Nitrogen cycling in a northern hardwood forest: Do species matter?

To investigate the influence of individual tree species on nitrogen (N) cycling in forests, we measured key characteristics of the N cycle in small single-species plots of five dominant tree species in the Catskill Mountains of New York State. The species studied were sugar maple (Acer saccharum), American beech (Fagus grandifolia), yellow birch (Betula alleghaniensis), eastern hemlock (Tsuga canadensis), and red oak (Quercus rubra). The five species varied markedly in N cycling characteristics. For example, hemlock plots consistently showed characteristics associated with "slow" N cycling, including low foliar and litter N, high soil C:N, low extractable N pools, low rates of potential net N mineralization and nitrification and low NO ₃ ^– amounts trapped in ion-exchange resin bags buried in the mineral soil. Sugar maple plots had the lowest soil C:N, and the highest levels of soil characteristics associated with NO ₃ ^– production and loss (nitrification, extractable NO ₃ ^– , and resin bag NO ₃ ^– ). In contrast, red oak plots had near-average net mineralization rates and soil C:N ratios, but very low values of the variables associated with NO ₃ ^– production and loss. Correlations between soil N transformations and litter concentrations of N, lignin, lignin:N ratio, or phenolic constituents were generally weak. The inverse correlation between net nitrification rate and soil C:N that has been reported in the literature was present in this data set only if red oak plots were excluded from the analysis. This study indicates that tree species can exert a strong control on N cycling in forest ecosystems that appears to be mediated through the quality of soil organic matter, but that standard measures of litter quality cannot explain the mechanism of control.     

The relative uptake of Ca and Sr into tree foliage using a whole-watershed calcium addition

The use of strontium isotopes and ratios of alkaline earth elements (i.e., ⁸⁷Sr/⁸⁶Sr and Ca/Sr) to trace Ca sources to plants has become common in ecosystem studies. Here we examine the relative uptake of Ca and Sr in trees and subsequent accumulation in foliage. Using a whole-watershed Ca addition experiment at the Hubbard Brook Experimental Forest in N.H., we measured the uptake of Ca relative to Sr in foliage and roots of sugar maple (Acer saccharum), yellow birch (Betula alleghaniensis), American beech (Fagus grandifolia), and red spruce (Picea rubens). Vegetation was analyzed for Ca and Sr concentrations and the ⁸⁷Sr/⁸⁶Sr ratio. A comparison of the Ca/Sr ratio in the vegetation and the Ca/Sr ratio of the applied mineral allows for the calculation of a discrimination factor, which defines whether Ca and Sr are incorporated and allocated in the same ratio as that which is available. A discrimination factor greater than unity indicates preferential uptake of Ca over Sr; a factor less than unity reflects preferential uptake of Sr over Ca. We demonstrate that sugar maple (SM) and yellow birch (YB) have similar and small discrimination factors (1.14 ± 0.12,1σ and 1.16 ± 0.09,1σ) in foliage formation and discrimination factors of less than 1 in root formation (0.55–0.70). Uptake into beech suggests a larger discrimination factor (1.9 ± 1.2) in foliage but a similar root discrimination factor to SM and YB (0.66 ± 0.06,1σ). Incorporation into spruce foliage occurs at a much slower rate than in these other tree species and precludes evaluation of Ca and Sr discrimination in spruce foliage at this time. Understanding the degree to which Ca is fractionated from Sr in different species allows for refinement in the use of ⁸⁷Sr/⁸⁶Sr and Ca/Sr ratios to trace Ca sources to foliage. Methods from this study can be applied to natural environments in which various soil cation pools have different ⁸⁷Sr/⁸⁶Sr and Ca/Sr ratios. The results reported herein have implications for re-evaluating Ca sources and fluxes in forest ecosystems.     

Hydraulic conductivity of branch junctions in three temperate tree species

Summary Hydraulic conductivities were measured in branch junctions and in the proximal segments of Quercus velutina Lam., Acer saccharum Marsh., and Tsuga canadensis (L.) Carr. In all three species, the basal proximal segment was more conductive than the junction by a factor of 1.1–1.5. There was no consistent pattern for the distal proximal segments, where the conductivities were higher, lower, or the same as the junction. These data are discussed in terms of Martin H. Zimmermann's concept of plant segmentation. It is concluded that junction constrictions to water flow contribute less to plant segmentation than the variation in leaf specific conductivity in the crown of these species.     

Spatial patterns of tree seedling establishment and their relationship to environmental variables in a cold-temperate deciduous forest of eastern North America

Patterns of seedling recruitment may have persistent effects on population and community processes. Assuming seed availability is not limiting, the environmental sieve (i.e., the suite of factors influencing seed germination and seedling emergence and survival) determines how many seedlings establish and, most importantly, where they do so. In this study, we identify the spatial structure of some resources and abiotic conditions known to be significant for tree seedling emergence and survival and determine how these environmental factors influence the establishment of Fagus grandifolia, Acer saccharum, Fraxinus americana, and Ostrya virginiana in a deciduous forest of southern Québec (Canada). We expect an increase from Fagus, through Acer and Fraxinus, to Ostrya in the control of environmental variables on seedling emergence and survival, because of differences in the seed size of these species. Density of newly-emerged seedlings of all four species showed positive spatial autocorrelation at distances of up to ca. 10 m. Environmental variables were also structured at the same spatial scale, except for soil moisture. Acer seedling emergence pattern was positively correlated to photosynthetic photon flux density (PPFD), and the pattern of Fraxinus to soil N and moisture. Seedling survival was not spatially autocorrelated for any of the four species, although it was positively density-dependent in Acer and Fagus. In only Ostrya was seedling survival correlated (positively) to one of the environmental variables studied, i.e., PPFD. Overall, environmental variables were spatially less heterogeneous than seedling emergence and survival. Either seed availability was not saturating or factors not considered here, such as competition and predation (the intensity of which often varies with resources and/or abiotic conditions), modified the influence that the physical environment had on patterns of seedling establishment. Our prediction of a greater environmental control on seedling emergence and survival in small-seed species was not totally confirmed.     

Dispersal versus climate: Expansion of Fagus and Tsuga into the Upper Great Lakes region

Pollen records for American beech (Fagus grandifolia) and eastern hemlock (Tsuga canadensis) compiled from 50 sites in Michigan and Wisconsin, USA, show that both species entered the Upper Great Lakes region about 7000 yr B.P., reaching their western and southwestern boundaries between 2000 and 1000 yr B.P. Fagus advanced northward into lower Michigan as a continuous front, except where Lake Michigan posed a geographic barrier. Colonies were established on the far side of the lake after a 1000 year lag, implying that longdistance dispersal across a 100-km wide barrier can occur. The Fagus range may not have been in equilibrium with climate for one or two thousand years before this time, when seeds were dispersed across the lake to Wisconsin. Tsuga seeds may have been dispersed 150 km or more from Ontario to reach Upper Michigan. Scattered colonies were established 6000–7000 yr B.P. on either side of Lake Michigan, which did not pose a significant barrier to this wind-dispersed species, Tsuga spread rapidly over a large region prior to 5000 yr B.P. Subsequent expansion to the west occurred more slowly, and may reflect gradual climatic changes in northern Wisconsin during the second half of the Holocene. Tsuga's range may have been limited by dispersal, rather than climate, for an unknown length of time prior to 5000 yr B.P. During this period Tsuga was expanding its range rapidly. The study shows, however, that it is difficult to devise rigorous tests to distinguish between dispersal limitations and climate as factors limiting range limits in the past.This work has been supported by the U.S. National Science Foundation.This work has been supported by the U.S. National Science Foundation.     

Competitive effects of the invasive shrub,Lonicera maackii (Rupr.) Herder (Caprifoliaceae), on the growth and survival of native tree seedlings

Invasive plants are often associated with reduced cover of native plants, but rarely has competition between invasives and natives been assessed experimentally. The shrub Lonicera maackii, native to northeastern Asia, has invaded forests and old fields in numerous parts of eastern North America, and is associated with reduced tree seedling density in Ohio forests. A field experiment was conducted to test the effects of established L. maackii on the survival and growth of transplanted native tree species. The experiment examined above-ground competition (by removing L. maackii shoots) and below-ground competition (by trenching around transplanted seedlings). The effects of above-ground competition with L. maackii were generally more important than below-ground competition, though both were detected. Shoot treatment was the key determinant for the survival of all species except P. serotina, whereas trenching only enhanced survival for A. saccharum caged and P. serotina, and only in the shoot removal treatment. For the surviving seedlings, L. maackii shoot removal increased growth of A. saccharum seedlings protected with cages, but actually reduced the growth of unprotected Q. rubra and A. saccharum seedlings, indicating that L. maackii shoots confer some protection from deer browsing. Significant interactions between root and shoot treatment on Q. rubra growth parameters, specifically greatest growth in the shoot present & trenched treatment, is attributed to protection from deer browsing combined with release from below-ground competition. Despite this protective function of L. maackii shoots, the overall effect of this invasive shrub is increased mortality of native tree seedlings, suggesting it impacts the natural regeneration of secondary forests.     

Effects of European Earthworm Invasion on Soil Characteristics in Northern Hardwood Forests of Minnesota, USA

European earthworms are colonizing worm-free hardwood forests across North America. Leading edges of earthworm invasion in forests of northern Minnesota provide a rare opportunity to document changes in soil characteristics as earthworm invasions are occurring. Across leading edges of earthworm invasion in four northern hardwood stands, increasing total earthworm biomass was associated with rapid disappearance of the O horizon. Concurrently, the thickness, bulk density and total soil organic matter content of the A horizon increased, and it’s percent organic matter and fine root density decreased. Different earthworm species assemblages influenced the magnitude and type of change in these soil parameters. Soil N and P availability were lower in plots with high earthworm biomass compared to plots with low worm biomass. Decreases in soil nitrogen availability associated with high earthworm biomass were reflected in decreased foliar nitrogen content for Carex pensylvanica, Acer saccharum and Asarum canadense but increased foliar N for Athyrium felix-femina. Overall, high earthworm biomass resulted in increased foliar carbon to nitrogen ratios. The effects of earthworm species assemblages on forest soil properties are related to their feeding and burrowing habits in addition to effects related to total biomass. The potential for large ecosystem consequences following exotic earthworm invasion has only recently been recognized by forest ecologists. In the face of rapid change and multiple pressures on native forest ecosystems, the impacts of earthworm invasion on forest soil structure and function must be considered.     

Non-timber forest product harvesting in alien-dominated forests: effects of frond-harvest and rainfall on the demography of two native Hawaiian ferns

Non-timber forest products (NTFP) represent culturally and economically important resources for millions of people worldwide. Although many NTFP are harvested from disturbed habitats and therefore subject to multiple pressures, few quantitative studies have addressed this issue. Similarly few NTFP studies have assessed seasonal variation in demographic rates even though this can confound harvest effects. In Hawaiȁ8i, the wild-gathered ferns, Microlepia strigosa and Sphenomeris chinensis, represent highly important cultural resources but declining populations have led to conservation concerns. Both ferns are harvested from disturbed, alien-dominated forests and contemporary Hawaiian gathering practices often consist of harvest and concurrent weeding of alien invasive species. We assessed the effects of concurrent frond-harvest and alien species weeding on frond structure, density, and rates of production by comparing experimentally harvested vs. control plots, and documented relationships between frond demographic patterns and precipitation. Gathering practices had no impact on frond density of either species or on most other demographic parameters over the short term. Exceptions included a significant decrease in the density of the longest S. chinensis fronds and a significant decrease in M. strigosa frond production when fronds were gathered without alien weeding. However, seasonal and annual changes in frond density and production occurred across all plots of both species and were significantly correlated with precipitation. The relatively low harvest effects for both species are likely due to several factors including short frond longevity and the strict criteria used by gatherers to select harvestable fronds. The potential for sustainable harvest in the context of alien-dominated forests is discussed.