Experimental biogeography: the role of environmental gradients in high geographic diversity in Cape Proteaceae.

One of the fundamental dimensions of biodiversity is the rate of species turnover across geographic distance. The Cape Floristic Region of South Africa has exceptionally high geographic species turnover, much of which is associated with groups of closely related species with mostly or completely non-overlapping distributions. A basic unresolved question about biodiversity in this global hotspot is the relative importance of ecological gradients in generating and maintaining high geographic turnover in the region. We used reciprocal transplant experiments to test the extent to which abiotic environmental factors may limit the distributions of a group of closely related species in the genus Protea (Proteaceae), and thus elevate species turnover in this diverse, iconic family. We tested whether these species have a "home site advantage" in demographic rates (germination, growth, mortality), and also parameterized stage-structured demographic models for the species. Two of the three native species were predicted to have a demographic advantage at their home sites. The models also predicted, however, that species could maintain positive population growth rates at sites beyond their current distribution limits. Thus the experiment suggests that abiotic limitation under current environmental conditions does not fully explain the observed distribution limits or resulting biogeographic pattern. One potentially important mechanism is dispersal limitation, which is consistent with estimates based on genetic data and mechanistic dispersal models, though other mechanisms including competition may also play a role.

Hierarchical models facilitate spatial analysis of large data sets: a case study on invasive plant species in the northeastern United States.

Many critical ecological issues require the analysis of large spatial point data sets - for example, modelling species distributions, abundance and spread from survey data. But modelling spatial relationships, especially in large point data sets, presents major computational challenges. We use a novel Bayesian hierarchical statistical approach, 'spatial predictive process' modelling, to predict the distribution of a major invasive plant species, Celastrus orbiculatus, in the northeastern USA. The model runs orders of magnitude faster than traditional geostatistical models on a large data set of c. 4000 points, and performs better than generalized linear models, generalized additive models and geographically weighted regression in cross-validation. We also use this approach to model simultaneously the distributions of a set of four major invasive species in a spatially explicit multivariate model. This multispecies analysis demonstrates that some pairs of species exhibit negative residual spatial covariation, suggesting potential competitive interaction or divergent responses to unmeasured factors.

Distribution and dynamics of two ferns: Dennstaedtia punctilobula (Dennstaedtiaceae) and Thelypteris noveboracensis (Thelypteridaceae) in a Northeast mixed hardwoods-hemlock forest.

Dennstaedtia punctilobula and Thelypteris noveboracensis are two native species that often arrest forest succession and reduce understory diversity. As part of a project to examine the feedback between forest understory and canopy dynamics, we studied the patterns of distribution and dynamics of these two fern species in an oak-transition hardwoods-hemlock forest. Dennstaedtia was least abundant under shade-tolerant tree species and most abundant in small (1-2 trees) canopy gaps, but did not show any distinct patterns across the sampled moisture regime. The light response was verified using light manipulation experiments and examination of plant size-abundance patterns across light environments. Thelypteris tended to be most prevalent under maple canopies and appeared to be more sensitive to soil moisture regime being restricted to more mesic sites than Dennstaedtia. Seasonal and year-to-year changes in abundance of established clones of both fern species were small, suggesting that once established, both species can maintain a strong hold on a site. Further work on the niche requirements of the two species is warranted, but any event that maintains or promotes canopy openness (tree death by disease or windthrow, forest harvesting, or the elimination of a shrub layer by browsing) will promote persistence of Dennstaedtia.

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

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.

The ecological importance of insect frass: allelopathy in eucalypts.

Our previously published experiments on allelopathic effects of insect frass in Eucalyptus communities (Silander et al. 1983) have been criticized on the grounds that our estimates of annual frass production were exscessive (Ohmart 1985). However, we spanned the entire array of estimates of frass fall available from eucalypt communities, and we demonstrated allelopathic effects at even the lowest levels suggested by Ohmart. We suggest that average values of frass fall per hectare are irrelevant because they ignore both large scale variation among sites in time and space, and small scale variation in patchiness of frass accumulation. At the ecological scales relevant to germinating seeds, frass concentrations in local pockets may be considerably higher than the averages calculated for the entire ground surface.

Chemical interference among plants mediated by grazing insects.

The experiments on Eucalyptus trees reported here demonstrate that allelopathic effects can be mediated by insects grazing on foliage. We show that the allelochemical nature of insect frass suppresses germination, growth and survival of herb layer species, that plant species vary in their tolerance of this material, and that as a result, the structure and composition of associated, herbaceous understory plant communities are markedly affected by frass fall.

Microevolution and Clone Structure in Spartina patens.

Analysis of the clone structures within a population of Spartina patens reveals considerable adaptive genetic divergence among adjacent dune, swale, and marsh sites. The dune subpopulation includes a small number of frequently encountered, spatially isolated clones that follow microtopography and have high reproductive output and colonizing potential. The marsh subpopulation consists of a large number of infrequent interdigitating clones with high vegetative biomass and competitive success. The swale subpopulation is generally intermediate for these traits.