Stochastic simulation of clonal growth in the tall goldenrod,Solidago altissima

Summary As clonal plants grow they move through space. The movement patterns that result can be complex and difficult to interpret without the aid of models. We developed a stochastic simulation model of clonal growth in the tall goldenrod, Solidago altissima. Our model was calibrated with field data on the clonal expansion of both seedlings and established clones, and model assumptions were verified by statistical analyses.When simulations were based on empirical distributions with long rhizome lengths, there was greater dispersal, less leaf overlap, and less spatial aggregation than when simulations were based on distributions with comparatively short rhizome lengths. For the field data that we utilized, variation in rhizome lengths had a greater effect than variation for either branching angles or rhizome initiation points (see text). We also found that observed patterns of clonal growth in S. altissima did not cause the formation of fairy rings. However, simulations with an artificial distribution of branching angles demonstrate that fairy rings can result solely from a plant's clonal morphology.Stochastic simulation models that incorporated variation in rhizome lengths, branching angles, and rhizome initiation points produced greater dispersal and less leaf overlap than deterministic models. Thus, variation for clonal growth parameters may increase the efficiency of substrate exploration by increasing the area covered and by decreasing the potential for intraclonal competition. We also demonstrated that ramet displacements were slightly, but consistently lower in stochastic simulation models than in random-walk models. This difference was due to the incorporation of details on rhizome bud initiation into stochastic simulation models, but not random-walk models. We discuss the advantages and disadvantages of deterministic, stochastic simulation, and random-walk models of clonal growth.     

Neighborhood predictors of plant performance

Summary We developed models of inter-individual interference to predict the fecundity of individuals in populations of the annual plant species, Arabidopsis thaliana. An individual plant is modeled as having a neighborhood which is a circular area of fixed radius with the plant at its center. Other plants which share the circle with the focal plant are termed neighbors of the focal plant. We developed an index of neighborhood interference which is the independent variable in a non-linear regression model that predicts individual plant fecundity. We present methods of exploratory data analysis that are useful in determining a best neighborhood radius, defined as that radius which minimizes residual sum of squares, and in deciding on the functional form of the interference index. In developing the interference index for Arabidopsis, we focus on aspects of the spatial distribution of neighbors: their number, distance and angular dispersion.We found that a best (or optimal) neighborhood radius can be resolved, which provides the best predictor of plant performance. Fecundity predictors based on adult neighbors were noticeably better than those based on neighbors at the seedling stage. Rosettes of Arabidopsis may change location during development (they fall over) and the new fallen positions do provide some improvement in the predictor. Taking into account distance to neighbors within the neighborhood provided only negligible improvement in the model. Finally, the incorporation of angular dispersion in the crowding index produced a considerably better fit. The fecundity predictor that included number of neighbors and angular dispersion in the crowding index explained about 70% of the variation in individual seed set.     

The lepidopteran mitochondrial control region: structure and evolution

For several species of lepidoptera, most of the approximately 350-bp mitochondrial control-region sequences were determined. Six of these species are in one genus, Jalmenus; are closely related; and are believed to have undergone recent rapid speciation. Recent speciation was supported by the observation of low interspecific sequence divergence. Thus, no useful phylogeny could be constructed for the genus. Despite a surprising conservation of control-region length, there was little conservation of primary sequences either among the three lepidopteran genera or between lepidoptera and Drosophila. Analysis of secondary structure indicated only one possible feature in common--inferred stem loops with higher-than-random folding energies-- although the positions of the structures in different species were unrelated to regions of primary sequence similarity. We suggest that the conserved, short length of control regions is related to the observed lack of heteroplasmy in lepidopteran mitochondrial genomes. In addition, determination of flanking sequences for one Jalmenus species indicated (i) only weak support for the available model of insect 12S rRNA structure and (ii) that tRNA translocation is a frequent event in the evolution of insect mitochondrial genomes.      

Predicting shifts in the functional composition of tropical forests under increased drought and CO2 from trade‐offs among plant hydraulic traits

Tropical forest responses are an important feedback on global change, but changes in forest composition with projected increases in CO₂ and drought are highly uncertain. Here we determine shifts in the most competitive plant hydraulic strategy (the evolutionary stable strategy or ESS) from changes in CO₂ and drought frequency and intensity. Hydraulic strategies were defined along a spectrum from drought avoidance to tolerance by physiology traits. Drought impacted competition more than CO₂, with elevated CO₂ reducing but not reversing drought‐induced shifts in the ESS towards more tolerant strategies. Trait plasticity and/or adaptation intensified these shifts by increasing the competitive ability of the drought tolerant relative to the avoidant strategies. These findings predict losses of drought avoidant evergreens from tropical forests under global change, and point to the importance of changes in precipitation during the dry season and constraints on plasticity and adaptation in xylem traits to forest responses.     

Modeling and analysis of stochastic invasion processes

In this paper we derive spatially explicit equations to describe a stochastic invasion process. Parents are assumed to produce a random number of offspring which then disperse according to a spatial redistribution kernel. Equations for population moments, such as expected density and covariance averaged over an ensemble of identical stochastic processes, take the form of deterministic integro-difference equations. These equations describe the spatial spread of population moments as the invasion progresses. We use the second order moments to analyse two basic properties of the invasion. The first property is permanence of form in the correlation structure of the wave. Analysis of the asymptotic form of the invasion wave shows that either (i) the covariance in the leading edge of the wave of invasion asymptotically achieves a permanence of form with a characteristic structure described by an unchanging spatial correlation function, or (ii) the leading edge of the wave has no asymptotic permanence of form with the length scales of spatial correlations continually increasing over time. Which of these two outcomes pertains is governed by a single statistic, φ which depends upon the shape of the dispersal kernel and the net reproductive number. The second property of the invasion is its patchy structure. Patchiness, defined in terms of spatial correlations on separate short (within patch) and long (between patch) spatial scales, is linked to the dispersal kernel. Analysis shows how a leptokurtic dispersal kernel gives rise to patchiness in spread of a population. Received: 11 August 1997 / Revised version: 22 September 1998 / Published online: 4 October 2000     

Intraspecific variation and species coexistence

We use a two-species model of plant competition to explore the effect of intraspecific variation on community dynamics. The competitive ability ("performance") of each individual is assigned by an independent random draw from a species-specific probability distribution. If the density of individuals competing for open space is high (e.g., because fecundity is high), species with high maximum (or large variance in) performance are favored, while if density is low, species with high typical (e.g., mean) performance are favored. If there is an interspecific mean-variance performance trade-off, stable coexistence can occur across a limited range of intermediate densities, but the stabilizing effect of this trade-off appears to be weak. In the absence of this trade-off, one species is superior. In this case, intraspecific variation can blur interspecific differences (i.e., shift the dynamics toward what would be expected in the neutral case), but the strength of this effect diminishes as competitor density increases. If density is sufficiently high, the inferior species is driven to extinction just as rapidly as in the case where there is no overlap in performance between species. Intraspecific variation can facilitate coexistence, but this may be relatively unimportant in maintaining diversity in most real communities.     

Local diversity in heterogeneous landscapes: quantitative assessment with a height-structured forest metacommunity model

“Mass effects,” in which “sink populations” of locally inferior competitors are maintained by dispersal from “source populations” elsewhere in the landscape, are thought to play an important role in maintaining plant diversity. However, due to the complexity of most quasi-realistic forest models, there is little theoretical understanding of the strength of mass effects in forests. Here, we develop a metacommunity version of a mathematically and computationally tractable height-structured forest model, the Perfect Plasticity Approximation, to quantify the strength of mass effects (i.e., the degree of mixing of locally dominant and subordinate species) in heterogeneous landscapes comprising different patch types (e.g., soil types). For realistic levels of inter-patch dispersal, mass effects are weak at equilibrium (i.e., in the absence of disturbance), even in some cases where differences in growth, mortality, and fecundity rates between locally dominant and subordinate species are too small to be reliably detected from field data. However, patch-scale transient dynamics are slow following catastrophic disturbance (in which post-disturbance initial abundances are determined exclusively by immigration) so that at any given time, subordinate species are present in appreciable numbers in most patches. Less severe disturbance regimes, in which some seeds or individuals survive the disturbance, should result in faster transient dynamics (i.e., faster approach to the low-diversity equilibrium). Our results suggest that in order for mass effects to play an important role in tree coexistence, niche differences must be strong enough to prevent neutral drift, yet too weak to be reliably detected from field data.     

Differential declines in Alaskan boreal forest vitality related to climate and competition

Rapid warming and changes in water availability at high latitudes alter resource abundance, tree competition, and disturbance regimes. While these changes are expected to disrupt the functioning of boreal forests, their ultimate implications for forest composition are uncertain. In particular, recent site‐level studies of the Alaskan boreal forest have reported both increases and decreases in productivity over the past few decades. Here, we test the idea that variations in Alaskan forest growth and mortality rates are contingent on species composition. Using forest inventory measurements and climate data from plots located throughout interior and south‐central Alaska, we show significant growth and mortality responses associated with competition, midsummer vapor pressure deficit, and increased growing season length. The governing climate and competition processes differed substantially across species. Surprisingly, the most dramatic climate response occurred in the drought tolerant angiosperm species, trembling aspen, and linked high midsummer vapor pressure deficits to decreased growth and increased insect‐related mortality. Given that species composition in the Alaskan and western Canadian boreal forests is projected to shift toward early‐successional angiosperm species due to fire regime, these results underscore the potential for a reduction in boreal productivity stemming from increases in midsummer evaporative demand.     

The role of succession in the evolution of flammability

Fire-prone ecosystems contain plants that are both fire-adapted and flammable. It has been hypothesized that these plants were under selection to become more flammable, but it is unclear whether this could be adaptive for an individual plant. We propose arrested succession as a robust mechanism that supports the evolution of flammability in surface fire ecosystems without the need to invoke group selection or additional fitness benefits. We used the natural history of lodgepole pine (Pinus ponderosa) forests, longleaf pine (Pinus palustris) forests, and tall grass prairies to create a general mathematical model of surface fire ecosystems and solved for the evolutionarily stable strategy (ESS) level of flammability. In our model, fires always kill understory plants and only sometimes kill overstory plants. Thus, more flammable plants suffer increased mortality due to fires, but also more frequently arrest succession by clearing their understory of late successional competitors. Increased flammability was selected for when the probability of an overstory plant dying from an individual fire was below a maximum threshold and the rate of succession relative to fires was above a minimum threshold. Future studies can test our model predictions and help resolve whether or not plants have been selected to be more flammable.