Phosphorus limitation of aboveground production in northern hardwood forests

Forest productivity on glacially derived soils with weatherable phosphorus (P) is expected to be limited by nitrogen (N), according to theories of long-term ecosystem development. However, recent studies and model simulations based on resource optimization theory indicate that productivity can be co-limited by N and P. We conducted a full factorial N × P fertilization experiment in 13 northern hardwood forest stands of three age classes in central New Hampshire, USA, to test the hypothesis that forest productivity is co-limited by N and P. We also asked whether the response of productivity to N and P addition differs among species and whether differential species responses contribute to community-level co-limitation. Plots in each stand were fertilized with 30 kg N·ha−1·yr−1, 10 kg P·ha−1·yr−1, N + P, or neither nutrient (control) for four growing seasons. The productivity response to treatments was assessed using per-tree annual relative basal area increment (RBAI) as an index of growth. RBAI responded significantly to P (P = 0.02) but not to N (P = 0.73). However, evidence for P limitation was not uniform among stands. RBAI responded to P fertilization in mid-age (P = 0.02) and mature (P = 0.07) stands, each taken as a group, but was greatest in N-fertilized plots of two stands in these age classes, and there was no significant effect of P in the young stands. Both white birch (Betula papyrifera Marsh.) and beech (Fagus grandifolia Ehrh.) responded significantly to P; no species responded significantly to N. We did not find evidence for N and P co-limitation of tree growth. The response to N + P did not differ from that to P alone, and there was no significant N × P interaction (P = 0.68). Our P limitation results support neither the N limitation prediction of ecosystem theory nor the N and P co-limitation prediction of resource optimization theory, but could be a consequence of long-term anthropogenic N deposition in these forests. Inconsistencies in response to P suggest that successional status and variation in site conditions influence patterns of nutrient limitation and recycling across the northern hardwood forest landscape

Soil nitrogen affects phosphorus recycling: foliar resorption and plant–soil feedbacks in a northern hardwood forest

Previous studies have attempted to link foliar resorption of nitrogen and phosphorus to their respective availabilities in soil, with mixed results. Based on resource optimization theory, we hypothesized that the foliar resorption of one element could be driven by the availability of another element. We tested various measures of soil N and P as predictors of N and P resorption in six tree species in 18 plots across six stands at the Bartlett Experimental Forest, New Hampshire, USA. Phosphorus resorption efficiency (P , 0.01) and proficiency (P ¼ 0.01) increased with soil N content to 30 cm depth, suggesting that trees conserve P based on the availability of soil N. Phosphorus resorption also increased with soil P content, which is difficult to explain based on single-element limitation, but follows from the correlation between soil N and soil P. The expected single-element relationships were evident only in the O horizon: P resorption was high where resin-available P was low in the Oe (P , 0.01 for efficiency, P , 0.001 for proficiency) and N resorption was high where potential N mineralization in the Oa was low (P , 0.01 for efficiency and 0.11 for proficiency). Since leaf litter is a principal source of N and P to the O horizon, low nutrient availability there could be a result rather than a cause of high resorption. The striking effect of soil N content on foliar P resorption is the first evidence of multiple-element control on nutrient resorption to be reported from an unmanipulated ecosystem

Community and population dynamics of spruce-fir forests on Whiteface Mountain, New York: recent trends, 1985-2000

We remeasured two sets of permanent plots in old-growth, spruce–fir forests on Whiteface Mountain to quantify ongoing vegetation dynamics at sites impacted by spruce decline. One set of plots was a stratified random sample of the vegetation in a subalpine watershed (Baldwin site). The other was selected to represent forest conditions in a high-elevation subset of the spruce–fir forest (Esther site). Between 1987 and 1997, there was a significant increase in aboveground tree biomass at Baldwin with the majority of the increment due to the growth of canopy-sized trees. This growth occurred with little change in either species composition or size structure. The annual mortality rate of 1.2%·year–1 for canopy-sized red spruce (Picea rubens Sarg.) in Baldwin almost matched the recruitment rate of 1.4 stems/ha per year. In addition, the relative growth rate of spruce was significantly faster than associated species. In contrast, spruce trees in Esther died at a rate of the 3.6%·year–1 (1985–1995), and survivors grew more slowly than other species. The most obvious community-level trend at Esther (1985–2000) was an increase in overall tree density with most of this increase due to ingrowth of small trees. The demography of the spruce population at Baldwin suggests that the decline is over for at least this population

Recovery from disturbance requires resynchronization of ecosystem nutrient cycles

Nitrogen (N) and phosphorus (P) are tightly cycled in most terrestrial ecosystems, with plant uptake more than 10 times higher than the rate of supply from deposition and weathering. This near-total dependence on recycled nutrients and the stoichiometric constraints on resource use by plants and microbes mean that the two cycles have to be synchronized such that the ratio of N:P in plant uptake, litterfall, and net mineralization are nearly the same. Disturbance can disrupt this synchronization if there is a disproportionate loss of one nutrient relative to the other. We model the resynchronization of N and P cycles following harvest of a northern hardwood forest. In our simulations, nutrient loss in the harvest is small relative to postharvest losses. The low N:P ratio of harvest residue results in a preferential release of P and retention of N. The P release is in excess of plant requirements and P is lost from the active ecosystem cycle through secondary mineral formation and leaching early in succession. Because external P inputs are small, the resynchronization of the N and P cycles later in succession is achieved by a commensurate loss of N. Through succession, the ecosystem undergoes alternating periods of N limitation, then P limitation, and eventually co-limitation as the two cycles resynchronize. However, our simulations indicate that the overall rate and extent of recovery is limited by P unless a mechanism exists either to prevent the P loss early in succession (e.g., P sequestration not stoichiometrically constrained by N) or to increase the P supply to the ecosystem later in succession (e.g., biologically enhanced weathering). Our model provides a heuristic perspective from which to assess the resynchronization among tightly cycled nutrients and the effect of that resynchronization on recovery of ecosystems from disturbance

Earthworm effects on the incorporation of litter C and N into soil organic matter in a sugar maple forest

To examine the mechanisms of earthworm effects on forest soil C and N, we double-labeled leaf litter with C-13 and N-15, applied it to sugar maple forest plots with and without earthworms, and traced isotopes into soil pools. The experimental design included forest plots with different earthworm community composition (dominated by Lumbricus terrestris or L. rubellus). Soil carbon pools were 37% lower in earthworm-invaded plots largely because of the elimination of the forest floor horizons, and mineral soil C:N was lower in earthworm plots despite the mixing of high C:N organic matter into soil by earthworms. Litter disappearance over the first winter-spring was highest in the L. terrestris (T) plots, but during the warm season, rapid loss of litter was observed in both L. rubellus (R) and T plots. After two years, 22.0% +/- 5.4% of C-13 released from litter was recovered in soil with no significant differences among plots. Total recovery of added C-13 (decaying litter plus soil) was much higher in no-worm (NW) plots (61-68%) than in R and T plots (20-29%) as much of the litter remained in the former whereas it had disappeared in the latter. Much higher percentage recovery of N-15 than C-13 was observed, with significantly lower values for T than R and NW plots. Higher overwinter earthworm activity in T plots contributed to lower soil N recovery. In earthworm-invaded plots isotope enrichment was highest in macroaggregates and microaggregates whereas in NW plots silt plus clay fractions were most enriched. The net effect of litter mixing and priming of recalcitrant soil organic matter (SOM), stabilization of SOM in soil aggregates, and alteration of the soil microbial community by earthworm activity results in loss of SOM and lowering of the C:N ratio. We suggest that earthworm stoichiometry plays a fundamental role in regulating C and N dynamics of forest SOM

The promise and peril of intensive-site-based ecological research: insights from the Hubbard Brook ecosystem study

Abstract. Ecological research is increasingly concentrated at particular locations or sites. This trend reflects a variety of advantages of intensive, site-based research, but also raises important questions about the nature of such spatially delimited research: how well does site based research represent broader areas, and does it constrain scientific discovery?We provide an overview of these issues with a particular focus on one prominent intensive research site: the Hubbard Brook Experimental Forest (HBEF), New Hampshire, USA. Among the key features of intensive sites are: long-term, archived data sets that provide a context for new discoveries and the elucidation of ecological mechanisms; the capacity to constrain inputs and parameters, and to validate models of complex ecological processes; and the intellectual cross-fertilization among disciplines in ecological and environmental sciences. The feasibility of scaling up ecological observations from intensive sites depends upon both the phenomenon of interest and the characteristics of the site. An evaluation of deviation metrics for the HBEF illustrates that, in some respects, including sensitivity and recovery of streams and trees from acid deposition, this site is representative of the Northern Forest region, of which HBEF is a part. However, the mountainous terrain and lack of significant agricultural legacy make the HBEF among the least disturbed sites in the Northern Forest region. Its relatively cool, wet climate contributes to high stream flow compared to other sites. These similarities and differences between the HBEF and the region can profoundly influence ecological patterns and processes and potentially limit the generality of observations at this and other intensive sites. Indeed, the difficulty of scaling up may be greatest for ecological phenomena that are sensitive to historical disturbance and that exhibit the greatest spatiotemporal variation, such as denitrification in soils and the dynamics of bird communities. Our research shows that end member sites for some processes often provide important insights into the behavior of inherently heterogeneous ecological processes. In the current era of rapid environmental and biological change, key ecological responses at intensive sites will reflect both specific local drivers and regional trends

Dynamics of Oxidized and Reduced Iron in a Northern Hardwood Forest

Iron (Fe) is ubiquitous in forest ecosystems and its cycle is thought to influence the development of soil, particularly Spodosols (podsolization), and the biogeochemistry of macronutrients such as carbon (C), nitrogen (N), and phosphorus (P), as well as many trace metals. The cycle of Fe in northern hardwood forests remains poorly understood. To address some of these uncertainties, we constructed a biogeochemical budget of Fe for a small catchment at the Hubbard Brook Experimental Forest in the White Mountains of New Hampshire, USA. Horizonal, temporal, and elevational patterns of concentrations and fluxes of oxidized and reduced Fe species were assessed in leaf litter, soil, soil solution, and stream water. The chemistry of dissolved Fe was evaluated in the context of its relationship with dissolved organic carbon, pH, and dissolved oxygen. Soil solution fluxes of Fe were highest in the organic (Oa, 52.5 mol ha−1 year−1) horizon and decreased with depth in the mineral (Bh, 50.5 mol ha−1 year−1, and Bs, 19.7 mol ha−1 year−1) horizons, consistent with podsolization theories predicting immobilization of Fe following downward transport to mineral soils. The export of Fe in stream water (1.8 mol ha−1 year−1) was lower than precipitation input (3.5 mol ha−1 year−1). The low stream flux indicates most Fe in drainage waters was immobilized in the soil and retained in the watershed. The portion of total Fe as Fe(II) was ~10–60% in soil solutions, seemingly high for soils that are considered to be well-drained, oxidizing environments. Organic complexes likely stabilized Fe(II) in solution under oxidizing conditions that would otherwise promote considerably higher Fe(III)-to-Fe(II) ratios. Our study indicates that there are organic matter-derived sources of dissolved Fe(II) as well as substantial mobilization of Fe(II), possibly the result of the reduction of Fe-bearing soil minerals