Early establishment of trees at the alpine treeline: idiosyncratic species responses to temperature-moisture interactions.

On a global scale, temperature is the main determinant of arctic and alpine treeline position. However on a local scale, treeline form and position vary considerably due to other climatic factors, tree species ecology and life-stage-dependent responses. For treelines to advance poleward or uphill, the first steps are germination and seedling establishment. These earliest life stages may be major bottlenecks for treeline tree populations and will depend differently on climatic conditions than adult trees. We investigated the effect of soil temperature and moisture on germination and early seedling survival in a field experiment in the French Alps near the local treeline (2100 m a.s.l.) using passive temperature manipulations and two watering regimes. Five European treeline tree species were studied: Larix decidua, Picea abies, Pinus cembra, Pinus uncinata and Sorbus aucuparia In addition, we monitored the germination response of three of these species to low temperatures under controlled conditions in growth chambers. The early establishment of these trees at the alpine treeline was limited either by temperature or by moisture, the sensitivity to one factor often depending on the intensity of the other. The results showed that the relative importance of the two factors and the direction of the effects are highly species-specific, while both factors tend to have consistent effects on both germination and early seedling survival within each species. We show that temperature and water availability are both important contributors to establishment patterns of treeline trees and hence to species-specific forms and positions of alpine treelines. The observed idiosyncratic species responses highlight the need for studies including several species and life-stages to create predictive power concerning future treeline dynamics.

A cool experimental approach to explain elevational treelines, but can it explain them?

At alpine treeline, trees give way to low-stature alpine vegetation. The main reason may be that tree canopies warm up less in the sun and experience lower average temperatures than alpine vegetation. Low growth temperatures limit tissue formation more than carbon gain, but whether this mechanism universally determines potential treeline elevations is the subject of debate. To study low-temperature limitation in two contrasting treeline tree species, Fajardo and Piper (American Journal of Botany 101: 788-795) grew potted seedlings at ground level or suspended at tree-canopy height (2 m), introducing a promising experimental method for studying the effects of alpine-vegetation and tree-canopy microclimates on tree growth. On the basis of this experiment, the authors concluded that lower temperatures at 2 m caused carbon limitation in one of the species and that treeline-forming mechanisms may thus be taxon-dependent. Here we contest that this important conclusion can be drawn based on the presented experiment, because of confounding effects of extreme root-zone temperature fluctuations and potential drought conditions. To interpret the results of this elegant experiment without logistically challenging technical modifications and to better understand how low temperature leads to treeline formation, studies on effects of fluctuating vs. stable temperatures are badly needed. Other treeline research priorities are interactions between temperature and other climatic factors and differences in microclimate between tree canopies with contrasting morphology and physiology. In spite of our criticism of this particular study, we agree that the development of a universal treeline theory should include continuing explorations of taxon-specific treeline-forming mechanisms.

Invertebrate herbivory increases along an experimental gradient of grassland plant diversity.

Plant diversity is a key driver of ecosystem functioning best documented for its influence on plant productivity. The strength and direction of plant diversity effects on species interactions across trophic levels are less clear. For example, with respect to the interactions between herbivorous invertebrates and plants, a number of competing hypotheses have been proposed that predict either increasing or decreasing community herbivory with increasing plant species richness. We investigated foliar herbivory rates and consumed leaf biomass along an experimental grassland plant diversity gradient in year eight after establishment. The gradient ranged from one to 60 plant species and manipulated also functional group richness (from one to four functional groups-legumes, grasses, small herbs, and tall herbs) and plant community composition. Measurements in monocultures of each plant species showed that functional groups differed in the quantity and quality of herbivory damage they experienced, with legumes being more damaged than grasses or non-legume herbs. In mixed plant communities, herbivory increased with plant diversity and the presence of two key plant functional groups in mixtures had a positive (legumes) and a negative (grasses) effect on levels of herbivory. Further, plant community biomass had a strong positive impact on consumed leaf biomass, but little effect on herbivory rates. Our results contribute detailed data from a well-established biodiversity experiment to a growing body of evidence suggesting that an increase of herbivory with increasing plant diversity is the rule rather than an exception. Considering documented effects of herbivory on other ecosystem functions and the increase of herbivory with plant diversity, levels of herbivory damage might not only be a result, but also a trigger within the diversity-productivity relationship.

Predicting invertebrate herbivory from plant traits: polycultures show strong nonadditive effects.

Plant functional traits affect the capacity of herbivores to find, choose, and consume plants. However, in a community composed of different plant species, it is unclear what proportion of herbivory on a focal plant is explained by its own traits and which is explained by the characteristics of the surrounding vegetation (i.e., nonadditive effects). Moreover, nonadditive effects could be positive or negative, and it is not known if they are related to community properties such as diversity. To quantify nonadditive effects, we developed four different additive models based on monoculture herbivory rates or plant traits and combined them with measurements of standing invertebrate herbivore damage along an experimental plant diversity gradient ranging from monocultures to 60-species mixtures. In all four models, positive nonadditive effects were detected, i.e., herbivory levels were higher in polycultures than what was expected from monoculture data, and these effects contributed up to 25% of the observed variance in herbivory. Importantly, the nonadditive effects, which were defined as the deviance of the models' predictions from the observed herbivory, were positively correlated with the communities' plant species richness. Consequently, interspecific interactions appear to have an important impact on the levels of herbivory of a community. Identifying those community properties that capture the effects of these interactions is a next important challenge for our understanding of how the environment interacts with plant traits to drive levels of herbivory.

Predicting invertebrate herbivory from plant traits: evidence from 51 grassland species in experimental monocultures.

Invertebrate herbivores can impact plant performance and plant communities. Conversely, plants can affect the ability of herbivores to find, choose, and consume them through their functional traits. While single plant traits have been related to rates of herbivory, most often involving single herbivore-plant pairs, much less is known about which suite of plant traits is important for determining herbivory for a pool of plant species interacting with a natural herbivore community. In this study we measured aboveground herbivore damage on 51 herbaceous species growing in monocultures of a grassland biodiversity experiment and collected 42 different plant traits representing four trait groups: physiological, morphological, phenological, and herbivore related. Using the method of random forests and multiple regression, we identified seven traits that are important predictors of herbivore damage (leaf nitrogen and lignin concentration, number of coleopteran and hemipteran herbivores potentially feeding on the plants, leaf life span, stem growth form, and root architecture); leaf nitrogen and lignin concentration were the two most important predictors. The final model accounted for 63% of the variation in herbivore damage. Traits from all four trait groups were selected, showing that a variety of plant characteristics can be statistically important when assessing folivory, including root traits. Our results emphasize that it is necessary to use a multivariate approach for identifying traits affecting complex ecological processes such as herbivory.