The accumulation of species-specific enemies around adults is hypothesized to maintain plant diversity by limiting the recruitment of conspecific seedlings relative to heterospecific seedlings. Although previous studies in forested ecosystems have documented patterns consistent with the process of negative feedback, these studies are unable to address which classes of enemies (for example, pathogens, invertebrates, mammals) exhibit species-specific effects strong enough to generate negative feedback, and whether negative feedback at the level of the individual tree is sufficient to influence community-wide forest composition. Here we use fully reciprocal shade-house and field experiments to test whether the performance of conspecific tree seedlings (relative to heterospecific seedlings) is reduced when grown in the presence of enemies associated with adult trees. Both experiments provide strong evidence for negative plant-soil feedback mediated by soil biota. In contrast, above-ground enemies (mammals, foliar herbivores and foliar pathogens) contributed little to negative feedback observed in the field. In both experiments, we found that tree species that showed stronger negative feedback were less common as adults in the forest community, indicating that susceptibility to soil biota may determine species relative abundance in these tropical forests. Finally, our simulation models confirm that the strength of local negative feedback that we measured is sufficient to produce the observed community-wide patterns in tree-species relative abundance. Our findings indicate that plant-soil feedback is an important mechanism that can maintain species diversity and explain patterns of tree-species relative abundance in tropical forests.

Recent studies have demonstrated the increasingly important role of lianas (woody vines) in forest regeneration, species diversity and ecosystem-level processes, particularly in the tropics. Mechanisms responsible for the maintenance of liana species diversity could yield new insights into the maintenance of overall species diversity. Lianas contribute to forest regeneration and competition, not only by competing directly with trees, but also by differentially affecting tree species and thus changing how trees compete among themselves. In addition, they contribute considerably to ecosystem-level processes, such as whole-forest transpiration and carbon sequestration. As the rate of tropical forest disturbance increases, they are likely to increase in relative abundance throughout the tropics and the importance of lianas to many aspects of forest dynamics will grow.

One of the main goals in ecology is determining the mechanisms that control the abundance and distribution of organisms. Using data from 69 tropical forests worldwide, I demonstrate that liana (woody vine) abundance is correlated negatively with mean annual precipitation and positively with seasonality, a pattern precisely the opposite of most other plant types. I propose a general mechanistic hypothesis integrating both ecological and ecophysiological approaches to explain this pattern. Specifically, the deep root and efficient vascular systems of lianas enable them to suffer less water stress during seasonal droughts while many competitors are dormant, giving lianas a competitive advantage during the dry season. Testing this hypothesis in central Panama, I found that lianas grew approximately seven times more in height than did trees during the dry season but only twice as much during the wet season. Over time, this dry season advantage may allow lianas to increase in abundance in seasonal forests. In aseasonal wet forests, however, lianas gain no such advantage because competing plants are rarely limited by water. I extend this theory to account for the local, within-forest increase in liana abundance in response to disturbance as well as the conspicuous decrease in liana abundance at high latitudes.

Abstract. Ecosystem productivity commonly increases asymptotically with plant species diversity, and determining the mechanisms responsible for this well-known pattern is essential to predict potential changes in ecosystem productivity with ongoing species loss. Previous studies attributed the asymptotic diversity-productivity pattern to plant competition and differential resource use (e.g., niche complementarity). Using an analytical model and a series of experiments, we demonstrate theoretically and empirically that host-specific soil microbes can be major determinants of the diversity-productivity relationship in grasslands. In the presence of soil microbes, plant disease decreased with increasing diversity, and productivity increased nearly 500%, primarily because of the strong effect of density-dependent disease on productivity at low diversity. Correspondingly, disease was higher in plants grown in conspecific-trained soils than heterospecific-trained soils (demonstrating host-specificity), and productivity increased and host-specific disease decreased with increasing community diversity, suggesting that disease was the primary cause of reduced productivity in speciespoor treatments. In sterilized, microbe-free soils, the increase in productivity with increasing plant species number was markedly lower than the increase measured in the presence of soil microbes, suggesting that niche complementarity was a weaker determinant of the diversityproductivity relationship. Our results demonstrate that soil microbes play an integral role as determinants of the diversity-productivity relationship.

Tropical forests are experiencing large-scale structural changes, the most apparent of which may be the increase in liana (woody vine) abundance and biomass. Lianas permeate most lowland tropical forests, where they can have a huge effect on tree diversity, recruitment, growth and survival, which, in turn, can alter tree community composition, carbon storage and carbon, nutrient and water fluxes. Consequently, increasing liana abundance and biomass have potentially profound ramifications for tropical forest composition and functioning. Currently, eight studies support the pattern of increasing liana abundance and biomass in American tropical and subtropical forests, whereas two studies, both from Africa, do not. The putative mechanisms to explain increasing lianas include increasing evapotranspirative demand, increasing forest disturbance and turnover, changes in land use and fragmentation and elevated atmospheric CO 2 . Each of these mechanisms probably contributes to the observed patterns of increasing liana abundance and biomass, and the mechanisms are likely to be interrelated and synergistic. To determine whether liana increases are occurring throughout the tropics and to determine the mechanisms responsible for the observed patterns, a widespread network of large-scale, long-term monitoring plots combined with observational and manipulative studies that more directly investigate the putative mechanisms are essential.

Summary1 Regeneration in forest canopy gaps is thought to lead invariably to the rapid recruitment and growth of trees and the redevelopment of the canopy. Our observations, however, suggest that an alternate successional pathway is also likely, whereby gap-phase regeneration is dominated by lianas and stalled in a low-canopy state for many years. We investigated gap-phase regeneration in an old-growth tropical forest on Barro Colorado Island (BCI) in Panama to test the following two hypotheses: (i) many gaps follow a pathway in which they remain at a low canopy height and are dominated by lianas and (ii) the paucity of trees in this pathway is a function of liana density. 2 We surveyed a total of 428 gaps of varying ages (c. 5, c. 10, and 13 years old) and identi®ed those which followed the conventional pathway of regeneration and others that remained stalled in a low-canopy state for many years and were dominated by either lianas or palms. Each of these pathways will likely have dierent successional trajectories that will favour the growth of a distinct suite of mature species and ultimately result in contrasting species composition. 3 The successional pathway of liana-dominated, stalled gaps is common throughout the forest. We estimate conservatively that 7.5% of the gaps that form each year will follow this pathway, probably due to the suppression of tree regeneration by lianas, and that many of these stalled gaps will persist for much longer than 13 years. Consequently, a high proportion of gaps in the forest at any given time will be stalled. Furthermore, liana tangles, which persist in the tropical forest understorey for extended periods of time, almost certainly originate in these gaps. 4 Liana abundance was positively correlated with pioneer tree abundance and diversity while negatively correlated with non-pioneer tree abundance and diversity. Thus, lianas appear to inhibit non-pioneer tree survival while indirectly enhancing that of pioneer trees. 5 Lianas are abundant in many types of tropical and temperate forests and a successional pathway involving liana-dominated, stalled gaps may therefore be frequent and widespread.

The abundance and diversity of lianas were examined along a tropical forest chronosequence at the Barro Colorado Nature Monument, Panama. Lianas ≥0.5 cm diameter were sampled along transects in two replicated stands in secondary (20, 40, 70 and 100 y after abandonment) and old-growth (>500 y) forests. Ordination of stands based on relative abundance, but not presence-absence, showed a significant separation of stands by age. Lianas were significantly more abundant and diverse (Fisher's α) in younger forests (20 and 40 y) than in older forests (70 and 100 y, and old-growth). The decline in liana abundance with stand age was offset by increased mean basal area per individual, resulting in a relatively constant total basal area and estimated biomass across stand age. The proportions of tendril climbers decreased and stem twiners increased over stand age. Decline in liana abundance and changes in liana composition may be related to changes in support and light availability. Although lianas are recognized as playing an important role in the early secondary sucession of many tropical forests, these results have shown that their important contribution to total basal area and biomass can continue as the forest matures, even as the numbers of established lianas declines.

Including lianas rooted within 2 m of the tree increased this percentage to 78%. Using both aboveand below-ground measures may provide a better estimate of liana competition than either measure alone. 6. Liana infestation is increasing on BCI. Lianas were present in the crowns of 73.6% of canopy trees (d.b.h. ‡20 cm). Liana canopy infestation was 57% higher than in 1980 and 65% higher than in 1967, which is consistent with reported increases in liana abundance, biomass, and leaf and flower production. 7. Synthesis. We used one of the largest studies ever conducted on lianas to confirm the negative effects of lianas on tree growth and survival over 10 years. Liana infestation of trees was widespread, dynamic and increasing on BCI.