Seedling establishment has long been believed to be rare on alpine tundra because of predicted life history trade-offs, the clonality of alpine species, and the harshness of the alpine climate. Contrary to the idea that seedlings are rare on alpine tundra, a 4-yr demographic study of seedlings at Niwot Ridge, Colorado, USA, found seedlings at high densities, particularly in wetter plant communities. Higher germination densities were associated with higher soil moistures both across communities and across time. Mortality of seedlings was highest in the first year and decreased in subsequent years. Species' abundances differed between seedling and adult populations. Many forbs that lacked vegetative reproduction were significantly more abundant among seedling populations, and many monocots and clonal forbs were more abundant among adult populations. In a comparison with published demographic rates, seedling recruitment and mortality rates of Niwot Ridge species fell above or within rates for a wide range of perennial species. Therefore, germination and seedling establishment stages are no more limiting to sexual reproduction in alpine plants than in other perennial plants.

Seed dormancy plays an important role in germination ecology and seed plant evolution. Morphological seed dormancy is caused by an underdeveloped embryo that must mature prior to germination. It has been suggested that the presence of an underdeveloped embryo is plesiomorphic among seed plants and that parallel directional change in embryo morphology has occurred separately in gymnosperms and in angiosperms. We test these hypotheses using original data on embryo morphology of key basal taxa, a published dataset, and the generalized least squares (GLS) method of ancestral character state reconstruction. Reconstructions for embryo to seed ratio (E:S) using family means for 179 families showed that E:S has increased between the ancestral angiosperm and almost all extant angiosperm taxa. Species in the rosid clade have particularly large embryos relative to the angiosperm ancestor. Results for the gymnosperms show a similar but smaller increase. There were no statistically significant differences in E:S between basal taxa and any derived group due to extremely large standard errors produced by GLS models. However, differences between reconstructed values for the angiosperm ancestor and more highly nested nodes are large and these results are robust to topological and branch-length manipulations. Our analysis supports the idea that the underdeveloped embryo is primitive among seed plants and that there has been a directional change in E:S within both angiosperms and gymnosperms. Our analysis suggests that dormancy enforced by an underdeveloped embryo is plesiomorphic among angiosperms and that nondormancy and other dormancy types probably evolved within the angiosperms. The shift in E:S was likely a heterochronic change, and has important implications for the life history of seed plants.

Seedling establishment is central to population maintenance for nonclonal plant species. Plants with low recruitment rates are expected to have high survival rates, and life history theory indicates there should be a single curve for the trade-off between recruitment and mortality that applies to most or all plant species. Alpine perennials are thought to have extraordinarily low recruitment rates because of the harsh environment, but the importance of recruitment in the life history of these plants is unknown. Two alpine cushion plant species, Minuartia obtusiloba and Paronychia pulvinata, were used to (1) determine the role of recruitment in population maintenance and (2) determine whether the fecundity/mortality trade-off for these alpine plants falls on or off of the curve for other perennial plant species. Using size-based population projection matrices, we determined that the life history of Minuartia and Paronychia emphasizes recruitment less than that of any other nonclonal species in a literature survey. Estimated maximum life spans of these two species are 200 and 324 yr, respectively, and a regression with other perennial species from the literature indicated that the relationship between fecundity and mortality in these alpine species is consistent with the predicted trade-off curve for perennial species from other environments.

This report describes a land management modeling effort that analyzed potential impacts of proposed actions under an updated Bureau of Land Management Resource Management Plan that will guide management for 20 years on 4.6 million hectares in the Great Basin ecoregion of the United States. State-and-transition models that included vegetation data, fire histories, and many parameters (i.e., rates of succession, fire return intervals, outcomes of management actions, and invasion rates of native and nonnative invasive species) were developed through workshops with scientific experts and range management specialists. Alternative restoration scenarios included continuation of current management, full fire suppression, wildfire use in designated fire use zones, wildfire use in resilient vegetation types only, restoration with a tenfold budget increase, no restoration treatments, and no livestock grazing. Under all the scenarios, cover of vegetation states with native perennial understory declined and was replaced by tree-invaded and weed-dominated states. The greatest differences among alternative management scenarios resulted from the use of fire as a tool to maintain native understory. Among restoration scenarios, only the scenario assuming a tenfold budget increase had a more desirable outcome than the current management scenario. Removal of livestock alone had little effect on vegetation resilience. Rather, active restoration was required. The predictive power of the model was limited by current understanding of Great Basin vegetation dynamics and data needs including statistically valid monitoring of restoration treatments, invasiveness and invasibility, and fire histories. The authors suggest that such computer models can be useful tools for systematic analysis of potential impacts in land use planning. However, for a modeling effort to be productive, the management situation must be conducive to open communication among land management agencies and partner entities, including nonprofit organizations.

Alpine plants offer unique opportunities to study the processes and economics of nutrient storage. The short alpine growing season forces rapid completion of plant growth cycles, which in turn causes competition between vegetative and reproductive growth sinks during the early part of the growing season. Mobilization of stored nitrogen and carbon reserves facilitates competing sinks and permits successful completion of reproduction before the onset of winter stress. We discuss the theoretical framework for assessing the costs and benefits of nutrient storage in alpine plants in order to lay the foundation for interpretation of observations. A principal point that has emerged from past theoretical treatments is the distinction between reserve storage, defined as storage that occurs with a cost to growth, and resource accumulation, defined as storage that occurs when resource supply exceeds demand, and thus when there is no cost to growth. We then discuss two case studies, one already published and one not yet published, pertaining to the storage and utilization of nitrogen and carbon compounds in alpine plants from Niwot Ridge, Colorado. In the first case, we tested the hypothesis that the seasonal accumulation of amino acids in the rhizome of N-fertilized plants of Bistorta bistortoides provides an advantage to the plant by not imposing a cost to growth at the time of accumulation, but providing a benefit to growth when the accumulated N is remobilized. We show that, as predicted, there is no cost during N accumulation but, not as predicted, there is no benefit to future growth. In the presence of N accumulation, reliance on stored N for growth increases, but reliance on current-season, soil-derived N decreases; thus the utilization of available N in this species is a 'zero sum' process. Inherent meristematic constraints to growth cause negative feedback that limits the utilization of accumulated N and precludes long-term advantages to this form of storage. In the second case study, we discuss new results showing high concentrations of cyclic polyol (cyclitol) compounds in the leaves of many alpine species dominant in the dry fellfield habitat. In Artemisia scopulorum, cyclitols were induced as the growing season progressed, and reached highest concentrations during the dry, late-summer months. Leaf cyclitol concentrations were high in all four species of the Caryophyllaceae that we examined and appeared to be constitutive components of the leaf carbohydrate pool as concentrations were high through the entire growing season. We observed correlations among seedling abundance, seeding survivorship and the presence of high leaf cyclitol concentrations. We propose that the primary function of cyclitols in the leaves of alpine, fellfield herbs is to promote drought tolerance through osmotic protection, and enhance fitness by improving seedling survival. We considered the possibility that cyclitols also function as carbon storage compounds that are remobilized at the end of the growing season and used to support growth...