North American tree migration paced by climate in the West, lagging in the East.

Tree fecundity and recruitment have not yet been quantified at scales needed to anticipate biogeographic shifts in response to climate change. By separating their responses, this study shows coherence across species and communities, offering the strongest support to date that migration is in progress with regional limitations on rates. The southeastern continent emerges as a fecundity hotspot, but it is situated south of population centers where high seed production could contribute to poleward population spread. By contrast, seedling success is highest in the West and North, serving to partially offset limited seed production near poleward frontiers. The evidence of fecundity and recruitment control on tree migration can inform conservation planning for the expected long-term disequilibrium between climate and forest distribution.

Globally, tree fecundity exceeds productivity gradients.

Lack of tree fecundity data across climatic gradients precludes the analysis of how seed supply contributes to global variation in forest regeneration and biotic interactions responsible for biodiversity. A global synthesis of raw seedproduction data shows a 250-fold increase in seed abundance from cold-dry to warm-wet climates, driven primarily by a 100-fold increase in seed production for a given tree size. The modest (threefold) increase in forest productivity across the same climate gradient cannot explain the magnitudes of these trends. The increase in seeds per tree can arise from adaptive evolution driven by intense species interactions or from the direct effects of a warm, moist climate on tree fecundity. Either way, the massive differences in seed supply ramify through food webs potentially explaining a disproportionate role for species interactions in the wet tropics.

Watershed-scale liming reveals the short- and long-term effects of pH on the forest soil microbiome and carbon cycling.

Soil microbial community composition routinely correlates with pH, reflecting both direct pH effects on microbial physiology and long-term biogeochemical feedbacks. We used two watershed-scale liming experiments to identify short- (2 years) and long-term (25 years) changes in the structure and function of bacterial and fungal communities in organic horizons (Oe and Oa ) of acid forest soils. Liming increased soil pH, extractable calcium, and soil carbon stocks, reduced biomass-specific respiration, and caused major changes in the soil microbiome in the short and long term. More taxa responded to liming in the short term (70%) than in the long term (30%), with most showing consistent directional responses at both sites. The ratio of change in relative abundance between limed and reference sites was twofold higher at the long than the short-term site, indicating that the effects of liming grew over time. Liming impacts were most pronounced in fungi, as steep declines of dominant ectomycorrhizal fungi (Cenococcum and Russula) occurred at both sites. Liming favoured neutrophilic bacteria over acidophilic populations according to estimated environmental pH optima. Collectively, these results demonstrate that a liming-induced change of one pH unit has an immediate and persistent effect on the structure and function of microbial communities in acid forest soils. The corresponding suppression of respiration indicates that anthropogenic alterations of soil pH, as driven by acid deposition or liming, can affect forest floor C stocks due to pH-driven shifts in community structure.

Microbial community shifts correspond with suppression of decomposition 25 years after liming of acidic forest soils.

Microbial community structure and function regularly covary with soil pH, yet effects of these interactions on soil carbon are rarely tested experimentally within natural ecosystems. We investigated the enduring (25 year) impacts of liming on microbial community structure and decomposition at an acidic northern hardwood forest, where experimental liming increased pH one unit and surprisingly doubled the organic carbon stocks of the forest floor. We show that this increase in carbon storage corresponded with restructuring of the bacterial and fungal communities that drive decomposition. In the Oe horizon, liming reduced the activities of five extracellular enzymes that mediate decomposition, while the Oa horizon showed an especially large (64%) reduction in the activity of a sixth, peroxidase, which is an oxidative enzyme central to lignocellulose degradation. Decreased enzyme activities corresponded with loss of microbial taxa important for lignocellulose decay, including large reductions in the dominant ectomycorrhizal genera Russula and Cenococcum, saprotrophic and wood decaying fungi, and Actinobacteria (Thermomonosporaceae). These results demonstrate the importance of pH as a dominant regulator of microbial community structure and illustrate how changes to this structure can produce large, otherwise unexpected increases in carbon storage in forest soils.

Survival and energy use of Ixodes scapularis nymphs throughout their overwintering period.

The blacklegged tick (Ixodes scapularis) spends up to 10 months in the soil between feeding as larvae and questing for hosts as nymphs the following year. We tracked the survival and energy use of 4320 engorged larvae evenly divided across 288 microcosms under field conditions from September to July on sites with high (>12 nymphs/150 m2) and low (<1.2 nymphs/150 m2) densities of naturally questing I. scapularis in New York State. Subsets of microcosms were destructively sampled periodically during this period to determine tick survivorship and physiological age. Across all sites tick mortality was low during the winter and increased in the spring and early summer, coincident with increasing energy use. Neither energy use nor mortality differed significantly between sites with high vs low natural tick density, but we did observe a significant positive relationship between soil organic matter content and the survival of I. scapularis during the spring. Our results suggest that the off-host mortality and energy use of I. scapularis nymphs is relatively low in the winter and increases significantly in the spring and early summer.

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.

The unseen invaders: introduced earthworms as drivers of change in plant communities in North American forests (a meta-analysis).

Globally, biological invasions can have strong impacts on biodiversity as well as ecosystem functioning. While less conspicuous than introduced aboveground organisms, introduced belowground organisms may have similarly strong effects. Here, we synthesize for the first time the impacts of introduced earthworms on plant diversity and community composition in North American forests. We conducted a meta-analysis using a total of 645 observations to quantify mean effect sizes of associations between introduced earthworm communities and plant diversity, cover of plant functional groups, and cover of native and non-native plants. We found that plant diversity significantly declined with increasing richness of introduced earthworm ecological groups. While plant species richness or evenness did not change with earthworm invasion, our results indicate clear changes in plant community composition: cover of graminoids and non-native plant species significantly increased, and cover of native plant species (of all functional groups) tended to decrease, with increasing earthworm biomass. Overall, these findings support the hypothesis that introduced earthworms facilitate particular plant species adapted to the abiotic conditions of earthworm-invaded forests. Further, our study provides evidence that introduced earthworms are associated with declines in plant diversity in North American forests. Changing plant functional composition in these forests may have long-lasting effects on ecosystem functioning.

Decreased water flowing from a forest amended with calcium silicate.

Acid deposition during the 20th century caused widespread depletion of available soil calcium (Ca) throughout much of the industrialized world. To better understand how forest ecosystems respond to changes in a component of acidification stress, an 11.8-ha watershed was amended with wollastonite, a calcium silicate mineral, to restore available soil Ca to preindustrial levels through natural weathering. An unexpected outcome of the Ca amendment was a change in watershed hydrology; annual evapotranspiration increased by 25%, 18%, and 19%, respectively, for the 3 y following treatment before returning to pretreatment levels. During this period, the watershed retained Ca from the wollastonite, indicating a watershed-scale fertilization effect on transpiration. That response is unique in being a measured manipulation of watershed runoff attributable to fertilization, a response of similar magnitude to effects of deforestation. Our results suggest that past and future changes in available soil Ca concentrations have important and previously unrecognized implications for the water cycle.

Redefining fine roots improves understanding of below-ground contributions to terrestrial biosphere processes.

Fine roots acquire essential soil resources and mediate biogeochemical cycling in terrestrial ecosystems. Estimates of carbon and nutrient allocation to build and maintain these structures remain uncertain because of the challenges of consistently measuring and interpreting fine-root systems. Traditionally, fine roots have been defined as all roots ≤ 2 mm in diameter, yet it is now recognized that this approach fails to capture the diversity of form and function observed among fine-root orders. Here, we demonstrate how order-based and functional classification frameworks improve our understanding of dynamic root processes in ecosystems dominated by perennial plants. In these frameworks, fine roots are either separated into individual root orders or functionally defined into a shorter-lived absorptive pool and a longer-lived transport fine-root pool. Using these frameworks, we estimate that fine-root production and turnover represent 22% of terrestrial net primary production globally - a c. 30% reduction from previous estimates assuming a single fine-root pool. Future work developing tools to rapidly differentiate functional fine-root classes, explicit incorporation of mycorrhizal fungi into fine-root studies, and wider adoption of a two-pool approach to model fine roots provide opportunities to better understand below-ground processes in the terrestrial biosphere.

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 = 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 basdd on single-element limitation, butfollows from the correlation between soil N and soil P. The expected single-element relationships were evident only in the 0 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 0 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.

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

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.

Simulating effects of changing climate and CO(2) emissions on soil carbon pools at the Hubbard Brook experimental forest.

Carbon (C) sequestration in forest biomass and soils may help decrease regional C footprints and mitigate future climate change. The efficacy of these practices must be verified by monitoring and by approved calculation methods (i.e., models) to be credible in C markets. Two widely used soil organic matter models - CENTURY and RothC - were used to project changes in SOC pools after clear-cutting disturbance, as well as under a range of future climate and atmospheric carbon dioxide (CO(2) ) scenarios. Data from the temperate, predominantly deciduous Hubbard Brook Experimental Forest (HBEF) in New Hampshire, USA, were used to parameterize and validate the models. Clear-cutting simulations demonstrated that both models can effectively simulate soil C dynamics in the northern hardwood forest when adequately parameterized. The minimum postharvest SOC predicted by RothC occurred in postharvest year 14 and was within 1.5% of the observed minimum, which occurred in year 8. CENTURY predicted the postharvest minimum SOC to occur in year 45, at a value 6.9% greater than the observed minimum; the slow response of both models to disturbance suggests that they may overestimate the time required to reach new steady-state conditions. Four climate change scenarios were used to simulate future changes in SOC pools. Climate-change simulations predicted increases in SOC by as much as 7% at the end of this century, partially offsetting future CO(2) emissions. This sequestration was the product of enhanced forest productivity, and associated litter input to the soil, due to increased temperature, precipitation and CO(2) . The simulations also suggested that considerable losses of SOC (8-30%) could occur if forest vegetation at HBEF does not respond to changes in climate and CO(2) levels. Therefore, the source/sink behavior of temperate forest soils likely depends on the degree to which forest growth is stimulated by new climate and CO(2) conditions.

Winter climate change affects growing-season soil microbial biomass and activity in northern hardwood forests.

Understanding the responses of terrestrial ecosystems to global change remains a major challenge of ecological research. We exploited a natural elevation gradient in a northern hardwood forest to determine how reductions in snow accumulation, expected with climate change, directly affect dynamics of soil winter frost, and indirectly soil microbial biomass and activity during the growing season. Soils from lower elevation plots, which accumulated less snow and experienced more soil temperature variability during the winter (and likely more freeze/thaw events), had less extractable inorganic nitrogen (N), lower rates of microbial N production via potential net N mineralization and nitrification, and higher potential microbial respiration during the growing season. Potential nitrate production rates during the growing season were particularly sensitive to changes in winter snow pack accumulation and winter soil temperature variability, especially in spring. Effects of elevation and winter conditions on N transformation rates differed from those on potential microbial respiration, suggesting that N-related processes might respond differently to winter climate change in northern hardwood forests than C-related processes.

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 13C and 15N, 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 13C released from litter was recovered in soil with no significant differences among plots. Total recovery of added 13C (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 15N than 13C 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.

Root morphology and architecture respond to N addition in Pinus tabuliformis, west China.

Belowground dynamics of terrestrial ecosystems are responding to global increases in anthropogenic N deposition with important consequences for productivity and ecosystem health. We compared root characteristics across five root orders in Pinus tabuliformis plantations treated for 3 years to a gradient of N addition (0-15 g m(-2) year(-1)). In reference plots, the roots of P. tabuliformis were finer and with higher specific root length than reported for other pine species, suggesting severe N limitation. Addition of N resulted in slightly reduced fine root biomass and significant changes in root morphology, responses that were associated primarily with first and second order roots. In particular, root number, cumulative root length, individual root length, and specific root length all declined with increasing N addition for first and second order roots, with most of the responses elicited at <9 g m(-2) year(-1) N addition. These responses (1) support the concept of ephemeral root modules consisting of first and second orders and (2) are consistent with a change in functional demand from uptake to transport with increasing soil resource availability. Traditionally, fine roots have been identified by a somewhat arbitrary diameter cut-off (e.g., 1 or 2 mm); as an index of fine root function, diameter would fail to reveal most of the functional response.

Masting is uncommon in trees that depend on mutualist dispersers in the context of global climate and fertility gradients.

The benefits of masting (volatile, quasi-synchronous seed production at lagged intervals) include satiation of seed predators, but these benefits come with a cost to mutualist pollen and seed dispersers. If the evolution of masting represents a balance between these benefits and costs, we expect mast avoidance in species that are heavily reliant on mutualist dispersers. These effects play out in the context of variable climate and site fertility among species that vary widely in nutrient demand. Meta-analyses of published data have focused on variation at the population scale, thus omitting periodicity within trees and synchronicity between trees. From raw data on 12 million tree-years worldwide, we quantified three components of masting that have not previously been analysed together: (i) volatility, defined as the frequency-weighted year-to-year variation; (ii) periodicity, representing the lag between high-seed years; and (iii) synchronicity, indicating the tree-to-tree correlation. Results show that mast avoidance (low volatility and low synchronicity) by species dependent on mutualist dispersers explains more variation than any other effect. Nutrient-demanding species have low volatility, and species that are most common on nutrient-rich and warm/wet sites exhibit short periods. The prevalence of masting in cold/dry sites coincides with climatic conditions where dependence on vertebrate dispersers is less common than in the wet tropics. Mutualist dispersers neutralize the benefits of masting for predator satiation, further balancing the effects of climate, site fertility and nutrient demands.

Foliar nutrient concentrations of six northern hardwood species responded to nitrogen and phosphorus fertilization but did not predict tree growth.

Foliar chemistry can be useful for diagnosing soil nutrient availability and plant nutrient limitation. In northern hardwood forests, foliar responses to nitrogen (N) addition have been more often studied than phosphorus (P) addition, and the interactive effects of N and P addition have rarely been described. In the White Mountains of central New Hampshire, plots in ten forest stands of three age classes across three sites were treated annually beginning in 2011 with 30 kg N ha-1 y-1 or 10 kg P ha-1 y-1 or both or neither-a full factorial design. Green leaves of American beech (Fagus grandifolia Ehrh.), pin cherry (Prunus pensylvanica L.f.), red maple (Acer rubrum L.), sugar maple (A. saccharum Marsh.), white birch (Betula papyrifera Marsh.), and yellow birch (B. alleghaniensis Britton) were sampled pre-treatment and 4-6 years post-treatment in two young stands (last cut between 1988-1990), four mid-aged stands (last cut between 1971-1985) and four mature stands (last cut between 1883-1910). In a factorial analysis of species, stand age class, and nutrient addition, foliar N was 12% higher with N addition (p < 0.001) and foliar P was 45% higher with P addition (p < 0.001). Notably, P addition reduced foliar N concentration by 3% (p = 0.05), and N addition reduced foliar P concentration by 7% (p = 0.002). When both nutrients were added together, foliar P was lower than predicted by the main effects of N and P additions (p = 0.08 for N × P interaction), presumably because addition of N allowed greater use of P for growth. Foliar nutrients did not differ consistently with stand age class (p ≥ 0.11), but tree species differed (p ≤ 0.01), with the pioneer species pin cherry having the highest foliar nutrient concentrations and the greatest responses to nutrient addition. Foliar calcium (Ca) and magnesium (Mg) concentrations, on average, were 10% (p < 0.001) and 5% lower (p = 0.01), respectively, with N addition, but were not affected by P addition (p = 0.35 for Ca and p = 0.93 for Mg). Additions of N and P did not affect foliar potassium (K) concentrations (p = 0.58 for N addition and p = 0.88 for P addition). Pre-treatment foliar N:P ratios were high enough to suggest P limitation, but trees receiving N (p = 0.01), not P (p = 0.64), had higher radial growth rates from 2011 to 2015. The growth response of trees to N or P addition was not explained by pre-treatment foliar N, P, N:P, Ca, Mg, or K.

Limits to reproduction and seed size-number trade-offs that shape forest dominance and future recovery.

The relationships that control seed production in trees are fundamental to understanding the evolution of forest species and their capacity to recover from increasing losses to drought, fire, and harvest. A synthesis of fecundity data from 714 species worldwide allowed us to examine hypotheses that are central to quantifying reproduction, a foundation for assessing fitness in forest trees. Four major findings emerged. First, seed production is not constrained by a strict trade-off between seed size and numbers. Instead, seed numbers vary over ten orders of magnitude, with species that invest in large seeds producing more seeds than expected from the 1:1 trade-off. Second, gymnosperms have lower seed production than angiosperms, potentially due to their extra investments in protective woody cones. Third, nutrient-demanding species, indicated by high foliar phosphorus concentrations, have low seed production. Finally, sensitivity of individual species to soil fertility varies widely, limiting the response of community seed production to fertility gradients. In combination, these findings can inform models of forest response that need to incorporate reproductive potential.

Nitrate production and availability in residential soils.

The rapid increase in residential land area in the United States has raised concern about water pollution associated with nitrogen fertilizers. Nitrate (NO3-) is the form of reactive N that is most susceptible to leaching and runoff; thus, a more thorough understanding of nitrification and NO3(-) availability is needed if we are to accurately predict the consequences of residential expansion for water quality. In particular, there have been few assessments of how the land use history, housing density, and age of residential soils influence NO3(-) pools and fluxes, especially at depth. In this study, we used 1 m deep soil cores to evaluate potential net nitrification and mineralization, microbial respiration and biomass, and soil NO3(-) and NH4+ pools in 32 residential home lawns that differed by previous land use and age, but had similar soil types. These were compared to eight forested reference sites with similar soils. Our results suggest that a change to residential land use has increased pools and production of reactive N, which has clear implications for water quality in the region. However, the results contradict the common assumption that NO3(-) production and availability is dramatically higher in residential soils than in forests in general. While net nitrification (128.6 +/- 15.5 mg m(-2) d(-1) vs. 4.7 +/- 2.3 mg m(-2) d(-1); mean +/- SE) and exchangeable NO3(-) (3.8 +/- 0.5 g/m2 vs. 0.7 +/- 0.3 g/m2) were significantly higher in residential soils than in forest soils in this study, these measures of NO3(-) production and availability were still notably low, comparable to deciduous forest stands in other studies. A second unexpected result was that current homeowner management practices were not predictive of NO3(-) availability or production. This may reflect the transient availability of inorganic N after fertilizer application. Higher housing density and a history of agricultural land use were predictors of greater NO3(-) availability in residential soils. If these factors are good predictors across a wider range of sites, they may be useful indicators of NO3(-) availability and leaching and runoff potential at the landscape scale.