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Conspecific negative density dependence (CNDD) promotes tree species diversity by reducing recruitment near conspecific adults due to biotic feedbacks from herbivores, pathogens, or competitors. While this process is well-described in tropical forests, tests of temperate tree species range from strong positive to strong negative density dependence. To explain this, several studies have suggested that tree species traits may help predict the strength and direction of density dependence: for example, ectomycorrhizal-associated tree species typically exhibit either positive or weaker negative conspecific density dependence. More generally, the strength of density dependence may be predictably related to other species-specific ecological attributes such as shade tolerance, or the relative local abundance of a species. To test the strength of density dependence and whether it affects seedling community diversity in a temperate forest, we tracked the survival of seedlings of three ectomycorrhizal-associated species experimentally planted beneath conspecific and heterospecific adults on the Prospect Hill tract of the Harvard Forest, in Massachusetts, USA. Experimental seedling survival was always lower under conspecific adults, which increased seedling community diversity in one of six treatments. We compared these results to evidence of CNDD from observed sapling survival patterns of 28 species over approximately 8 years in an adjacent 35-ha forestmore »plot. We tested whether species-specific estimates of CNDD were associated with mycorrhizal association, shade tolerance, and local abundance. We found evidence of significant, negative conspecific density dependence (CNDD) in 23 of 28 species, and positive conspecific density dependence in two species. Contrary to our expectations, ectomycorrhizal-associated species generally exhibited stronger (e.g., more negative) CNDD than arbuscular mycorrhizal-associated species. CNDD was also stronger in more shade-tolerant species but was not associated with local abundance. Conspecific adult trees often have a negative influence on seedling survival in temperate forests, particularly for tree species with certain traits. Here we found strong experimental and observational evidence that ectomycorrhizal-associating species consistently exhibit CNDD. Moreover, similarities in the relative strength of density dependence from experiments and observations of sapling mortality suggest a mechanistic link between negative effects of conspecific adults on seedling and sapling survival and local tree species distributions.« less

Abstract Hemlock woolly adelgid (HWA; Adelges tsugae Annand (Hemiptera: Adelgidae)) is the cause of widespread mortality of Carolina and eastern hemlock (Tsuga caroliniana Engelmann and T. canadensis (L.) Carrière) throughout the eastern United States (U.S.). Since its arrival in the northeastern U.S., HWA has steadily invaded and established throughout eastern hemlock stands. However, in 2018, anecdotal evidence suggested a sharp, widespread HWA decline in the northeastern U.S. following above-average summer and autumn rainfall. To quantify this decline in HWA density and investigate its cause, we surveyed HWA density in hemlock stands from northern Massachusetts to southern Connecticut and analyzed HWA density and summer mortality in Pennsylvania. As native fungal entomopathogens are known to infect HWA in the northeastern U.S. and rainfall facilitates propagation and spread of fungi, we hypothesized high rainfall facilitates fungal infection of aestivating nymphs, leading to a decline in HWA density. We tested this hypothesis by applying a rain-simulation treatment to hemlock branches with existing HWA infestations in western MA. Our results indicate a regional-scale decline and subsequent rebound in HWA density that correlates with 2018 rainfall at each site. Experimental rain treatments resulted in higher proportions of aestivating nymphs with signs of mortality compared to controls.more »In conjunction with no evidence of increased mortality from extreme winter or summer temperatures, our results demonstrate an indirect relationship between high rainfall and regional HWA decline. This knowledge may lead to better prediction of HWA population dynamics.« less

Land-use history is the template upon which contemporary plant and tree populations establish and interact with one another and exerts a legacy on the structure and dynamics of species assemblages and ecosystems. We use the first census (2010–2014) of a 35-ha forest-dynamics plot at the Harvard Forest in central Massachusetts to describe the composition and structure of the woody plants in this plot, assess their spatial associations within and among the dominant species using univariate and bivariate spatial point-pattern analysis, and examine the interactions between land-use history and ecological processes. The plot includes 108,632 live stems ≥ 1 cm in diameter (2,215 individuals/ha) and 7,595 standing dead stems ≥ 5 cm in diameter. Live tree basal area averaged 42.25 m 2 /ha, of which 84% was represented by Tsuga canadensis (14.0 m 2 / ha), Quercus rubra (northern red oak; 9.6 m2/ ha), Acer rubrum (7.2 m 2 / ha) and Pinus strobus (eastern white pine; 4.4 m 2 / ha). These same four species also comprised 78% of the live aboveground biomass, which averaged 245.2 Mg/ ha. Across all species and size classes, the forest contains a preponderance (> 80,000) of small stems (<10-cm diameter) that exhibit a reverse-Jmore »size distribution. Significant spatial clustering of abundant overstory species was observed at all spatial scales examined. Spatial distributions of A. rubrum and Q. rubra showed negative intraspecific correlations in diameters up to at least a 150-m spatial lag, likely indicative of crowding effects in dense forest patches following intensive past land use. Bivariate marked point-pattern analysis, showed that T. canadensis and Q. rubra diameters were negatively associated with one another, indicating resource competition for light. Distribution and abundance of the common overstory species are predicted best by soil type, tree neighborhood effects, and two aspects of land-use history: when fields were abandoned in the late 19th century and the succeeding forest types recorded in 1908. In contrast, a history of intensive logging prior to 1950 and a damaging hurricane in 1938 appear to have had little effect on the distribution and abundance of present-day tree species. Our findings suggest that current day composition and structure are still being influenced by anthropogenic disturbances that occurred over a century ago.« less

Abstract Background and Aims Terrestrial laser scanners (TLSs) have successfully captured various properties of individual trees and have potential to further increase the quality and efficiency of forest surveys. However, TLSs are limited to line of sight observations, and forests are complex structural environments that can occlude TLS beams and thereby cause incomplete TLS samples. We evaluate the prevalence and sources of occlusion that limit line of sight to forest stems for TLS scans, assess the impacts of TLS sample incompleteness, and evaluate sampling strategies and data analysis techniques aimed at improving sample quality and representativeness. Methods We use a large number of TLS scans (761), taken across a 255 650-m2 area of forest with detailed field survey data: the Harvard Forest Global Earth Observatory (ForestGEO) (MA, USA). Sets of TLS returns are matched to stem positions in the field surveys to derive TLS-observed stem sets, which are compared with two additional stem sets derived solely from the field survey data: a set of stems within a fixed range from the TLS and a set of stems based on 2-D modelling of line of sight. Stem counts and densities are compared between the stem sets, and four alternative derivations ofmore »area to correct stem densities for the effects of occlusion are evaluated. Representation of diameter at breast height and species, drawn from the field survey data, are also compared between the stem sets. Key Results Occlusion from non-stem sources was the major influence on TLS line of sight. Transect and point TLS samples demonstrated better representativeness of some stem properties than did plots. Deriving sampled area from TLS scans improved estimates of stem density. Conclusions TLS sampling efforts should consider alternative sampling strategies and move towards in-progress assessment of sample quality and dynamic adaptation of sampling.« less

Forest insects and pathogens have significant impacts on U.S. forests, annually affecting an area nearly three times that of wildfires and timber harvesting combined. However, coupled with these direct effects of forest insects and pathogens are the indirect impacts through influencing forest management practices, such as harvesting. In an earlier study, we surveyed private woodland owners in the northeastern U.S. and 84% of respondents indicated they intended to harvest in at least one of the presented insect invasion scenarios. This harvest response to insects represents a potentially significant shift in the timing, extent, and species selection of harvesting. Here we used the results from the landowner survey, regional forest inventory data, and characteristics of the emerald ash borer (Species: Agrilus planipennis Fairmaire, 1888) invasion to examine the potential for a rapidly spreading invasive insect to alter harvest regimes and affect regional forest conditions. Our analysis suggests that 25% of the woodland parcels in the Connecticut River Watershed in New England may intend to harvest in response to emerald ash borer. If the emerald ash borer continues to spread at its current rate within the region, and therefore the associated management response occurs in the next decade, this could result inmore »an increase in harvest frequencies, from 2.6% year−1 (historically) to 3.7% year−1 through to approximately 2030. If harvest intensities remain at levels found in remeasured Forest Inventory and Analysis plots, this insect-initiated harvesting would result in the removal of 12%–13% of the total aboveground biomass. Eighty-one percent of the removed biomass would be from species other than ash, creating a forest disturbance that is over twice the magnitude than that created by emerald ash borer alone, with the most valuable co-occurring species most vulnerable to biomass loss.« less

The ability to automatically delineate individual tree crowns using remote sensing data opens the possibility to collect detailed tree information over large geographic regions. While individual tree crown delineation (ITCD) methods have proven successful in conifer-dominated forests using Light Detection and Ranging (LiDAR) data, it remains unclear how well these methods can be applied in deciduous broadleaf-dominated forests. We applied five automated LiDAR-based ITCD methods across fifteen plots ranging from conifer- to broadleaf-dominated forest stands at Harvard Forest in Petersham, MA, USA, and assessed accuracy against manual delineation of crowns from unmanned aerial vehicle (UAV) imagery. We then identified tree- and plot-level factors influencing the success of automated delineation techniques. There was relatively little difference in accuracy between automated crown delineation methods (51–59% aggregated plot accuracy) and, despite parameter tuning, none of the methods produced high accuracy across all plots (27—90% range in plot-level accuracy). The accuracy of all methods was significantly higher with increased plot conifer fraction, and individual conifer trees were identified with higher accuracy (mean 64%) than broadleaf trees (42%) across methods. Further, while tree-level factors (e.g., diameter at breast height, height and crown area) strongly influenced the success of crown delineations, the influence of plot-level factorsmore »varied. The most important plot-level factor was species evenness, a metric of relative species abundance that is related to both conifer fraction and the degree to which trees can fill canopy space. As species evenness decreased (e.g., high conifer fraction and less efficient filling of canopy space), the probability of successful delineation increased. Overall, our work suggests that the tested LiDAR-based ITCD methods perform equally well in a mixed temperate forest, but that delineation success is driven by forest characteristics like functional group, tree size, diversity, and crown architecture. While LiDAR-based ITCD methods are well suited for stands with distinct canopy structure, we suggest that future work explore the integration of phenology and spectral characteristics with existing LiDAR as an approach to improve crown delineation in broadleaf-dominated stands.« less