this mirid bug by using the sweeping method, vegetation surveys, and statistical analysis revealed that L. multiflorum was the only plant species that was positively correlated with the density of adult S. rubrovittatus through two generations and thus may be the most stable and important host of the mirid bug early in the season before the colonization of rice paddies. The risk and cost of such an indirect negative effect on a crop plant through facilitation of a native pest by a non-native plant in the agricultural landscape should not be overlooked.

Facilitation of ground-dwelling wolf spider predation on mirid bugs by horizontal webs built by Tetragnatha spiders in organic paddy fields

Abstract Trait-mediated effects of predators can impact prey population dynamics by affecting prey behavior. The mirid bug Stenotus rubrovittatus (Matsumura) (Hemiptera: Miridae), a major insect pest in Japanese rice production, usually remains in the upper layer of paddies to feed on rice ears. However, the mirids are frequently trapped by horizontal webs of Tetragnatha spp. spiders, which are highly abundant in organic rice paddies, and fall to the bottom layers of paddies where they are preyed upon by ground-dwelling predators. It is hypothesized that Tetragnatha spp. spiders facilitate bug predation by wolf spiders through trait-mediated effects, in which their horizontal webs force the bugs onto or near the ground and thereby into the hunting zones of wolf spiders. Molecular gut-content analysis of 619 wolf spiders coupled with field measurements revealed that the number of wolf spiders that tested positive for mirid bug predation increased significantly with the density of Tetragnatha spp. spiders in the paddies. We also observed a positive relationship between Tetragnatha spp. abundance and total cover by their webs in paddies. We identified the potential for an unexpected interaction between an herbivorous insect pest and ground-dwelling spiders that usually inhabit different microhabitats in paddy fields by focusing on trait-mediated effects of webs built by Tetragnatha spp. Because spider webs occupy a certain proportion of the available space in terrestrial ecosystems, consideration of trait-mediated effects on interactions between flying insects and other predators may lead to a better understanding of local food webs.

Comparison of generalist predators in winter- flooded and conventionally managed rice paddies and identification of their limiting factors

Winter-flooding of rice paddies without the application of agricultural chemicals is attracting attention as a new agricultural method for enhancing the habitat conditions of wintering waterfowl in rice paddy ecosystems throughout Japan and east Asia. Conditions in these paddies are expected to result in restoration of not only the winter habitats of waterfowl but also those of other taxonomic groups during the rice growing season. In this study, we tested whether the diversity of summer spiders––ubiquitous predators in rice paddies––was higher in the winter-flooded paddies than in the conventional ones by conducting field measurements in 31 winter-flooded and 7 conventional paddies. Limiting factors of spiders in the winter-flooded paddies were then examined. Results revealed that both the density and species richness of spiders were significantly higher in the winter-flooded paddies than in the conventional ones both before and after the insecticide application against pecky rice bug Stenotus rubrovittatus (Matsumura)(Hemiptera: Miridae) to conventional paddies. In addition, spider density and species richness in the winter-flooded paddies correlated with the availability of two prey groups––chironomids and other nematocera. These findings suggest that in the winter-flooded paddies the diversity of generalist predators is higher than in the conventional ones during the rice-growing season and that the combination of management at both the landscape and field level is likely more effective for increasing spider abundance in winter-flooded paddies.

Can spines deter deer browsing?: a field experiment using a shrub Damnacanthus indicus

We examined the defensive function of spines of Damnacanthus indicus against deer herbivory by experimentally removing spines in the field where deer density is high. Individuals of D. indicus whose spines had been removed had a higher probability of being browsed by deer than control individuals. In addition, plant height was significantly lower for individuals that were browsed than those that were not. These results support the notion that spines of D. indicus have a function to deter deer browsing. This study is the first to demonstrate experimentally the defensive function of spines against large mammals in temperate areas.

Landscape Effects of a Non-Native Grass Facilitate Source Populations of a Native Generalist Bug, Stenotus rubrovittatus, in a Heterogeneous Agricultural Landscape

Abstract Non-native plant species can provide native generalist insects, including pests, with novel food and habitats. It is hypothesized that local and landscape-level abundances of non-native plants can affect the population size of generalist insects, although generalists are assumed to be less sensitive to habitat connectivity than specialists. In a heterogeneous landscape in Japan, the relationship between the density of a native pest of rice (Stenotus rubrovittatus (Matsumura) (Heteroptera: Miridae)) and the abundance of Italian ryegrass (Lolium multiflorum Lam. (Poales: Poaceae)), a non-native meadow grass known to facilitate S. rubrovittatus, was analyzed. Statistical analyses of data on bug density, vegetation, and the spatial distribution of fallow fields and meadows dominated by Italian ryegrass, obtained by field surveys, demonstrated that local and landscape-level abundances of Italian ryegrass (the unmowed meadow areas within a few hundred meters of a sampling plot) positively affected bug density before its immigration into rice fields. Our findings suggest that a generalist herbivorous insect that prefers non-native plants responds to spatial availability and connectivity of plant species patches at the metapopulation level. Fragmentation by selective mowing that decreases the total area of source populations and increases the isolation among them would be an effective and environmentally-friendly pest management method.

Dead or alive? Sexual conflict and lethal copulatory interactions in long-jawed Tetragnatha spiders

Yuki G. Baba,a, Akio Tanikawa,b Mayura B. Takada,c and Kyoko Futamid aInstitute for Agro-Environmental Sciences NARO, 3-1-3 Kannondai, Tsukuba-shi, 305–8604 Ibaraki, Japan, bLaboratory of Biodiversity Science, School of Agriculture and Life Sciences, The University of Tokyo, Yayoi, Bunkyo-ku, Tokyo 113–8657, Japan, cInstitute for Sustainable Agro-ecosystem Services, Graduate School of Agricultural and Life Sciences, The University of Tokyo, 1-1-1 Midoricho, Nishitokyo, Tokyo 188-0002, Japan, and dInstitute of Tropical Medicine, Nagasaki University, 1-12-4 Sakamoto, Nagasaki 852–8523, Japan