Factors limiting the potential range expansion of lodgepole pine in Interior Alaska.

Understanding the factors influencing species range limits is increasingly crucial in anticipating migrations due to human-caused climate change. In the boreal biome, ongoing climate change and the associated increases in the rate, size, and severity of disturbances may alter the distributions of boreal tree species. Notably, Interior Alaska lacks native pine, a biogeographical anomaly that carries implications for ecosystem structure and function. The current range of lodgepole pine (Pinus contorta var. latifolia) in the adjacent Yukon Territory may expand into Interior Alaska, particularly with human assistance. Evaluating the potential for pine expansion in Alaska requires testing constraints on range limits such as dispersal limitations, environmental tolerance limits, and positive or negative biotic interactions. In this study, we used field experiments with pine seeds and transplanted seedlings, complemented by model simulations, to assess the abiotic and biotic factors influencing lodgepole pine seedling establishment and growth after fire in Interior Alaska. We found that pine could successfully recruit, survive, grow, and reproduce across our broadly distributed network of experimental sites. Our results show that both mammalian herbivory and competition from native tree species are unlikely to constrain pine growth and that environmental conditions commonly found in Interior Alaska fall well within the tolerance limits for pine. If dispersal constraints are released, lodgepole pine could have a geographically expansive range in Alaska, and once established, its growth is sufficient to support pine-dominated stands. Given the impacts of lodgepole pine on ecosystem processes such as increases in timber production, carbon sequestration, landscape flammability, and reduced forage quality, natural or human-assisted migration of this species is likely to substantially alter responses of Alaskan forest ecosystems to climate change.

Absence of net long-term successional facilitation by alder in a boreal Alaska floodplain.

Long-term experiments provide a way to test presumed causes of successional or environmentally driven vegetation changes. Early-successional nitrogen (N)-fixing plants are widely thought to facilitate productivity and vegetation development on N-poor sites, thus accounting for observed vegetation patterns later in succession. We tested this facilitative impact on vegetation development in a 23-yr field experiment on an Interior Alaska (USA) floodplain. On three replicate early-successional silt bars, we planted late-successional white spruce (Picea glauca) seedlings in the presence and absence of planted seedlings of an early-successional N-fixing shrub, thinleaf alder (Alnus incana). Alder initially facilitated survivorship and growth of white spruce. Within six years, however, after canopy closure, alder negatively affected spruce survivorship and growth. Our three replicate sites followed different successional trajectories. One site was eliminated by erosion and supported no vegetation development during our study. The other two sites, which differed in site moisture, diverged in vegetation composition. Structural equation modeling (SEM) suggested that, in the drier of these two sites, alder inhibited spruce growth directly (presumably by competition) and indirectly through effects mediated by competition with other woody species. However, at the wetter site, alder had both positive and negative effects on spruce growth, with negative effects predominating. Snowshoe hares (Lepus americanus) in alder thickets further reduced height growth of spruce in the wetter site. We conclude that net effects of alder on white spruce, the late-successional dominant, were primarily inhibitory and indirect, with the mechanisms depending on initial site moisture. Our results highlight the importance of long-term research showing that small differences among initial replicate sites can cause divergence in successional trajectories, consistent with individualistic distributions of species and communities along environmental gradients. This divergence was detectable only decades later.

Seasonal uptake and allocation of phosphorus in Eriophorum vaginatum L measured by labelling with 32 P.

Seasonal measurements of phosphorus (P) pools in the tussock-forming sedge Eriophorum vaginatum L. indicated low annual P uptake, but pronounced fluxes of P to growing biomass from below-ground stores in spring, and a return in autumn. Additions of 32 P to tussocks in early spring and mid summer confirmed this deduction. Addition of 32 P caused a rapid incorporation of about 10% of the labelled P in the tillers within a few days. Subsequent uptake was negligible, presumably because the 32 P not immediately absorbed was fixed in the soil and unavailable to plants. This demonstrates that nutrient pulses were more important than steady-state mineralization in supporting P uptake by Eriophorum vaginatum. After the 32 P additions, roots initially retained 60% of the total 32 P taken up. During the following weeks, part of the 32 P was transported slowly to stem bases and leaf sheaths, the principal storage organs. A lower proportion was transported to the leaf blades. We suggest that storage and recirculation of nutrients are the main sources of the annual nutrient supply to growth of E. vaginatum under normal field conditions, where P fixation in the soil and low rate of P diffusion towards the roots limit P uptake from soil except during periods of pulsed release.

Effect of defoliation upon root growth, phosphate absorption and respiration in nutrient-limited tundra graminoids.

Moderate experimental defoliation stimulated root respiration and phosphate absorption in two tundra graminoids, Eriophorum vaginatum and Carex aquatilis, growing under nutrient-limited field conditions in northern Alaska. The increase in phosphate absorption rate following defoliation of Eriophorum was associated with a decrease in root phosphate and available carbohydrate contents per unit root length but a constant root nitrogen content. Only after four repeated defoliations did phosphate absorption rate decrease below control levels. We suggest that the stimulation of root respiration and phosphate absorption immediately following defoliation resulted from lowered root phosphorus status as nutrient reserves were reallocated to support shoot regrowth. Root growth was affected more severely by defoliation than was root activity. Two or more defoliations reduced root elongation, initiation and weight per unit length, but root mortality increased only after four defoliations. Carex aquatilis, a species with large belowground biomass, was less sensitive to defoliation than Eriophorum. Phosphate absorption rate increased only after four defoliations in this species, and root elongation, initiation and mortality were affected only by the most severe clipping regimes. Responses of plants to repeated defoliation over two growing seasons were consistent with results of short-term studies.

Significance of sequential leaf development for nutrient balance of the cotton sedge,Eriophorum vaginatum L.

The sedgeEriophorum vaginatum in an interior Alaskan muskeg site produced leaves sequentially at about 1.5-month intervals. Each leaf remained active for two growing seasons. Young leaves (even those initiated late in the season) always had high concentrations of N, P, K and Mg and were low in Ca. Stems had high concentrations of nutrients, sugar, amino acid N and soluble organic P in autumn and spring but low concentrations in summer. Growth of leaves in spring was strongly supported by translocation from storage. Leaves approached their maximum nutrient pool before nutrient uptake began in late spring, one month before maximum biomass. Retranslocation of nutrients from aging leaves could support nutrient input into new, actively growing leaves as a consequence of the sequential leaf development. For instance retranslocation from aging leaves accounted for more than 90 and 85% of P and N input to new leaves appearing in early summer and 100% to leaves that appeared later. Leaching losses were negligible. Half time for decay of standing dead litter was 10 years. We suggest that sequential leaf development paired with highly efficient remobilization of nutrients from senescing leaves enables plants to recycle nutrients within the shoot and minimize dependence upon soil nutrients. This may be an important mechanism enablingEriophorum vaginatum to dominate nutrient-poor sites. This may also explain why graminoids with sequential leaf production cooccur with evergreen shrubs and dominate over forbs and deciduous shrubs in nutrient-poor sites in the boreal forest (e.g., in bogs) and at the northern limit of the tundra zone.