North American Temperate Forests Research Articles

Interannual variability and seasonality of litterfall in three temperate and boreal forest ecosystems of eastern Canada: A synthesis of long-term monitoring

Litterfall is a major pathway for transferring aboveground biomass to the forest floor and thus plays an important role in building forest soil carbon stocks. However, inter- and intra-annual variability of litterfall remains poorly documented, especially in North American temperate and boreal forests, due to the lack of recent long-term studies at high sampling frequencies. This potentially creates uncertainties in estimates of forest carbon budget models. The objectives of the present study were to 1) quantify the mean annual flux, interannual variability, and seasonality of litterfall in three sites (dominated respectively by sugar maple (Acer saccharum Marsh.), balsam fir (Abies balsamea (L.) Mill. 1768), and black spruce (Picea mariana (Mill.) B.S.P.)) in eastern Canada over a period of 22–32 years, 2) relate the litterfall amounts and temporal variations to the changes in the size of major organic matter pools in these ecosystems, and 3) compare our litterfall estimates with reference values used in national greenhouse gas inventories. Litterfall production decreased from the sugar maple to the balsam fir and black spruce sites, preponderantly due to species composition. Litterfall evolution was related to the aboveground biomass of live trees in both conifer sites; in contrast, in the broadleaf site, changes in forest composition and structure were apparently the main drivers. The litterfall seasonality varied between broadleaf and conifer sites and could be explained by a sigmoidal model. Substantial departures from the seasonality for some given years were likely due to important climatic anomalies. Forest floor biomass remained stable over time at all three sites despite the increase in litterfall at the balsam fir and sugar maple sites and rapid forest floor turnover at the latter site. Our analyses of litterfall suggest that reference values from the literature used for national greenhouse gas inventories underestimate annual litterfall and forest floor carbon stocks for temperate and boreal forests.

Examining climate and composition differences in eastern North American temperate forests using pollen ratios and pollen assemblage cluster analysis

Examining climate and composition differences in eastern North American temperate forests using pollen ratios and pollen assemblage cluster analysis

The impact of spongy moth (Lymantria dispar dispar) defoliation on carbon balance of a temperate deciduous forest in North America

Temperate deciduous forests are an important contributor to the global carbon (C) sink. However, changes in environmental conditions and natural disturbances such as insect infestations can impact carbon sequestration capabilities of these forests. While, insect infestations are expected to increase in warmer future climates, there is a lack of knowledge on the quantitative impact of these natural disturbances on the carbon balance of temperate deciduous forests. In 2021, a record-breaking defoliation, caused by the spongy moth (Lymantria dispar dispar (LDD), formerly knows as the gypsy moth) occurred in eastern North America. In this study, we assess the impact of this spongy moth defoliation on carbon uptake in a mature oak-dominated temperate forest in the Great Lakes region in Canada, using eddy covariance flux data from 2012 to 2022. Study results showed that the forest was a large C sink with mean annual net ecosystem productivity (NEP) of 207 ± 77 g C m–2 yr−1 from 2012 to 2022, excluding 2021, which experienced the infestation. Over this period mean annual gross ecosystem productivity (GEP) was 1,398 ± 137 g C m–2 yr−1, while ecosystem respiration (RE) was 1,209 ± 139 g C m–2 yr−1. However, in 2021 due to defoliation in the early growing season, annual GEP of the forest declined to 959 g C m–2 yr−1, while annual RE increased to 1,345 g C m–2 yr−1 causing the forest to become a large source of C with annual NEP of -351 g C m–2 yr−1. The forest showed a rapid recovery from this major disturbance event, with annual GEP, RE, and NEP values of 1,671, 1,287, and 298 g C m–2 yr−1, respectively in 2022 indicating that the forest was once again a large C sink. This study demonstrates that major transient natural disturbances under changing climate can have a significant impact on forest C dynamics. The extent to which North American temperate forests will remain a major C sink will depend on the severity and intensity of these disturbance events and the rate of recovery of forests following disturbances. Lay abstractTemperate deciduous forests play an important role in carbon sequestration from the atmosphere. However, the impact of climate change, extreme weather, and disturbance events can alter the extent to which these forests sequester carbon, in some cases shifting their role from being a carbon sink to becoming a carbon source to the atmosphere. In 2021, a spongy moth infestation severely defoliated a mature oak-dominated temperate forest north of Lake Erie, Ontario, Canada, turning the forest from a 10-year mean carbon sink of about 2 tons of carbon per hectare to a carbon source of about 3.5 tons of carbon per hectare. Our analysis indicates that meteorological conditions during the early growing season might have influenced the severity of this infestation. Specifically, the prevalence of dry and warm weather conditions enabled the moth to survive and thrive longer. This study shows the significant influence of natural disturbances on forest carbon dynamics as temperatures continue to rise due to climate change. The future role forests play in carbon sequestration will be determined by the severity of disturbance events and the effectiveness of forests to recover in the aftermath of these events.

Genetic variation in growth and leaf traits associated with local adaptation to climate in yellow birch (Betula alleghaniensis Britton)

Understanding patterns of variation in functional traits of hardwood trees is crucial for conserving and managing North American temperate forests under climate change. This study examined provenance variation of yellow birch ( Betula alleghaniensis Britton) in growth, biomass allocation, leaf morphology, and stable carbon isotope composition. Trees were grown from 10 seed sources originating from across Canada and the northern USA. Height and diameter were not significantly related to climate at seed origin, suggesting that variation may be better explained by site factors, such as soil pH and soil moisture. In contrast, carbon isotope composition and leaf morphological traits were significantly correlated to climate variables including temperature, precipitation, and solar radiation. Provenances from warmer, drier localities tended to have higher stable carbon isotope ratio (δ13C), greater specific leaf area, and narrower leaf width than their counterparts from cooler, wetter climates. Thus, variation in leaf morphological traits appears to be involved in adaptation of yellow birch to variation in temperature and moisture availability across the species’ range. Our results suggest that there may exist potential for selection and breeding of drought-resistant yellow birch genotypes to aid in reforestation under climate change.

Linked disturbance in the temperate forest: Earthworms, deer, and canopy gaps.

Despite the large body of theory concerning multiple disturbances, relatively few attempts have been made to test the theoretical assumptions of how and if disturbances interact. Of particular importance is whether disturbance events are linked, as this can influence the probability and intensity of ecological change. Disturbances are linked when one disturbance event increases or decreases the likelihood or extent of another. To this end, we used two long-term, multi-disturbance experiments in northern Wisconsin to determine whether earthworm invasion is linked to canopy gap creation and white-tailed deer browsing. These three disturbances are common and influential within North American temperate forests, making any interactions among them particularly important to understand. We expected both deer and canopy gaps to favor invasive earthworms, particularly species that live close to or on the soil surface. However, we found only partial support of our hypotheses, as both deer exclosures and canopy gaps decreased earthworms in each experiment. Further, earthworm density increased the most over time in areas far from the gap center and in areas with deer present. Deer exclosures primarily decreased Aporrectodea and Lumbricus species, while gaps decreased Dendrobaena and Lumbricus species. Our findings show that earthworm invasion is linked to deer presence and gap-creating disturbances, which provides new insight in multiple disturbance theory, aboveground-belowground dynamics, and temperate forest management.

Before-and-after evidence that urbanization contributes to the decline of a migratory songbird

Although urbanization is widely believed to be an increasing threat to biodiversity, drawing strong inferences about its effects on wildlife has been challenging because the state of a population prior to development is rarely known. The Wood Thrush (<em>Hylocichla mustelina</em>) is a steeply declining migratory songbird that breeds in North American temperate deciduous forests and winters in Central America. From 1987 to 2001, Wood Thrush abundance and daily nest survival were measured in 72 forest fragments across Waterloo Region, Ontario, Canada. Some of these fragments have since been surrounded by urbanization while others have remained rural. In 2020 and 2021, we reassessed Wood Thrush abundance and daily nest survival in these same fragments by replicating point count and nest monitoring protocols. Our results suggest that Wood Thrush have declined most steeply in fragments with development built within 1 km during the intervening 20-year period (-79% vs. only -57% elsewhere), although experiencing no decline in daily nest survival over the same time period, regardless of development. Our findings provide rare before-and-after evidence that urbanization near breeding habitat is contributing to a pronounced localized decline of a migratory songbird, likely by mechanisms unrelated to nest survival.

Predicting Lumber Grade Occurrence and Volume in Sugar Maple and Yellow Birch Logs

AbstractThe North American temperate deciduous forests are an important source of hardwoods sought after by the appearance wood products industries. The purpose of this study was to model the relationships between log characteristics and sawn board attributes in sugar maple (Acer saccharum Marsh.) and yellow birch (Betula alleghaniensis Britt.). We harvested sixty-four sugar maple and thirty-two yellow birch trees from two locations in southern Quebec, Canada, which were then processed into 189 sawlogs and 2,236 boards. We developed statistical models linking log characteristics to the volume recovery of the various lumber grades and color specifications according to the National Hardwood Lumber Association rules. In both species, board grades were strongly related to log length, position of the log in the stem, and small-end diameter and decay of the log. Color classes were related to small-end diameter of the log and red heartwood diameter of the log at both ends for sugar maple, and to log length, diameter of the log at the small-end, and red heartwood diameter at the large-end of the log for yellow birch. These models outperformed a log classification used in practice for predicting lumber volume recovery from different log grades.

Effects of whole-tree and stem-only clearcutting on forest floor and soil carbon and nutrients in a balsam fir (Abies balsamea (L.) Mill.) and red spruce (Picea rubens Sarg.) dominated ecosystem

Intensive harvesting of forest biomass for bioenergy has the potential to degrade forest soils and productivity if ecosystem carbon and other nutrients are depleted faster than replenished naturally or by management inputs. Climate change mitigation potential associated with bioenergy may be threatened if forest management operations reduce soil carbon stocks. Research reported here was initiated in 1979 to determine the effects of whole-tree harvesting on long-term site productivity and ecosystem carbon and nutrient pools and fluxes in a northeastern North American temperate balsam fir-red spruce forest. A sequence of harvesting and silvicultural treatments were applied from 1981 to 1993 using a paired-watershed experimental design. Treatments included whole-tree clearcutting, stem-only harvesting, simulated by return of chipped or lopped delimbing residue, conifer release by applying herbicide, and pre-commercial thinning and fertilizer application. Quantitative samples of the forest floor were excavated in 2016. Quantitative soil pits were sampled in 2017 to a depth of 50 cm below the forest floor, where possible; the basal till subsoil was sampled to a depth of 100 cm with an auger. Forest floor samples were analyzed for total carbon, nitrogen, phosphorus, potassium, calcium, magnesium and aluminum. Organic and mineral soil layer samples from the quantitative pits were analyzed for pH, short and long-lived soil carbon via 13C cross polarization magic angle spinning (CP-MAS) nuclear magnetic resonance (NMR), total carbon and nitrogen and extractable phosphorus, potassium, calcium, magnesium and aluminum. Forest floor carbon and nitrogen on whole-tree harvested plots 35 years after harvest were about 50% of the amount on the unharvested watershed. Stem-only harvesting partially mitigated the reduction, and gross losses were apparently offset by increases in the mineral soil solum to depth 50 cm, suggesting that downward translocation of soil organic matter was a significant ecosystem process during the 35 years since harvest. Accumulated forest floor and mineral soil stock of total carbon and nitrogen and extractable nutrients to depth 100 cm did not differ significantly between the harvested and unharvested watershed. A corresponding similarity in mineral soil pools of extractable phosphorus and non-acid cations in the two watersheds suggests that inputs from atmospheric deposition, primary and secondary mineral weathering, organic matter mineralization and litterfall balanced exports including leaching and plant uptake. Ecosystem stocks of extractable phosphorus, base cations, and total organic carbon and nitrogen were maintained 35 years after whole-tree harvesting of a primary spruce-fir forest.

Climate Change Increases the Severity and Duration of Soil Water Stress in the Temperate Forest of Eastern North America

Under climate change, drought conditions are projected to intensify and soil water stress is identified as one of the primary drivers of the decline of forests. While there is strong evidence of such megadisturbance in semi-arid regions, large uncertainties remain in North American temperate forests and fine-scale assessments of future soil water stress are needed to guide adaptation decisions. The objectives of this study were to (i) assess the impact of climate change on the severity and duration of soil water stress in a temperate forest of eastern North America and (ii) identify environmental factors driving the spatial variability of soil water stress levels. We modeled current and future soil moisture at a 1 km resolution with the Canadian Land Surface Scheme (CLASS). Despite a slight increase in precipitation during the growing season, the severity (95th percentile of absolute soil water potential) and duration (number of days where absolute soil water potential is greater than or equal to 9,000 hPa) of soil water stress were projected to increase on average by 1,680 hPa and 6.7 days in 80 years under RCP8.5, which correspond to a 33 and 158% increase compared to current levels. The largest increase in severity was projected to occur in areas currently experiencing short periods of soil water stress, while the largest increase in duration is rather likely to occur in areas already experiencing prolonged periods of soil water stress. Soil depth and, to a lesser extent, soil texture, were identified as the main controls of the spatial variability of projected changes in the severity and duration of soil water stress. Overall, these results highlight the need to disentangle impacts associated with an increase in the severity vs. in the duration of soil water stress to guide the management of temperate forests under climate change.

Beech Bark Disease in an Unmanaged Temperate Forest: Patterns, Predictors, and Impacts on Ecosystem Function

Beech Bark Disease (BBD) is a devastating threat to American beech (Fagus grandifolia), spreading through eastern mixed deciduous forests of North America at unprecedented rates. Understanding how and why some beech trees escape severe BBD effects is important; however, the biotic and abiotic factors that underpin the progression of BBD within unmanaged forests at local scales are not well explored. We surveyed 651 individual beech trees ≥ 10 cm diameter at breast height (DBH) for BBD, in a 13.5-ha unmanaged forest dynamics plot in Ontario, Canada, where &amp;gt;46,000 trees have been identified to species, mapped, and DBH measured at ∼5-year intervals. For each beech tree, BBD severity was ranked on a 5-point severity index, which was then evaluated as a function of tree characteristics including DBH and relative growth rate (RGR). Most beech trees were at either the insect or fungal stage of BBD, with only 22% of beech trees being free of symptoms. Ordinal logistic regression analysis indicated both DBH and RGR were significant predictors of BBD severity. These models, along with both randomization and Moran’s Eigenvector Maps (MEM) analyses, indicated that DBH and RGR and their spatial variation accounted for ∼44.6% of BBD severity in trees. Our MEMs also indicated ∼4.2% of variation in BBD severity was associated with unmeasured spatial variables, which may reflect either the spread of BBD through our study site, or the influence of abiotic variables on BBD severity. At our site, BBD is responsible for at least ∼6.0 Mg C ha–1, or ∼6.5% of the average 92.5 Mg of aboveground biomass C ha–1, transitioning from the live to dead biomass pool. Our study enhances the understanding of the factors predicting the severity of a major forest pathogen in North American temperate forests, assists the integration of BBD severity risk into forest C budget models, and provides insight into how large-scale forest inventories can inform screening for pest or pathogen resistance in trees.

Changes in Understory Composition of Rural North American Temperate Forests after a 14-Year Period with Focus on Exotic and Sensitive Plant Species

A better understanding of the mechanisms influencing compositional changes in understory plant communities is crucial to protect temperate forests against global change stressors, including anthropogenic disturbances and invasion by exotic species. We assessed changes in species composition after a 14-year period in 20 rural temperate forest stands located in the northern hardwood biome of eastern Canada. We identified species that underwent the largest changes in relative occurrence during that period and assessed the influence of biotic and anthropogenic filters on the trajectory of those understory communities. We found small but significant compositional changes after 14 years, mostly related to a decrease in species diversity in the younger forest stands originating from abandoned pasture. The largest occurrence gains for understory species were observed in these stands, but also in stands with lower understory community diversity. Understory species occurrence losses could not be linked to any of the biotic and anthropogenic filters considered in this study presumably because they were difficult to isolate from secondary successional dynamics. Shade tolerant exotic species showed a small but significant increase in relative occurrence over 14 years, with notable gains in only a few stands. We observed generalized occurrence gains for sensitive spring geophytes during the same period throughout most sites. Overall, the understory vegetation community of rural temperate forests in this region was found to be relatively stable and seems to retain its potential for natural recovery after disturbance.

Heat and drought impact on carbon exchange in an age-sequence of temperate pine forests

BackgroundMost North American temperate forests are plantation or regrowth forests, which are actively managed. These forests are in different stages of their growth cycles and their ability to sequester atmospheric carbon is affected by extreme weather events. In this study, the impact of heat and drought events on carbon sequestration in an age-sequence (80, 45, and 17 years as of 2019) of eastern white pine (Pinus strobus L.) forests in southern Ontario, Canada was examined using eddy covariance flux measurements from 2003 to 2019.ResultsOver the 17-year study period, the mean annual values of net ecosystem productivity (NEP) were 180 ± 96, 538 ± 177 and 64 ± 165 g C m–2 yr–1 in the 80-, 45- and 17-year-old stands, respectively, with the highest annual carbon sequestration rate observed in the 45-year-old stand. We found that air temperature (Ta) was the dominant control on NEP in all three different-aged stands and drought, which was a limiting factor for both gross ecosystem productivity (GEP) and ecosystems respiration (RE), had a smaller impact on NEP. However, the simultaneous occurrence of heat and drought events during the early growing seasons or over the consecutive years had a significant negative impact on annual NEP in all three forests. We observed a similar trend of NEP decline in all three stands over three consecutive years that experienced extreme weather events, with 2016 being a hot and dry, 2017 being a dry, and 2018 being a hot year. The youngest stand became a net source of carbon for all three of these years and the oldest stand became a small source of carbon for the first time in 2018 since observations started in 2003. However, in 2019, all three stands reverted to annual net carbon sinks.ConclusionsOur study results indicate that the timing, frequency and concurrent or consecutive occurrence of extreme weather events may have significant implications for carbon sequestration in temperate conifer forests in Eastern North America. This study is one of few globally available to provide long-term observational data on carbon exchanges in different-aged temperate plantation forests. It highlights interannual variability in carbon fluxes and enhances our understanding of the responses of these forest ecosystems to extreme weather events. Study results will help in developing climate resilient and sustainable forestry practices to offset atmospheric greenhouse gas emissions and improving simulation of carbon exchange processes in terrestrial ecosystem models.

Intermediate disturbances drive long‐term fluctuation in old‐growth forest biomass: an 84‐yr temperate forest record

AbstractWhat are typical values and dynamic status of live‐tree biomass pools in old‐growth, mesic, cool temperate forests? A handful of biomass density estimates in eastern North American temperate forests show large biomass/carbon reserves on a per‐area basis. However, it is less clear whether these ecosystems are, over multi‐decade scales, typically steady‐state or non‐equilibrial carbon pools. Previous studies have suggested both possibilities, but claims are based on inferences from short‐term studies or proxy data sets. An unusually long‐term and extensive data set from repeatedly sampled permanent plots (84 yr, ca. 10 ha sample area, 6–8 measurements), from old‐growth conifer‐hardwood forest in northern Michigan, USA, allows direct estimation of multi‐decade trends in aboveground live‐tree biomass. Results confirm prior suggestions of high‐biomass density for old‐growth temperate forests (averaging &gt;300 Mg/ha), but, despite significant decade‐scale variation, show no overall, long‐term directional change. Study plots typically show multi‐decade trends of gradually increasing biomass density, interrupted by sharp declines attributed to intermediate‐severity disturbances, with recovery of pre‐disturbance biomass density requiring upwards of a half‐century. At the stand scale, biomass dynamics are strongly historically contingent, and short‐term studies may yield biased or misleading results. Disturbance legacies, through demographic and structural effects, can have multi‐decade effects on vulnerability to further disturbance. While this study shows no general trend in aboveground biomass pools, it suggests that changes in disturbance regime may drive important feedbacks in biomass pool dynamics.

Woodpeckers and other excavators maintain the diversity of cavity-nesting vertebrates.

Woodpeckers and other excavators create most of the holes used by secondary cavity nesters (SCNs) in North American temperate mixedwood forests, but the degree to which excavators release SCNs from nest-site limitation is debated. Our goal was to quantify how excavators maintain the diversity and abundance of secondary cavity nesters in a temperate forest through the creation of tree cavities. We examined the short- and long-term (legacy) effects of excavators (principally woodpeckers, but also red-breasted nuthatches and black-capped chickadees) on forest biodiversity using longitudinal monitoring data (1,732 nest cavities, 25 sites, 16years) in British Columbia, Canada. Sites with higher densities of excavator nests had more cavities available, higher species richness of SCNs and higher nest density of SCNs, indicating the importance of a standing stock of cavities. Years with higher nesting densities of excavators were followed by years with higher SCN diversity, indicating that the creation of nesting opportunities through fresh excavation releases SCNs from community-wide nest-site limitation. We also show that excavators leave a 'legacy' of biodiversity (species richness and abundance) at a site by accumulating cavities at rates faster than they become unusable by decay or destruction. By quantifying site-level effects of cavity excavation on the SCN community, our study highlights the key role of excavators as ecosystem engineers that maintain forest wildlife biodiversity.

Are we missing the forest for the trees? Conspecific negative density dependence in a temperate deciduous forest.

One of the central goals of ecology is to determine the mechanisms that enable coexistence among species. Evidence is accruing that conspecific negative density dependence (CNDD), the process by which plant seedlings are unable to survive in the area surrounding adults of their same species, is a major contributor to tree species coexistence. However, for CNDD to maintain community-level diversity, three conditions must be met. First, CNDD must maintain diversity for the majority of the woody plant community (rather than merely specific groups). Second, the pattern of repelled recruitment must increase in with plant size. Third, CNDD should extend to the majority of plant life history strategies. These three conditions are rarely tested simultaneously. In this study, we simultaneously test all three conditions in a woody plant community in a North American temperate forest. We examined whether understory and canopy woody species across height categories and dispersal syndromes were overdispersed-a spatial pattern indicative of CNDD-using spatial point pattern analysis across life history stages and strategies. We found that there was a strong signal of overdispersal at the community level. Across the whole community, larger individuals were more overdispersed than smaller individuals. The overdispersion of large individuals, however, was driven by canopy trees. By contrast, understory woody species were not overdispersed as adults. This finding indicates that the focus on trees for the vast majority of CNDD studies may have biased the perception of the prevalence of CNDD as a dominant mechanism that maintains community-level diversity when, according to our data, CNDD may be restricted largely to trees.

Phylogenomics and biogeography of Wisteria: Implications on plastome evolution among inverted repeat‐lacking clade (IRLC) legumes

AbstractThe genus Wisteria (Fabaceae) is disjunctly distributed in eastern Asian and eastern North American temperate deciduous forests, and it is widely cultivated around the world as spectacular garden plants. It is a member of inverted repeat‐lacking clade (IRLC). The IRLC Species are characterized by the loss of an IR region in their plastomes, which has long been of great interest. In this research, we report whole plastome sequences from all four Wisteria species and a Wisteriopsis japonica, combining these with existing data to explore phylogenetic relationships and biogeography of Wisteria, as well as plastome evolution of IRLC species. Phylogenetic analyses recognized a clade containing Glycyrrhiza–Wisteriopsis–Wisteria as sister to the remaining genera of IRLC. North American Wisteria frutescens and the three Asian species formed reciprocal clades, and Wisteria brachybotrys was sister to Wisteria floribunda and Wisteria sinensis. Wisteria may have originated in Japan near the boundary of the Oligocene and Miocene. The disappearance of Bering Land Bridge in the late Miocene might lead to the Eastern Asian–Eastern North American disjunction of Wisteria. Allopatric speciation of Wisteria between the Japanese archipelago and the Asian continent in the Quaternary increased the species richness of eastern Asia in comparison with eastern North America. Synonymous substitution rates (dS) of protein‐coding genes in the IRLC species were around 2‐fold (SC genes) or 11‐fold (IR genes) higher than those of non‐IRLC species. For both SC and IR genes, herbaceous legumes have around 3‐fold higher dS than woody ones. Both loss of one IR region and herbaceous habit elevated substitution rates of the plastomes.

Insect reproductive behaviors are important mediators of carrion nutrient release into soil

Current declines in terrestrial insect biomass and abundance have raised global concern for the fate of insects and the ecosystem services they provide. However, the ecological and economic contributions of many insects have yet to be quantified. Carrion-specializing invertebrates are important mediators of carrion decomposition; however, the role of their reproductive activities in facilitating this nutrient pulse into ecosystems is poorly understood. Here, we investigate whether insects that sequester carrion belowground for reproduction alter soil biotic and abiotic properties in North American temperate forests. We conducted a field experiment that measured soil conditions in control, surface carrion alone, and beetle-utilized carrion treatments. Our data demonstrate that Nicrophorus beetle reproduction and development results in changes in soil characteristics which are consistent with those observed in surface carrion decomposition alone. Carrion addition treatments increase soil labile C, DON and DOC, while soil pH and microbial C:N ratios decrease. This study demonstrates that the decomposition of carrion drives soil changes but suggests that the behaviors of insect scavengers play an important role in the release of carrion nutrients directly into the soil by sequestering carrion resources in the ecosystem where they were deposited.

Density dependence and habitat heterogeneity regulate seedling survival in a North American temperate forest

Survival and growth of forest tree seedlings are influenced by many abiotic and biotic forces that may vary across space and time. The simultaneous influences of habitat heterogeneity, temporal environmental variability (including disturbance regimes), and biotic interactions are difficult to disentangle, yet understanding the relative importance of these factors for tree seedling dynamics is critical for conservation of forest biodiversity. Most long term, spatially explicit studies of tree seedling ecology have been set in the tropics; much less attention has been given to tree seedlings in temperate forests. We monitored the survival of over 3000 individual seedlings over an eight-year period in a temperate forest in northeast Wisconsin, USA. Results from four seedling censuses demonstrated that both conspecific density and environmental variables significantly affected seedling survival. Higher densities of conspecific trees consistently reduced the probability of seedling survival over time. At the community level, relatively common species were negatively influenced by neighboring conspecific trees and exhibited higher per capita and per basal area mortality. The negative impacts of high conspecific tree density were most pronounced in areas of higher light and moisture, but these interactions varied over time, as did the importance of other abiotic variables. Patterns of less common species were more clearly explained by abiotic variables, with shifting relevance of specific variables according to species abundance. Conspecific negative density dependence, which interacts with resource gradients, and habitat conditions that shift over time are influencing community composition in this northern temperate forest.

Response of carbon and water fluxes to meteorological and phenological variability in two eastern North American forests of similar age but contrasting species composition – a multiyear comparison

Abstract. The annual carbon and water dynamics of two eastern North American temperate forests were compared over a 6-year period from 2012 to 2017. The geographic location, forest age, soil, and climate were similar between the two stands; however, stand composition varied in terms of tree leaf-retention and shape strategy: one stand was a deciduous broadleaf forest, while the other was an evergreen needleleaf forest. The 6-year mean annual net ecosystem productivity (NEP) of the coniferous forest was slightly higher and more variable (218±109 g C m−2 yr−1) compared to that of the deciduous forest NEP (200±83 g C m−2 yr−1). Similarly, the 6-year mean annual evapotranspiration (ET) of the coniferous forest was higher (442±33 mm yr−1) than that of the deciduous forest (388±34 mm yr−1), but with similar interannual variability. Summer meteorology greatly impacted the carbon and water fluxes in both stands; however, the degree of response varied among the two stands. In general, warm temperatures caused higher ecosystem respiration (RE), resulting in reduced annual NEP values – an impact that was more pronounced at the deciduous broadleaf forest compared to the evergreen needleleaf forest. However, during warm and dry years, the evergreen forest had largely reduced annual NEP values compared to the deciduous forest. Variability in annual ET at both forests was related most to the variability in annual air temperature (Ta), with the largest annual ET observed in the warmest years in the deciduous forest. Additionally, ET was sensitive to prolonged dry periods that reduced ET at both stands, although the reduction at the coniferous forest was relatively larger than that of the deciduous forest. If prolonged periods (weeks to months) of increased Ta and reduced precipitation are to be expected under future climates during summer months in the study region, our findings suggest that the deciduous broadleaf forest will likely remain an annual carbon sink, while the carbon sink–source status of the coniferous forest remains uncertain.

Tracking invasions of a destructive defoliator, the gypsy moth (Erebidae: Lymantria dispar): Population structure, origin of intercepted specimens, and Asian introgression into North America.

Genetic data can help elucidate the dynamics of biological invasions, which are fueled by the constant expansion of international trade. The introduction of European gypsy moth (Lymantria dispar dispar) into North America is a classic example of human‐aided invasion that has caused tremendous damage to North American temperate forests. Recently, the even more destructive Asian gypsy moth (mainly L. d. asiatica and L. d. japonica) has been intercepted in North America, mostly transported by cargo ships. To track invasion pathways, we developed a diagnostic panel of 60 DNA loci (55 nuclear and 5 mitochondrial) to characterize worldwide genetic differentiation within L. dispar and its sister species L. umbrosa. Hierarchical analyses supported strong differentiation and recovered five geographic groups that correspond to (1) North America, (2) Europe plus North Africa and Middle East, (3) the Urals, Central Asia, and Russian Siberia, (4) continental East Asia, and (5) the Japanese islands. Interestingly, L. umbrosa was grouped with L. d. japonica, and the introduced North American population exhibits remarkable distinctiveness from contemporary European counterparts. Each geographic group, except for North America, shows additional lower‐level structures when analyzed individually, which provided the basis for inference of the origin of invasive specimens. Two assignment approaches consistently identified a coastal area of continental East Asia as the major source for Asian invasion during 2014–2015, with Japan being another source. By analyzing simulation and laboratory crosses, we further provided evidence for the occurrence of natural Asian–North American hybrids in the Pacific Northwest, raising concerns for introgression of Asian alleles that may accelerate range expansion of gypsy moth in North America. Our study demonstrates how genetic data contribute to bio‐surveillance of invasive species with results that can inform regulatory management and reduce the frequency of trade‐associated invasions.