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

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

Stink Bugs (Hemiptera: Heteroptera: Pentatomidae) of Minnesota: An Annotated Checklist and New State Records

Pentatomidae have been relatively poorly documented in Minnesota. Based on literature and museum records, an annotated checklist of the Pentatomidae of Minnesota was created. State-level and county-level records for Minnesota and the distribution of each species in North America are provided. Fifty-one species of Pentatomidae (12 Asopinae, 37 Pentatominae, and 2 Podopinae) are recorded for Minnesota. Of this total, 15 species are newly recorded for the state. Knowledge of the fauna of Pentatomidae in Minnesota will be important for providing baseline data for monitoring of potential shifts in the fauna resulting from the invasions of exotic Pentatomidae. Furthermore, a list of native Pentatomidae will be necessary for monitoring non-target impacts, if classical biological control is implemented for management of exotic Pentatomidae

Big Blue Bump and Transient Active Regions in Seyfert Galaxies

An important feature of the EUV spectrum (known as the Big Blue Bump, hereafter BBB) in Seyfert Galaxies is the narrow range in its cutoff energy E0 from source to source, even though the luminosity changes by 4 orders of magnitude. Here we show that if the BBB is due to accretion disk emission, then in order to account for this ``universality'' in the value of E0, the emission mechanism is probably optically thin bremsstrahlung. In addition, we demonstrate that the two-phase model with active regions localized on the surface of the cold disk is consistent with this constraint if the active regions are very compact and are highly transient, i.e., they evolve faster than one dynamical time scale.Comment: Accepted for publication in ApJL, uses aas2pp4.st

Error bounds on block Gauss Seidel solutions of coupled\ud multiphysics problems

Mathematical models in many fields often consist of coupled sub–models, each of which describe a different physical process. For many applications, the quantity of interest from these models may be written as a linear functional of the solution to the governing equations. Mature numerical solution techniques for the individual sub–models often exist. Rather than derive a numerical solution technique for the full coupled model, it is therefore natural to investigate whether these techniques may be used by coupling in a block Gauss–Seidel fashion. In this study, we derive two a posteriori bounds for such linear functionals. These bounds may be used on each Gauss–Seidel iteration to estimate the error in the linear functional computed using the single physics solvers, without actually solving the full, coupled problem. We demonstrate the use of the bound first by using a model problem from linear algebra, and then a linear ordinary differential equation example. We then investigate the effectiveness of the bound using a non–linear coupled fluid–temperature problem. One of the bounds derived is very sharp for most linear functionals considered, allowing us to predict very accurately when to terminate our block Gauss–Seidel iteration.\ud \ud Copyright c 2000 John Wiley & Sons, Ltd

Efficient algorithms for tensor scaling, quantum marginals and moment polytopes

We present a polynomial time algorithm to approximately scale tensors of any format to arbitrary prescribed marginals (whenever possible). This unifies and generalizes a sequence of past works on matrix, operator and tensor scaling. Our algorithm provides an efficient weak membership oracle for the associated moment polytopes, an important family of implicitly-defined convex polytopes with exponentially many facets and a wide range of applications. These include the entanglement polytopes from quantum information theory (in particular, we obtain an efficient solution to the notorious one-body quantum marginal problem) and the Kronecker polytopes from representation theory (which capture the asymptotic support of Kronecker coefficients). Our algorithm can be applied to succinct descriptions of the input tensor whenever the marginals can be efficiently computed, as in the important case of matrix product states or tensor-train decompositions, widely used in computational physics and numerical mathematics. We strengthen and generalize the alternating minimization approach of previous papers by introducing the theory of highest weight vectors from representation theory into the numerical optimization framework. We show that highest weight vectors are natural potential functions for scaling algorithms and prove new bounds on their evaluations to obtain polynomial-time convergence. Our techniques are general and we believe that they will be instrumental to obtain efficient algorithms for moment polytopes beyond the ones consider here, and more broadly, for other optimization problems possessing natural symmetries