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.

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.

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.

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.

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.

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.

Biological Control of Hemlock Woolly Adelgid: Implications of Adult Emergence Patterns of Two Leucopis spp. (Diptera: Chamaemyiidae) and Laricobius nigrinus (Coleoptera: Derodontidae) Larval Drop.

The hemlock woolly adelgid (Hemiptera: Adelgidae Adelges tsugae Annand) poses a serious threat to hemlocks in eastern North America, and ongoing research is focused on the identification and development of biological controls to protect and manage hemlock resources. Three predators native to the Pacific Northwest of North America that have been the focus of much research are Leucopis argenticollis (Zetterstedt), Leucopis piniperda (Malloch) (Diptera: Chamaemyiidae), and Laricobius nigrinus (Fender) (Coleoptera: Derodontidae). This study addresses the knowledge gap of adult Leucopis spp. emergence patterns, with comparisons to the timing of larval La. nigrinus drop for pupation. Adult Leucopis spp. emergence was observed in the lab from field-collected, adelgid-infested foliage from Washington state in 2019 and 2020. Adult Leucopis spp. were collected daily as they emerged from foliage collections and identified to species using morphological features; a subset was validated using DNA barcoding. Accumulated heating degree days were calculated to compare a standardized emergence timing across collections made at different locations and temperature regimes. The abundance of the two Leucopis spp. and of the combined Leucopis spp. and La. nigrinus varied among sites and years, and no species was consistently more abundant than the other. Evaluations of seasonal emergence trends of the three species determine the predator complex behaves in a temporally stratified and predictable way. Emergence of adult Le. argenticollis was observed first, followed by La. nigrinus larval drop, with Le. piniperda emerging at the end of larval drop, and finally a second emergence of Le. argenticollis.

Continent-wide tree fecundity driven by indirect climate effects.

Indirect climate effects on tree fecundity that come through variation in size and growth (climate-condition interactions) are not currently part of models used to predict future forests. Trends in species abundances predicted from meta-analyses and species distribution models will be misleading if they depend on the conditions of individuals. Here we find from a synthesis of tree species in North America that climate-condition interactions dominate responses through two pathways, i) effects of growth that depend on climate, and ii) effects of climate that depend on tree size. Because tree fecundity first increases and then declines with size, climate change that stimulates growth promotes a shift of small trees to more fecund sizes, but the opposite can be true for large sizes. Change the depresses growth also affects fecundity. We find a biogeographic divide, with these interactions reducing fecundity in the West and increasing it in the East. Continental-scale responses of these forests are thus driven largely by indirect effects, recommending management for climate change that considers multiple demographic rates.

Simulation of the effects of forest harvesting under changing climate to inform long-term sustainable forest management using a biogeochemical model.

Process ecosystem models are useful tools to provide insight on complex, dynamic ecological systems, and their response to disturbances. The biogeochemical model PnET-BGC was modified and tested using field observations from an experimentally whole-tree harvested northern hardwood watershed (W5) at the Hubbard Brook Experimental Forest (HBEF), New Hampshire, USA. In this study, the confirmed model was used as a heuristic tool to investigate long-term changes in hydrology, biomass accumulation, and soil solution and stream water chemistry for three different watershed cutting intensities (40%, 60%, 80%) and three rotation lengths (30, 60, 90 years) under both constant (current climate) and changing (MIROC5-RCP4.5) future climate scenarios and atmospheric CO2 through the year 2200. For the no future cutting scenario, total ecosystem stored carbon (i.e., sum of aboveground biomass, woody debris and soil) reached a maximum value of 207 t C ha-1 under constant climate but increased to 452 t C ha-1 under changing climate in 2200 due to a CO2 fertilization effect. Harvesting of trees decreased total ecosystem stored carbon between 7 and 36% for constant climate and 7-60% under changing climate, respectively, with greater reductions for shorter logging rotation lengths and greater watershed cutting intensities. Harvesting under climate change resulted in noticeable losses of soil organic matter (12-56%) coinciding with loss of soil nutrients primarily due to higher rates of soil mineralization associated with increases in temperature, compared with constant climate conditions (3-22%). Cumulative stream leaching of nitrate under climate change (181-513 kg N ha-1) exceeded constant climate values (139-391 kg N ha-1) for the various cutting regimes. Under both climate conditions the model projected greater sensitivity to varying the length of cutting period than cutting intensities. Hypothetical model simulations highlight future challenges in maintaining long-term productivity of managed forests under changing climate due to a potential for a deterioration of soil fertility.

Mean annual temperature influences local fine root proliferation and arbuscular mycorrhizal colonization in a tropical wet forest.

Mean annual temperature (MAT) is an influential climate factor affecting the bioavailability of growth-limiting nutrients nitrogen (N) and phosphorus (P). In tropical montane wet forests, warmer MAT drives higher N bioavailability, while patterns of P availability are inconsistent across MAT. Two important nutrient acquisition strategies, fine root proliferation into bulk soil and root association with arbuscular mycorrhizal fungi, are dependent on C availability to the plant via primary production. The case study presented here tests whether variation in bulk soil N bioavailability across a tropical montane wet forest elevation gradient (5.2°C MAT range) influences (a) morphology fine root proliferation into soil patches with elevated N, P, and N+P relative to background soil and (b) arbuscular mycorrhizal fungal (AMF) colonization of fine roots in patches. We created a fully factorial fertilized root ingrowth core design (N, P, N+P, unfertilized control) representing soil patches with elevated N and P bioavailability relative to background bulk soil. Our results show that percent AMF colonization of roots increased with MAT (r 2 = .19, p = .004), but did not respond to fertilization treatments. Fine root length (FRL), a proxy for root foraging, increased with MAT in N+P-fertilized patches only (p = .02), while other fine root morphological parameters did not respond to the gradient or fertilized patches. We conclude that in N-rich, fine root elongation into areas with elevated N and P declines while AMF abundance increases with MAT. These results indicate a tradeoff between P acquisition strategies occurring with changing N bioavailability, which may be influenced by higher C availability with warmer MAT.

Experimental approach and initial forest response to a simulated ice storm experiment in a northern hardwood forest.

Ice storms are a type of extreme winter weather event common to north temperate and boreal forests worldwide. Recent climate modelling studies suggest that these storms may become more frequent and severe under a changing climate. Compared to other types of storm events, relatively little is known about the direct and indirect impacts of these storms on forests, as naturally occurring ice storms are inherently difficult to study. Here we describe a novel experimental approach used to create a suite of ice storms in a mature hardwood forest in New Hampshire, USA. The experiment included five ice storm intensities (0, 6.4, 12.7 and 19.1 mm radial ice accretion) applied in a single year, and one ice storm intensity (12.7 mm) applied in two consecutive years. Results demonstrate the feasibility of this approach for creating experimental ice storms, quantify the increase in fine and coarse woody debris mass and nutrients transferred from the forest canopy to the soil under the different icing conditions, and show an increase in the damage to the forest canopy with increasing icing that evolves over time. In this forest, little damage occurred below 6.4 mm radial ice accretion, moderate damage occurred with up to 12.7 mm of accretion, and significant branch breakage and canopy damage occurred with 19.1 mm of ice. The icing in consecutive years demonstrated an interactive effect of ice storm frequency and severity such that some branches damaged in the first year of icing appeared to remain in the canopy and then fall to the ground in the second year of icing. These results have implications for National Weather Service ice storm warning levels, as they provide a quantitative assessment of ice-load related inputs of forest debris that will be useful to municipalities creating response plans for current and future ice storms.

Simulating Impacts of Ice Storms on Forest Ecosystems.

Ice storms can have profound and lasting effects on the structure and function of forest ecosystems in regions that experience freezing conditions. Current models suggest that the frequency and intensity of ice storms could increase over the coming decades in response to changes in climate, heightening interest in understanding their impacts. Because of the stochastic nature of ice storms and difficulties in predicting when and where they will occur, most past investigations of the ecological effects of ice storms have been based on case studies following major storms. Since intense ice storms are exceedingly rare events it is impractical to study them by waiting for their natural occurrence. Here we present a novel alternative experimental approach, involving the simulation of glaze ice events on forest plots under field conditions. With this method, water is pumped from a stream or lake and sprayed above the forest canopy when air temperatures are below freezing. The water rains down and freezes upon contact with cold surfaces. As the ice accumulates on trees, the boles and branches bend and break; damage that can be quantified through comparisons with untreated reference stands. The experimental approach described is advantageous because it enables control over the timing and amount of ice applied. Creating ice storms of different frequency and intensity makes it possible to identify critical ecological thresholds necessary for predicting and preparing for ice storm impacts.

Ticks as Soil-Dwelling Arthropods: An Intersection Between Disease and Soil Ecology.

Ticks are widespread vectors for many important medical and veterinary infections, and a better understanding of the factors that regulate their population dynamics is needed to reduce risk for humans, wildlife, and domestic animals. Most ticks, and all non-nidicolous tick species, spend only a small fraction of their lives associated with vertebrate hosts, with the remainder spent in or on soils and other substrates. Ecological studies of tick-borne disease dynamics have emphasized tick-host interactions, including host associations, burdens, and efficiencies of pathogen transmission, while under emphasizing tick biology during off-host periods. Our ability to predict spatiotemporal trends in tick-borne diseases requires more knowledge of soil ecosystems and their effect on host and tick populations. In this review, we focus on tick species of medical and veterinary concern and describe: 1) the relationships between soil factors and tick densities; 2) biotic and abiotic factors within the soil ecosystem that directly affect tick survival; 3) potential indirect effects on ticks mediated by soil ecosystem influences on their vertebrate hosts; 4) the potential for tick-mediated effects on vertebrate host populations to affect ecosystems; and 5) possible nontarget impacts of tick management on the soil ecosystem. Soils are complex ecosystem components with enormous potential to affect the survival and behavior of ticks during their off-host periods. Hence, tick-borne disease systems present an excellent opportunity for soil ecologists and public health researchers to collaborate and improve understanding of these medically important and ecologically complex disease cycles.

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.

The application of an integrated biogeochemical model to simulate dynamics of vegetation, hydrology and nutrients in soil and streamwater following a whole-tree harvest of a northern hardwood forest.

Understanding the impacts of clear-cutting is critical to inform sustainable forest management associated with net primary productivity and nutrient availability over the long-term. Few studies have rigorously tested model simulations against field measurements which would provide more confidence in efforts to quantify logging impacts over the long-term. The biogeochemical model, PnET-BGC has been used to simulate forest production and stream chemistry at the Hubbard Brook Experimental Forest (HBEF), New Hampshire, USA. Previous versions of PnET-BGC could accurately simulate the longer-term biogeochemical response to harvesting, but were unable to reproduce the marked changes in stream NO3- immediately after clear-cutting which is an important impact of forest harvesting. Moreover, the dynamics of nutrients in major pools including mineralization and plant uptake were poorly predicted. In this study, the model was modified and parametrized allowing for a lower decomposition rate during the earlier years after the clear-cut and increased NH4+ plant uptake with the regrowth of new vegetation to adequately reproduce hydrology, aboveground forest biomass, and soil solution and stream water chemistry in response to a whole-tree harvest of a northern hardwood forest watershed (W5) at the HBEF. Modeled soil solution and stream water chemistry successfully captured the rapid recovery of leaching nutrients to pre-cut levels within four years after the treatment. The model simulated a substantial increase in aboveground net primary productivity (NPP) from around 36% to 97% of pre-cut aboveground values within years 2 to 4 of the cut, which closely reproduced the measured values. The projected accumulation of aboveground biomass 70 years following the harvest was almost 190 t ha-1, which is close to the pre-cut measured value. A first-order sensitivity analysis showed greater sensitivity of projections of the model outputs for the mature forest than the strongly aggrading forest.

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.