Facilitation in the conceptual melting pot

1. Here we present an introduction to this issue's Special Feature arising from the British Ecological Society Symposium: Facilitation in Plant Communities (20–22 April 2009).
2. Papers in the Special Feature demonstrate the benefits that arise from cross-system application of general concepts, for example, the well-known stress gradient hypothesis. Such comparisons challenge our definition of facilitation, as well as our pre-conceptions on the nature of intermediary organisms.
3. We suggest that under some circumstances a clear definition of the two-way nature of interactions is essential, e.g. when considering the evolutionary implications of facilitation. In other cases, however, we can perhaps be more relaxed, e.g. when facilitation is a component of conservation ecology.
4. Synthesis . Overall we believe that establishing facilitation as an independent concept has driven substantial progress towards a clearer understanding of how ecological systems work. Through the links established by work such as that presented in this Special Feature, we believe this field will continue to make rapid progress and aid ecological understanding in general.     

Similar tree seedling responses to shrubs and to simulated environmental changes at Pyrenean and subarctic treelines

Background: Climate, land-use and disturbance regimes are key drivers of treeline dynamics worldwide, but local and regional spatio-temporal patterns indicate that additional factors play an important role. Some studies suggest that shrub-tree interactions control tree seedling recruitment patterns across the treeline ecotone, but little is known about the generality of this interaction.

Aims: We established an experiment in a treeline ecotone in the central Pyrenees to investigate the role of such interactions and other environmental factors on tree seedling growth and survival. It was based on a similar experiment we completed recently in the subarctic Scandes. By comparing local phenomena between both experiments we assessed the generality of the findings across regions with different biogeographic histories and species characteristics.

Methods: We followed the survival and growth of transplanted Pinus uncinata seedlings during three growing seasons in a multi-factorial design (forest vs. treeline, +/– shrub removal, +/– temperature increase and +/– nutrient addition).

Results: There was better seedling growth at the treeline compared to the forest, and the presence of shrubs prevented seedling winter damage and herbivory. Both temperature and nutrient increase enhanced seedling performance.

Conclusions: Although some particular details were exclusive to the Pyrenees or to the Scandes, the similarities indicated general patterns and help to interpret the underlying mechanisms of treeline dynamics in both regions. This convergence of responses points to the potential of the development of a robust mechanistically founded predictive framework for scaling up shrub impacts on treeline dynamics to other regions.     

Mycorrhizal networks mediate overstorey-understorey competition in a temperate forest

In forests, common mycorrhizal networks (CMNs) often connect the roots of neighbouring plants. Observations of material flows between hosts connected by CMNs have given rise to the hypothesis that CMNs limit the negative effects of competition by overstorey trees on seedlings recruiting underneath them. I conducted an experiment in a temperate forest dominated by ectomycorrhizal conifers and hardwoods to isolate the effects of CMNs on the growth and survival of four tree species that co‐occur in the understorey. Ectomycorrhizal networks had strong negative effects on the survival of an arbuscular mycorrhizal species, Acer rubrum, and neutral effects on the survival of three ectomycorrhizal species, Betula allegheniensis, Pinus strobus, and Tsuga canadensis. CMNs had positive effects on the growth of at least one ectomycorrhizal species, P. strobus. Interspecific differences in juvenile responses to CMNs may influence forest community development, promoting coexistence of some tree species while limiting the success of others.     

Low impact of climate change on subalpine grasslands in the Swiss Northern Alps

While phenological shifts and migration of isolated species under climate change have already been observed on alpine summits, very few studies have focused on community composition changes in subalpine grasslands. Here we use permanent plots monitored since 1954 and precisely located phytosociological censuses from 1970 to study compositional changes of subalpine grasslands in two distinct regions of the Swiss Northern Alps. In both areas, warming trends during the monitoring period were associated with changes in land management (abandonment of goat and sheep pasturing or grazing replaced by mowing). Old and recent inventories were compared with correspondence analyses (CA). Ecological indicator values, community‐affinities and biological traits of the species were used to infer the factors responsible for triggering the observed changes. In both regions, subalpine grasslands were stable with smaller changes than have previously been observed in alpine environments. Only a few species appeared or disappeared and changes were generally limited to increasing or decreasing frequency and cover of certain taxa. At one site, grazing abandonment favored fallow species. Some of these species were located at their upper altitudinal distribution limits and may have spread because of rising temperatures. In both areas, declining species were predominantly alpine and low‐growing species; their decline was probably due to increased competition (e.g., shadow) with more vigorous subalpine taxa no longer limited by grazing. We conclude that vegetation communities can respond rapidly to warming as long as colonization is facilitated by available space or structural change. In the subalpine grasslands studies, changes were mainly driven by land management. These communities have a dense vegetation cover and newly arriving herbaceous species preferring warmer conditions may take some time to establish themselves. However, climate disturbances, such as exceptional drought, may accelerate community changes by opening gaps for new species.     

Morphological plasticity following species-specific recognition and competition in two perennial grasses

Morphological characteristics and biomass allocation of two perennial grasses, Pseudoroegneria spicata (Pursh) A. Löve ssp. spicata (bluebunch wheatgrass) and Agropyron desertorum (Fisch. ex Link) Schult. (crested wheatgrass), were compared under different competition and nutrient treatments. The competitive responses of two plants grown in containers under field conditions were assessed in monocultures and mixtures in two experiments using different scales of nutrient application. In the Small-Scale Experiment, a localized fertilization was applied in the rooting zone between two plants; in the Large-Scale Experiment the entire container was supplied with nutrients. Agropyron responded more vigorously to fertilization than did Pseudoroegneria, but based on the relative performance of Agropyron in monoculture and mixture, it was not superior to Pseudoroegneria in resource competition. Pseudoroegneria was apparently able to recognize neighboring plants as either conspecifics or individuals of the other species. The responses included changes in shoot architecture, root morphology, and allocation between roots and shoots. Agropyron generally did not exhibit such morphological flexibility. In field plot plantings of 4-yr-old tussocks similar shoot differences were seen in Pseudoroegneria. There was, however, no indication of superior resource competition for Agropyron. Thus, any early advantage of Agropyron in vigorous growth of young plants in response to nutrients was apparently lost by the time the plants had reached this stage of development. Morphological and allocation flexibility of Pseudoroegneria may have compensated for slower, less vigorous growth. If species-specific recognition and morphological plasticity are common in nature, this complicates our attempts to understand mechanisms of competition.