Human activities have increased N availability dramatically in terrestrial and aquatic ecosystems. Extensive research demonstrates that local plant species diversity generally declines in response to nutrient enrichment, yet the mechanisms for this decline remain unclear. Based on an analysis of >900 species responses from 34 N-fertilization experiments across nine terrestrial ecosystems in North America, we show that both trait-neutral and trait-based mechanisms operate simultaneously to influence diversity loss as production increases. Rare species were often lost because of soil fertilization, randomly with respect to traits. The risk of species loss due to fertilization ranged from >60% for the rarest species to 10% for the most abundant species. Perennials, species with N-fixing symbionts, and those of native origin also experienced increased risk of local extinction after fertilization, regardless of their initial abundance. Whereas abundance was consistently important across all systems, functional mechanisms were often system-dependent. As N availability continues to increase globally, management that focuses on locally susceptible functional groups and generally susceptible rare species will be essential to maintain biodiversity.functional traits ͉ metaanalysis ͉ productivity ͉ random loss ͉ rarity

We know a great deal about the plastic responses of plant phenotypes to the abiotic and biotic environment, but very little about the consequences of phenotypic plasticity for plant communities. In other words, we know that plant traits can vary widely for a given genotype, but we know little about the importance of trait-mediated interactions (TMI) among plants. Here, we discuss three major factors that affect the expression of phenotypic plasticity: variation in the abiotic environment, variation in the presence or identity of neighbors, and variation in herbivory. We consider how plastic responses to these factors might affect interactions among plants. Plastic responses to the abiotic environment have important consequences for conditionality in competitive effects, to the point of causing shifts from competitive to facilitative interactions. Because plants show a high degree of plasticity in response to neighbors, and even to the specific identify of neighbors, phenotypic plasticity may allow species to adjust to the composition of their communities, promoting coexistence and community diversity. Likewise, plastic responses to consumers may have various and counterintuitive consequences: induction of plant resistance, compensatory growth, and increased resource uptake may affect interactions among plants in ways that cannot be predicted simply by considering biomass lost to consumers. What little we know about TMI among plants suggests that they should not be ignored in plant community theory. Although work to date on the community consequences of phenotypic plasticity has been hampered by experimental constraints, new approaches such as manipulating phenotypes by using signals instead of actual environmental conditions and the use of transgenic plants should allow us to rapidly expand our understanding of the community consequences of plant plasticity.

In Carpinteria Salt Marsh, Salicornia virginica (pickleweed) grows at lower marsh elevations than does Arthrocnemum subterminalis (Parish's glasswort). Standing biomass of both species was greatest immediately adjacent to their abrupt border, suggesting that conditions for plant growth were best here. We utilized field experiments, in which growth rates of naturally occurring and transplanted individuals of both species were measured in four marsh zones, to investigate the role of edaphic factors and competition in maintaining this zonation pattern. The frequency of flooding, and hence soil waterlogging, was greatest at lower marsh elevations, whereas salinity was highest at higher marsh elevations. Consequently, it was not clear, a priori, which part of the marsh had the most severe physical conditions. In our field experiments, both Salicornia and Arthrocnemum grew better in the two middle marsh zones (high Salicornia zone and Arthrocnemum zone) than in either the low marsh (low Salicornia zone), where flooding was frequent and soils were waterlogged, or the high marsh (transition zone), where soil salinity was extremely high during much of the year and plant water potentials very low. However, Salicornia appeared better able to tolerate flooding, and so persisted in the low Salicornia zone, whereas Arthrocnemum appeared better able to tolerate high salinities, and so persisted in the transition zone. Interspecific competition was most important in the relatively benign middle marsh zones, where each species excluded the other from a portion of this prime habitat. In this marsh, flooding, soil salinity, and competition all interacted to determine plant zonation patterns, but the relative importance of these factors varied at different elevations.

Summary 1We investigated the factors producing zonation patterns of the dominant plants in south-eastern USA salt marshes where Juncus roemerianus dominates the high marsh, and Spartina alterniflora the middle and low marsh. 2 Juncus did not occur naturally in the Spartina zone and performed poorly when transplanted there, irrespective of whether neighbours were present or removed, indicating that its lower limit was set by physical stress. 3 In contrast, although Spartina occurred naturally at low densities in the Juncus zone, it performed well if transplanted there only if neighbours were removed, indicating that its upper limit was set by competition. 4 Parallel laboratory and field manipulations of flooding, salinity and competition indicated that the lower limit of Juncus was mediated by both flooding and salinity, but not by competition. 5 The general mechanisms producing zonation patterns of vegetation in coastal salt marshes may be universal, as suggested by previous studies, but the importance of particular factors is likely to vary geographically. In particular, salinity stress probably plays a much more important role in mediating plant zonation patterns at lower latitudes. 6 Our results suggest that the nature of ecological interactions is likely to vary geographically because of variation in the physical environment, and this variation must be taken into account in order to successfully generalize the results of field studies across geographical scales.

Global energy use and food production have increased nitrogen inputs to ecosystems worldwide, impacting plant community diversity, composition, and function. Previous studies show considerable variation across terrestrial herbaceous ecosystems in the magnitude of species loss following nitrogen (N) enrichment. What controls this variation remains unknown. We present results from 23 N-addition experiments across North America, representing a range of climatic, soil and plant community properties, to determine conditions that lead to greater diversity decline. Species loss in these communities ranged from 0 to 65% of control richness. Using hierarchical structural equation modelling, we found greater species loss in communities with a lower soil cation exchange capacity, colder regional temperature, and larger production increase following N addition, independent of initial species richness, plant productivity, and the relative abundance of most plant functional groups. Our results indicate sensitivity to N addition is co-determined by environmental conditions and production responsiveness, which overwhelm the effects of initial community structure and composition.

We investigated species-specific relationships among two species of vascular epiphytes and ten host tree species in a coastal plain forest in the southeastern United States. The epiphytes Tillandsia usneoides and Polypodium polypodioides were highly associated with particular host species in the field, but host traits that favored colonization were inadequate to fully explain the epiphyte-host associations for either epiphyte. Field transplant experiments that bypassed epiphyte colonization demonstrated that the growth of epiphytes was significantly higher on host tree species that naturally bore high epiphyte loads than on host species with few or no epiphytes. These species-specific relationships were highly correlated with the water-holding capacity of the host tree's bark. Positive and negative effects of throughfall, light attenuation by the canopy, and bark stability did not explain the overall patterns of host specificity, but did correlate with some epiphyte-host species relationships. The relative importance of particular host traits differed between the "atmospheric epiphyte" Tillandsia, and the fern Polypodium, which roots in the bark of its hosts. Species-specific interactions among plants, such as those described here, suggest that communities are more than individualistic assemblages of co-occurring species.

Despite long-standing interest in latitudinal variation in ecological patterns and processes, there is to date weak and conflicting evidence that herbivore pressure varies with latitude. We used three approaches to examine latitudinal variation in herbivore pressure in Atlantic Coast salt marshes, focusing on five abundant plant taxa: the grass Spartina alterniflora, the congeneric rushes Juncus gerardii and J. roemerianus, the forb Solidago sempervirens, and the shrubs Iva frutescens and Baccharis halimifolia. Herbivore counts indicated that chewing and gall-making herbivores were typically > or = 10 times more abundant at low-latitude sites than at high-latitude sites, but sucking herbivores did not show a clear pattern. For two herbivore taxa (snails and tettigoniid grasshoppers), correctly interpreting latitudinal patterns required an understanding of the feeding ecology of the species, because the species common at high latitudes did not feed heavily on plant leaves whereas the related species common at low latitudes did. Damage to plants from chewing herbivores was 2-10 times greater at low-latitude sites than at high-latitude sites. Damage to transplanted "phytometer" plants was 100 times greater for plants transplanted to low- than to high-latitude sites, and two to three times greater for plants originating from high- vs. low-latitude sites. Taken together, these results provide compelling evidence that pressure from chewing and gall-making herbivores is greater at low vs. high latitudes in Atlantic Coast salt marshes. Sucking herbivores do not show this pattern and deserve greater study. Selective pressure due to greater herbivore damage at low latitudes is likely to partially explain documented patterns of low plant palatability to chewing herbivores and greater plant defenses at low latitudes, but other factors may also play a role in mediating these geographic patterns.

Ecological interactions may vary geographically as a function of diversity, density, or per capita interaction strengths, but we know little about the relative importance of these three mechanisms. We examined variation in species richness, abundance, and interactions among leaf‐chewing herbivores and the dominant salt‐marsh plant Spartina alterniflora along the Atlantic Coast of the United States. High‐latitude S. alterniflora plants are more palatable to herbivores than are low‐latitude plants. Within this range of latitude, diversity and density of the dominant leaf‐chewing consumers, snails and grasshoppers, in Spartina‐dominated portions of the marsh varied little. Low‐latitude plants, however, experienced much greater levels of leaf damage from consumers than did high‐latitude plants. Per capita feeding rates of low‐latitude snails (Littoraria irrorata) and grasshoppers (Orchelimum fidicinum) in the laboratory were greater than feeding rates of high‐latitude snails (Melampus bidentatus) and grasshoppers (Conocephalus spartinae). In field experiments, low‐latitude snails strongly suppressed S. alterniflora growth, but high‐latitude snails had no effect on primary production. Thus, latitudinal differences in the effect of herbivores on plants (i.e., interaction strength), driven by differences in per capita effects among species, rather than differences in diversity or density, may contribute to selection for latitudinal differences in plant palatability. Because geographical differences in interaction strength can occur in the absence of differences in diversity or density, linking biogeography with community ecology will require experimental studies that explicitly measure interaction strength at multiple geographic locations.