Abstract. Removal of top-down control on herbivores can result in a trophic cascade where herbivore pressure on plants results in changes in plant communities. These altered plant communities are hypothesized to exert bottom-up control on subsequent herbivory via changes in plant quality or productivity. But it remains untested whether top-down perturbation causes long term changes in plants that ricochet back up the new food chain that depends on them. In a large-scale, 30-yr controlled field experiment, we show that 10 yr of top-down control of an ungulate herbivore (white-tailed deer, Odocoileus virginianus) created contrasting forest tree communities exerting bottom-up effects that ricochet back up 3 trophic levels 20-30 yr later. Higher ungulate densities during stand initiation caused significant reductions in tree species diversity, canopy foliage density, canopy insect density, and bird density in young (ca. 30 yr old) forests. Because recruitment of trees from seedlings to the canopy occurs over a relatively brief period (ca. 10 yr), with membership in the canopy lasting an order of magnitude longer, our results show that even short-term perturbations in ungulate density may cause centuries-long disruptions to forest ecosystem structure and function. In documenting this five-step trophic ricochet, we unite key concepts of trophic theory with the extensive literature on effects of ungulate overabundance. As predators decline and ungulate herbivores increase worldwide, similar impacts may result that persist long after herbivore density becomes effectively managed.

A physical method is described for the rapid isolation of plant trichomes, with emphasis on stalked glandular types. The technique involved breaking frozen trichomes with powdered dry ice and collection of glandular heads by sieving from larger tissue fragments. This method was applied to several plants that bear similar stalked trichomes: geranium (Pelargonium), potato (Solanum tuberosum), tomato (Lycopersicon esculentum), squash (Cucurbita pepo), and velvetleaf (Abutilon theophrasti). The tissue preparation was of sufficient quality without further purification for biochemical and molecular studies. The preparation maintained the biochemical integrity of the trichomes for active enzymes and usable nucleic acids. A large quantity of tissue can be harvested; for example, 351 milligrams dry weight of glandular trichomes were harvested from geranium pedicels in 12 hours. The utility of the technique was demonstrated by examining the fatty acid composition of tall glandular trichomes of geraniums, Pelargonium xhortorum L.H. Bailey. These purified cells contained high concentrations of unusual omega5-unsaturated fatty acids, proportionally 23.4% of total fatty acids in the trichomes. When the trichomes were removed, the supporting tissue contained no omega5-fatty acids, thereby unequivocally localizing omega5-fatty acids to the trichomes. Because omega5-fatty acids are unique precursors for the biosynthesis of omega5-anacardic acids, we conclude that anacardic acid synthesis must occur in the glandular trichomes.

Invasive species often cause enormous economic and ecological damage, and this is especially true for invasive plants in the Asteraceae family. Arbuscular mycorrhizal fungi (AMF) play an important role in the successful invasion by exotic plant species because of their ability to promote growth and influence interspecific competition. However, few studies have evaluated the effects of invasive Asteraceae species on AMF diversity and how feedback mechanisms during competition with native species subsequently affect the accumulation of nutrient resources. Two exotic Asteraceae, Ambrosia artemisiifolia and Bidens pilosa, were monitored during competition with a native grass species, Setaria viridis, which is being replaced by these exotic species in natural areas around the study site. From these species continuously maintained in a field plot for 5 years, we collected the rhizosphere soil and cloned and identified soil AMF. Furthermore, AM fungal spores were isolated from rhizosphere soil of the two invasive species and used as inoculum in greenhouse experiments, to compare growth and nutrient accumulation during competition. The results indicate that although the AMF diversity in the rhizosphere soil of A. artemisiifolia and B. pilosa differed, the three most abundant species (Septoglomus viscosum, Septoglomus constrictum, Glomus perpusillum) were identical. The addition of AMF inoculum changed the competition between the plants, increasing the competitive ability of the invasives and decreasing that of the native. The results show a similar AMF community composition between A. artemisiifolia and B. pilosa, increased AMF root colonization of the invasive species during competition, AMF-enhanced N accumulation, and AMF-facilitated competitive growth of the invasive species.

Externally feeding phytophagous insect larvae (i.e., caterpillars, here, larval Lepidoptera and sawflies, Hymenoptera: Symphyta) are important canopy herbivores and prey resources in temperate deciduous forests. However, composition of forest trees has changed dramatically in the eastern United States since 1900. In particular, browsing by high densities of white-tailed deer (Odocoileus virginianus) has resulted in forests dominated by browse-tolerant species, such as black cherry (Prunus serotina), and greatly reduced relative abundance of other tree species, notably pin cherry (Prunus pensylvanica) and birches (Betula spp.). To quantify effects of these changes on caterpillars, we sampled caterpillars from 960 branch tips of the 8 tree species that comprise 95% of trees in Allegheny hardwood forests: red maple (Acer rubrum), striped maple (Acer pensylvanicum), sugar maple (Acer saccharum), sweet birch (Betula lenta), yellow birch (Betula allegheniensis), American beech (Fagus grandifolia), black cherry, and pin cherry. We collected 547 caterpillar specimens that belonged to 66 Lepidoptera and 10 Hymenoptera species. Caterpillar density, species richness, and community composition differed significantly among tree species sampled. Pin cherry, nearly eliminated at high deer density, had the highest density and diversity of caterpillars. Pin cherry shared a common caterpillar community with black cherry, which was distinct from those of other tree hosts. As high deer density continues to replace diverse forests of cherries, maples, birches, and beech with monodominant stands of black cherry, up to 66% of caterpillar species may be eliminated. Hence, deer-induced changes in forest vegetation are likely to ricochet back up forest food webs and therefore negatively affect species that depend on caterpillars and moths for food and pollination.

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