A Mechanistic Explanation for Global Patterns of Liana Abundance and Distribution

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

The Distribution of Lianas and Their Change in Abundance in Temperate Forests Over the Past 45 Years

Lianas (woody vines) are an important and dynamic component of many forests throughout the world, and increases in CO2, mean winter temperature, and forest fragmentation may promote their growth and proliferation in temperate forests. In this study, we used a 45‐year data set to test the hypothesis that lianas have increased in abundance and basal area in the interiors of 14 deciduous temperate forests in Wisconsin (USA) since 1959. We also censused woody plants along a gradient from the forest edge to the interior in seven of these forests to test the hypothesis that the abundance of lianas declines significantly with increasing distance from the forest edge. We found that lianas did not increase in abundance within the interiors of temperate forests in Wisconsin over the last 45 years. However, relative and absolute liana abundance decreased sharply with increasing distance from forest edges. Our findings suggest that forest fragmentation, not climate change, may be increasing the abundance of lianas in northern deciduous temperate forests, and that lianas may further increase in abundance if the severity of forest fragmentation intensifies

Disturbance and Clonal Reproduction Determine Liana Distribution and Maintain Liana Diversity in a Tropical Forest

Negative density dependence (NDD) and habitat specialization have received strong empirical support as mechanisms that explain tree species diversity maintenance and distribution in tropical forests. In contrast, disturbance appears to play only a minor role. Previous studies have rarely examined the relative strengths of these diversity maintenance mechanisms concurrently, and few studies have included plant groups other than trees. Here we used a large, spatially explicit data set from Barro Colorado Island, Panama (BCI) to test whether liana and tree species distribution patterns are most consistent with NDD, habitat specialization, or disturbance. We found compelling evidence that trees responded to habitat specialization and NDD; however, only disturbance explained the distribution of the majority of liana species and maintained liana diversity. Lianas appear to respond to disturbance with high vegetative (clonal) reproduction, and liana species\u27 ability to produce clonal stems following disturbance results in a clumped spatial distribution. Thus, clonal reproduction following disturbance explains local liana spatial distribution and diversity maintenance on BCI, whereas negative density dependence and habitat specialization, two prominent mechanisms contributing to tree species diversity and distribution, do not

Treefall Gaps and the Maintenance of Species Diversity in a Tropical Forest

The maintenance of species diversity by treefall gaps is a long‐standing paradigm in forest ecology. Gaps are presumed to provide an environment in which tree species of differing competitive abilities partition heterogeneous resources. The empirical evidence to support this paradigm, however, remains scarce, and some recent studies even suggest that gaps do not maintain the diversity of shade‐tolerant species. Although there is evidence that gaps maintain the diversity of pioneer trees, most of this evidence comes from studies that did not make comparisons between gaps and intact forest sites (controls). Further, nearly all studies on the maintenance of diversity by gaps have ignored lianas, an important component of both old‐world and neotropical forests. We tested the hypothesis that treefall gaps maintain shade‐tolerant tree, pioneer tree, and liana species diversity in an old‐growth forest on Barro Colorado Island (BCI), Panama. We compared the density and species richness of these guilds between paired gap and non‐gap sites on both a per‐area and a per‐individual (per capita) basis. We found no difference in shade‐tolerant tree density and species richness between the gap and non‐gap sites. Both pioneer tree and liana density and species richness, however, were significantly higher in the gap than in the non‐gap sites on both a per‐area and a per‐individual basis. These results suggest that gaps maintain liana species diversity and that this effect is not merely a consequence of increased density. Furthermore, our data confirm the long‐held belief that gaps maintain pioneer tree species diversity. Because lianas and pioneer trees combined account for ∼43% of the woody plant species on BCI, and in other forests, our results are likely to be broadly applicable and suggest that gaps play a strong role in the maintenance of woody species diversity

CP-Violating Yukawa Couplings in the Skyrme Model and the Neutron Electric Dipole Moment

We argue that the large-\Nc behaviour of the Yukawa couplings in the Skyrme model involves issues more subtle than the vanishing of linear fluctuations needed for classical stability of the skyrmion. The chiral fluctuations about the skyrmion must be quantized in order to reach a conclusion. An improved quantization procedure allows us to confront this question directly. The pion-nucleon coupling constants \gcup (CP conserving) and \bgcup (CP violating) are calculated in the large-\Nc, three-flavour Skyrme model by direct evaluation of the leading matrix elements appearing in the LSZ reduction formula. We find that \gcup \sim \Nc^{{3 \over 2}}, but that, at most, \bgcup \sim m^2_\pi \Nc^{-\shalf}. These results show that the leading contribution to the neutron electric dipole moment in large-\Nc Skyrme model is the direct one (\Dn \sim \Nc m^2_\pi), rather than the pion loop contribution.Comment: 12 pages, Latex with no macros, BRX-TH-33

Have we forgotten the forest because of the trees?

Recently, Brokaw and Busing argued that there is limited evidence for niche partitioning of tree species within forest gaps1. Consequently, gaps appear to play a relatively minor role in the maintenance of tree species diversity in forests via traditional resource partitioning. This conclusion is strongly supported by the existing empirical evidence, particularly for shade-tolerant tree species. Most studies of gaps, however, have failed to take into account plant groups other than trees2-4. Gaps may be a necessary habitat for the persistence of a large proportion of the vascular plant species other than shade-tolerant trees; specifically, pioneer trees, lianas, herbs, shrubs, and herbaceous vines2-8. For example, in a study on Barro Colorado Island (BCI), Panama, gaps had higher liana and pioneer tree diversity on both a per area and per stem basis (thereby removing the effect of density) compared to the surrounding forest2,3. These two plant groups alone account for approximately 43% of the woody species in this tropical forest2. There is also evidence that many forest herbs are gap dependent5,6. The role of gaps in the maintenance of shrubs is less clear, although there is some evidence that gaps promote shrub growth and reproduction7,8. Overall, when the major vascular plant groups are considered, as much as 65% of the flora of BCI may be gap dependent (Table 1). The specific mechanism that leads to the higher diversity of these groups in gaps remains unknown. Nonetheless, because these vascular plant groups represent a majority of the plant species in tropical forests worldwide4,9, gaps may often play a strong role in the maintenance of species diversity. Brokaw and Busing also argued that gaps might maintain diversity via the density effect10; Specifically, that gaps will have a higher diversity of trees solely because they have a higher density of trees compared to the surrounding forest. Tree density in gaps, however, declines (thins) with age, and thus the density effect could maintain diversity in the mature forest primarily in two ways. First, if individuals in gaps reach reproductive age prior to thinning then they could potentially colonize new gaps. Data are lacking, however, on whether trees reach reproductive age sooner (i.e., smaller size or age class) in gaps than in non-gap sites. Second, there must be niche partitioning. Without niche partitioning, thinning of individuals occurs randomly, and the initial increase in diversity would be merely a transitory result of the short-term increase in plant density2,11. Consequently, given the scanty evidence for niche partitioning and accelerated reproduction in gaps, the evidence for the density effect as a viable mechanism to explain the maintenance of diversity in forests is equivocal at best. We argue that papers sounding the death knell for the role of gaps in the maintenance of diversity in forests (e.g., Ref. 11) may be premature. The focus of most previous research on the ability of tree species to partition resources in gaps may have caused us to overlook the importance of gaps for many other groups of vascular plants (Table 1). Future research is necessary to quantify further the proportion of species in these and other groups (e.g., epiphytes) that require gaps for persistence in the community

Lianas Suppress Seedling Growth and Survival of 14 Tree Species in a Panamanian Tropical Forest

Lianas are a common plant growth form in tropical forests, where they compete intensely with trees, decreasing tree recruitment, growth, and survival. If the detrimental effects of lianas vary significantly with tree species identity, as is often assumed, then lianas may influence tree species diversity and community composition. Furthermore, recent studies have shown that liana abundance and biomass are increasing relative to trees in neotropical forests, which will likely magnify the detrimental effects of lianas and may ultimately alter tree species diversity, relative abundances, and community composition. Few studies, however, have tested the responses of multiple tree species to the presence of lianas in robust, well‐replicated experiments. We tested the hypotheses that lianas reduce tree seedling growth and survival, and that the effect of lianas varies with tree species identity. We used a large‐scale liana removal experiment in Central Panama in which we planted 14 replicate seedlings of 14 different tree species that varied in shade tolerance in each of 16 80 × 80 m plots (eight liana‐removal and eight unmanipulated controls; 3136 total seedlings). Over a nearly two‐yr period, we found that tree seedlings survived 75% more, grew 300% taller, and had twice the aboveground biomass in liana‐removal plots than seedlings in control plots, consistent with strong competition between lianas and tree seedlings. There were no significant differences in the response of tree species to liana competition (i.e., there was no species by treatment interaction), indicating that lianas had a similar negative effect on all 14 tree species. Furthermore, the effect of lianas did not vary with tree species shade tolerance classification, suggesting that the liana effect was not solely based on light. Based on these findings, recently observed increases in liana abundance in neotropical forests will substantially reduce tree regeneration, but will not significantly alter tropical tree species diversity, relative abundance, or community composition

Living Close to Your Neighbors: The Importance of Both Competition and Facilitation in Plant Communities

Recent work has demonstrated that competition and facilitation likely operate jointly in plant communities, but teasing out the relative role of each has proven difficult. Here we address how competition and facilitation vary with seasonal fluctuations in environmental conditions, and how the effects of these fluctuations change with plant ontogeny. We planted three sizes of pine seedlings (Pinus strobus) into an herbaceous diversity experiment and measured pine growth every two weeks for two growing seasons. Both competition and facilitation occurred at different times of year between pines and their neighbors. Facilitation was important for the smallest pines when environmental conditions were severe. This effect decreased as pines got larger. Competition was stronger than facilitation overall and outweighed facilitative effects at annual time scales. Our data suggest that both competition and the counter‐directional effects of facilitation may be more common and more intense than previously considered