Ecological and metabolomic responses of plants to deer exclosure in a suburban forest.

Trees and shrubs in suburban forests can be subject to chronic herbivory from abundant white-tailed deer, influencing survival, growth, secondary metabolites, and ecological success in the community. We investigated how deer affect the size, cover, and metabolomes of four species in the understory of a suburban forest in central New Jersey, USA: the woody shrubs Euonymus alatus and Lindera benzoin, the tree Nyssa sylvatica, and the semi-woody shrub Rosa multiflora. For each species, we compared plants in 38 16 m2 plots with or without deer exclosure, measuring proportion cover and mean height after 6.5 years of fencing. We scored each species in all plots for deer browsing over 8 years and assessed selection by deer among the species. We did untargeted metabolomics by sampling leaves from three plants of each species in an equal number of fenced and unfenced plots, conducting chloroform-methanol extractions followed by LC-MS/MS, and conducting statistical analysis on MetaboAnalyst. The proportion of a species browsed ranged from 0.24 to 0.35. Nyssa sylvatica appeared most selected by and susceptible to deer; in unfenced plots, both its cover and mean height were significantly lower. Only cover or height was lower for E. alatus and L. benzoin in unfenced plots, while R. multiflora height was greater. The metabolomic analysis identified 2333 metabolites, which clustered by species but not fencing treatment. However, targeted analysis of the top metabolites grouped by fencing for all samples and for each species alone and was especially clear in N. sylvatica, which also grouped by fencing using all metabolites. The most significant metabolites that were upregulated in fenced plants include some involved in defense-related metabolic pathways, e.g., monoterpenoid biosynthesis. In overbrowsed suburban forests, variation of deer impact on species' ecological success, potentially mediated by metabolome-wide chemical responses to deer, may contribute to changes in community structure.

Woody plant secondary chemicals increase in response to abundant deer and arrival of invasive plants in suburban forests.

Plants in suburban forests of eastern North America face the dual stressors of high white-tailed deer density and invasion by nonindigenous plants. Chronic deer herbivory combined with strong competition from invasive plants could alter a plant's stress- and defense-related secondary chemistry, especially for long-lived juvenile trees in the understory, but this has not been studied. We measured foliar total antioxidants, phenolics, and flavonoids in juveniles of two native trees, Fraxinus pennsylvanica (green ash) and Fagus grandifolia (American beech), growing in six forests in the suburban landscape of central New Jersey, USA. The trees grew in experimental plots subjected for 2.5 years to factorial treatments of deer access/exclosure × addition/no addition of the nonindigenous invasive grass Microstegium vimineum (Japanese stiltgrass). As other hypothesized drivers of plant secondary chemistry, we also measured nonstiltgrass herb layer cover, light levels, and water availability. Univariate mixed model analysis of the deer and stiltgrass effects and multivariate structural equation modeling (SEM) of all variables showed that both greater stiltgrass cover and greater deer pressure induced antioxidants, phenolics, and flavonoids, with some variation between species. Deer were generally the stronger factor, and stiltgrass effects were most apparent at high stiltgrass density. SEM also revealed that soil dryness directly increased the chemicals; deer had additional positive, but indirect, effects via influence on the soil; in beech photosynthetically active radiation (PAR) positively affected flavonoids; and herb layer cover had no effect. Juvenile trees' chemical defense/stress responses to deer and invasive plants can be protective, but also could have a physiological cost, with negative consequences for recruitment to the canopy. Ecological implications for species and their communities will depend on costs and benefits of stress/defense chemistry in the specific environmental context, particularly with respect to invasive plant competitiveness, extent of invasion, local deer density, and deer browse preferences.

Effects of white-tailed deer and invasive plants on the herb layer of suburban forests.

Lack of hunting and predators and proximity to human communities make suburban forests prone to high deer abundance and non-native plant invasions. I investigated these likely drivers of community structure in the herb layers of six suburban forests in one region of New Jersey, USA. In 223 plots I assessed the herb layer response to 2.5 years with or without deer fencing and the early stage of invasion from seed additions of Microstegium vimineum, an invasive, annual grass. Non-native plants and herbaceous native plants were affected very little by fencing or M. vimineum invasion. In contrast, across all forests the combination of deer access and M. vimineum addition had a strongly negative effect on woody native percent cover. Forests differed in overall fencing effects on woody natives; their cover was greater in fenced plots in just three forests, suggesting greater deer pressure in those forests during the experiment. The early invasion by M. vimineum was greatest in two of these same forests, but was not influenced by fencing. Multi-group structural equation modelling compared two groups of forests that differed in vegetation abundance and other characteristics. It paralleled the results above and also showed no negative influence of non-native cover on native cover, even in the forests where non-native cover was greater. It identified a positive effect of light level on herb layer plants in the forests with less vegetation, and also revealed a positive effect of soil water potential (SWP) on non-native plants in the forests with more vegetation, which had higher SWP. These suburban forests within a common region varied widely in native and non-native herb layer abundance, the early success of M. vimineum invasion and the herb layer's response to early invasion and protection from deer.

The Interaction between Competition and Predation: A Meta-analysis of Field Experiments.

Ecologists working with a range of organisms and environments have carried out manipulative field experiments that enable us to ask questions about the interaction between competition and predation (including herbivory) and about the relative strength of competition and predation in the field. Evaluated together, such a collection of studies can offer insight into the importance and function of these factors in nature. Using a new factorial meta-analysis technique, we combined the results of 20 articles reporting on 39 published field experiments to ask whether the presence of predators affects the intensity of competitive effects and to compare the average effects of competition and predation. Across all studies, the effects of competition in the presence of predators were less than in the absence of predators, and the interaction between competition and predation for most response variables was statistically significant. Removal of competitors had much more positive effects on organisms' growth and mass than did exclusion of predators. Predator exclusion had much more beneficial effects on organisms' survival than did competition. The mean effects of competition and predation on density did not differ from one another. The results differed among trophic levels. Further understanding would benefit greatly from more field experiments that manipulate both competition and predation, that focus on a wider range of organisms and environments, that focus on population-level parameters such as density, and that report results more completely, including data such as sample sizes and variances.