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

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

Connectivity Explains Local Ant Community Structure in A Neotropical Forest Canopy: A Large‐Scale Experimental Approach

Understanding how habitat structure and resource availability affect local species distributions is a key goal of community ecology. Where habitats occur as a mosaic, variation in connectivity among patches influences both local species richness and composition, and connectivity is a key conservation concern in fragmented landscapes. Similarly, availability of limiting resources frequently determines species coexistence or exclusion. For primarily cursorial arthropods like ants, gaps between neighboring trees are a significant barrier to movement through the forest canopy. Competition for limited resources such as nest sites also promotes antagonistic interactions. Lianas (woody vines) connect normally isolated neighboring tree crowns and often have hollow stems inhabited by ants. We used two large‐scale liana‐removal experiments to determine how connectivity and nest site availability provided by lianas affect arboreal ant species richness, species composition, and β‐diversity in a lowland tropical forest in Panama. Removing lianas from a tree crown reduced ant species richness up to 35%, and disproportionately affected species that require large foraging areas. Adding artificial connectivity to trees mitigated the effects of liana removal. Ant colonization of artificial nests was higher (73% occupied) in trees without lianas vs. trees with lianas (28% occupied). However, artificial nests typically were colonized by existing polydomous, resident ant species. As a result, nest addition did not affect ant community structure. Collectively, these results indicate that lianas are important to the maintenance of arboreal ant diversity specifically by providing connectivity among neighboring tree crowns. Anticipated increases in liana abundance in this forest could increase the local (tree‐level) species richness of arboreal ants, with a compositional bias toward elevating the density of broad‐ranging specialist predators

The impact of lianas on tree regeneration in tropical forest canopy gaps: evidence for an alternative pathway of gap‐phase regeneration

1 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 identified 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 different 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

Esophageal Involvement in Lymphoma

Peer Reviewedhttp://deepblue.lib.umich.edu/bitstream/2027.42/75337/1/j.1572-0241.1985.tb02134.x.pd

Effects of Lightning on Trees: A Predictive Model Based on in situ Electrical Resistivity

The effects of lightning on trees range from catastrophic death to the absence of observable damage. Such differences may be predictable among tree species, and more generally among plant life history strategies and growth forms. We used field‐collected electrical resistivity data in temperate and tropical forests to model how the distribution of power from a lightning discharge varies with tree size and identity, and with the presence of lianas. Estimated heating density (heat generated per volume of tree tissue) and maximum power (maximum rate of heating) from a standardized lightning discharge differed 300% among tree species. Tree size and morphology also were important; the heating density of a hypothetical 10 m tall Alseis blackiana was 49 times greater than for a 30 m tall conspecific, and 127 times greater than for a 30 m tall Dipteryx panamensis. Lianas may protect trees from lightning by conducting electric current; estimated heating and maximum power were reduced by 60% (±7.1%) for trees with one liana and by 87% (±4.0%) for trees with three lianas. This study provides the first quantitative mechanism describing how differences among trees can influence lightning–tree interactions, and how lianas can serve as natural lightning rods for trees

Lianas suppress tree regeneration and diversity in treefall gaps

Treefall gaps are hypothesized to maintain diversity by creating resource‐rich, heterogeneous habitats necessary for species coexistence. This hypothesis, however, is not supported empirically for shade‐tolerant trees, the dominant plant group in tropical forests. The failure of gaps to maintain shade‐tolerant trees remains puzzling, and the hypothesis implicated to date is dispersal limitation. In central Panama, we tested an alternative ‘biotic interference’ hypothesis: that competition between growth forms (lianas vs. trees) constrains shade‐tolerant tree recruitment, survival and diversity in gaps. We experimentally removed lianas from eight gaps and monitored them for 8 years, while also monitoring nine un‐manipulated control gaps. Removing lianas increased tree growth, recruitment and richness by 55, 46 and 65%, respectively. Lianas were particularly harmful to shade‐tolerant species, but not pioneers. Our findings demonstrate that competition between plant growth forms constrains diversity in a species‐rich tropical forest. Because lianas are abundant in many tropical systems, our findings may apply broadly