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Ants and Ecosystem Function in Hemlock Removal Experiment at Harvard Forest 2006-2014
Eastern hemlock (Tsuga canadensis) is a foundation species in eastern North American forests. Hemlock stands host unique assemblages of flora and fauna, and the structure of these assemblages is expected to change as hemlock declines due to infestation by the hemlock woolly adelgid (Adelges tsugae) and is removed by pre-emptive salvage logging. A 2003 survey of hemlock stands in central Connecticut and Massachusetts showed that ant species richness and abundance is substantially greater in logged and adelgid-infested hemlock stands than it is in intact hemlock stands (see HF065) and we hypothesized that we would see similar changes in ant species diversity following application of treatments in the Hemlock Removal Experiment at the Simes Tract (see HF118). We further hypothesized that because ants are known to modulate ecosystem function in other habitats that observed changes in ant species diversity would be accompanied by changes in ecosystem processes including soil respiration rate and nitrogen availability. In April 2006, we established a set of sub-plots in all eight canopy manipulation plots of the Hemlock Removal Experiment. In each canopy manipulation plot, we installed two ant exclosure plots, two disturbance control plots, and two control plots. Ant species richness and abundance, as well as effectiveness of the exclosures, is monitored with pitfall traps monthly during the summer. Soil nitrogen availability is measured using resins that are collected every three months, and soil respiration is measured bi-weekly during the growing season.
Net Carbon Exchange of an Old-Growth Hemlock Forest at Harvard Forest HEM Tower since 2000
This project estimates carbon exchange rates of multiple forest types at Harvard Forest (see HF072) and compares them to long-term ongoing carbon exchange measurements at the EMS, which is located in a mesic, 60-90 year old red oak and red maple dominated forest on abandoned farmland (HF004). Measurements in each forest type are used to investigate climatic influences on carbon exchange. This mesic hemlock-dominated forest with most trees 100-200 years old on undisturbed soils stored only about 3 Mg/ha of carbon in 2001, compared to over 4 Mg/ha in the 60-90 year old oak/maple stand. However, both sites stored more carbon in 2001 than was measured in the oak/maple stand in any previous year since 1991 (see HF004). The hemlock forest behaved very differently from the oak-maple stand in that the highest rates of carbon storage occurred in spring, while there was very little carbon storage in mid to late summer. Statistical models of carbon exchange in the hemlock forest showed that carbon storage was positively related to daily minimum air temperature in spring, but negatively correlated with soil temperature in the summer. The first effect was attributable to a positive influence of above-freezing minimum temperatures on photosynthesis by hemlock foliage. The negative relationship of soil temperature to carbon storage by hemlock forest in summer was due to exponentially increasing soil and ecosystem respiration, accompanied by a neutral or negative effect of high air temperature on photosynthesis by hemlock trees (see HF063). These effects indicate that carbon storage in the hemlock forest could be strongly affected by climate warming, but the effects will probably be in opposite directions in spring and summer.
Adult Population Survey of Hemlock Woolly Adelgid in Massachusetts 2003-2005
To effectively document insect presence and abundance a statistically based sampling plan is necessary. No such plan for the hemlock woolly adelgid is currently available in the published literature. Two interrelated sampling strategies allow a researcher to enter a site and determine with a given level of probability whether or not an HWA infestation is present and the level of infestation. A ground based sampling plan is also critical to the ‘ground truthing’ of aerial survey systems. A fully developed sampling plan requires relatively extensive datasets obtained over time, and may include the use of computer-generated data obtained through re-sampling techniques. The results obtained through the Harvard Forest Summer Research Program should provide a firm foundation for further development of an HWA sampling program. Data collection is on-going.
Seed Bank in Hemlock Removal Experiment at Harvard Forest 2004-2015
The impending loss of hemlock trees due to hemlock woolly adelgid (Adelges tsugae) infestation prompted the need to identify species of plants able to colonize areas where hemlock has been removed. We investigated species present in the seed bank (in 2004 and 2010), the seedling bank (in 2004 and 2010), and seed rain (from 2005-2015) in the Harvard Forest Hemlock Removal Experiment. In 2004, Kelley Sullivan and Aaron Ellison determined the distribution of seeds buried in the soil in the six hemlock and two hardwood stands, before canopy treatments were applied (data files hf105-01, hf105-02). In 2010, Elizabeth Farnsworth and Aaron Ellison examined the distribution of plant species present five years after the simulated hemlock removal experiment was performed and six years after the baseline, pre-treatment study of the seedbank (data files hf105-03, hf105-04, hf105-05). In the 2010 study, we hypothesized that the composition of the seed and seedling banks and seed rain would diverge among the two treatments and the controls, based on the differential impacts of harvesting versus adelgid attack on the standing vegetation. We further hypothesized that the older (deeper) strata of the seedbanks of the treatment plots would have been exhausted over time, thus yielding poor germination relative to the top strata that continually receive seed rain. Identical methods of characterizing the composition of the seedbank and the aboveground vegetation at seedbank core locations were applied in both 2004 and 2010. Additional comparisons were made with long-term data collected on overall plant species composition in the Simes plots (datasets HF 106, HF126), and with data on seed rain (data file hf105-05). This study provides a unique temporal documentation of changes in the seedbank, seed rain, and vegetation under conditions of a changing overstory.
Understory Vegetation in Hemlock Removal Experiment at Harvard Forest since 2003
Hemlock decline in New England is caused by direct and indirect effects of invasion of the hemlock woolly adelgid. Direct damage from the insect is causing gradual mortality of hemlock, and widespread harvesting of hemlock in advance of mortality, in contrast, causes immediate mortality and removal of biomass from the site. Although both processes affect thousands of acres of forest annually we have only a limited understanding of their effects on forest ecosystem function and productivity and the nature of the subsequent forest community. We anticipate that harvesting will yield different consequences than gradual mortality from the insect. Therefore we designed an experiment to simulate these contrasting impacts, by logging or girdling hemlock stands. Results from the experimental treatments are compared to the changes observed in forests that are being infested by the adelgid, and can also be included in integrated analyses of a suite of large experiments that form a core component of the Harvard Forest LTER program.
Light Environment in Hemlock Removal Experiment at Harvard Forest since 2003
The impending loss of hemlock trees due to hemlock woolly adelgid (Adelges tsugae) infestation is expected to lead to changes in the light environment of the forest understory. These changes will both drive succession and will themselves be altered by successional processes. The light reaching the forest floor is measured using hemispherical canopy photographs. Photographs were taken in summer of 2003 and in December 2004 and May 2005 (both deciduous-tree leaf-off condition), prior to the application of the logging and girdling treatments, and then in September 2005 (deciduous tree leaf-on condition) after logging and girdling. Subsequent photographic series will be taken annually in spring (leaf-off) and summer/fall (leaf-on).
Air and Soil Temperature in Hemlock Removal Experiment at Harvard Forest since 2004
The impending loss of hemlock trees due to hemlock woolly adelgid (Adelges tsugae) infestation is expected to lead to changes in the temperature regime of the forest understory. These changes will both drive succession and will themselves be altered by successional processes. Air temperature (1 m above forest floor) and soil temperature (at 10 cm depth) are measured at 1-minute intervals (hourly averages, minimum, and maximum are stored) using thermistors connected to Campbell 21-X dataloggers. Temperature measurements began 60-120 days prior to applications of the logging and girdling treatments, and will continue for the duration of the study.
Ant Diversity and Vegetation Composition in Hemlock Removal Experiment at Harvard Forest 2006
Ants comprise a considerable amount of animal biomass in terrestrial ecosystems and play major roles in ecological processes ranging from seed dispersal to soil turnover. Invasion by the hemlock woolly adelgid will transform late-successional hemlock forests into earlier successional mixed hardwood-white pine forests or red-maple wetlands. Understanding how ant assemblages vary in different habitat types allows for predictions of how hemlock decline could alter the composition of ant assemblages, with implications for a wide range of ecosystem processes. An ongoing study at the Simes Tract of Harvard Forest is documenting the effects of invasion and land-use history on ant biodiversity. Surveys from 2003 to 2005 focused on ant structure in hemlock and hardwood microhabitats in the Harvard Forest Hemlock Removal Experiment, in which hemlock forest response to deforestation by the hemlock woolly adelgid (Adelges tsugae) and to selective logging is being examined (Ellison et al. 2005). In the summer of 2006, we surveyed a greater range of microhabitat types with two objectives. First, to collect rare or elusive species in hemlock and hardwood stands that may have gone uncollected in previous years. Second, to sample forest communities not included in previous years - white pine, swamp, and rocky slope - for ant species unique to these microhabitats. We found fourteen newly documented species of ants in the Simes Tract - nine of which were in an open, swamp. Aphaenogaster rudis and Camponotus pennsylvanicus were the only ant species found in all microhabitat types. In a canonical correspondence analysis, A. rudis and C. pennsylvanicus were associated most strongly with hemlock stands and low species richness of understory plants.
Deer and Moose Browsing in Hemlock Removal Experiment at Harvard Forest 2008
Hemlock decline in New England is caused by direct and indirect effects of invasion of the hemlock woolly adelgid. Direct damage from the insect is causing gradual mortality of hemlock, and widespread harvesting of hemlock in advance of mortality, in contrast, causes immediate mortality and removal of biomass from the site. Although both processes affect thousands of acres of forest annually we have only a limited understanding of their effects on forest ecosystem function and productivity and the nature of the subsequent forest community. We anticipate that harvesting will yield different consequences than gradual mortality from the insect. Therefore we designed an experiment to simulate these contrasting impacts, by logging or girdling hemlock stands. Results from the experimental treatments will be compared to the changes observed in forests that are being infested by the adelgid, and can also be included in integrated analyses of a suite of large experiments that form a core component of the Harvard Forest LTER program. Deer and moose foraging can play a key role in shaping forest regeneration after disturbance in temperate forest. In 2008, we initiated a browsing survey of woody stems in the Simes hemlock removal experiment plots. There are regular moose sightings in the study area, and moose pellets are commonly found within the plots. Also, extensive browsing of tree regeneration in the logged plots was noted starting in 2007. Sampling of the 8 study plots was completed in Summer 2008. The next sampling is tentatively scheduled for Summer 2013.
Coarse Woody Debris in Hemlock Removal Experiment at Harvard Forest since 2005
The woody detritus survey is designed to measure coarse woody detritus that includes snags, logs, and stumps, and to estimate fine woody detritus which includes smaller pieces of downed wood. To capture both standing and downed wood, we based our survey around two main types of methods, the line intercept method and the fixed radius plot method. Surveys have been completed for 2005, 2007, 2009, 2011, 2013, 2015, 2017, and 2021.
Overstory Vegetation in Hemlock Removal Experiment at Harvard Forest since 2003
Hemlock decline in New England is caused by direct and indirect effects of invasion of the hemlock woolly adelgid. Direct damage from the insect is causing gradual mortality of hemlock. Widespread harvesting of hemlock in advance of mortality, in contrast, causes immediate mortality and removal of biomass from the site. Although both processes affect thousands of acres of forest annually we have only a limited understanding of their effects on forest ecosystem function and productivity and the nature of the subsequent forest community. We anticipate that harvesting will yield different consequences than gradual mortality from the insect. Therefore, we designed an experiment to simulate these contrasting impacts, by logging or girdling hemlock stands. Results from the experimental treatments will be compared to the changes observed in forests that are being infested by adelgid and can also be included in integrated analyses of a suite of large experiments that form a core component of the Harvard Forest LTER program.
Impacts of Hemlock Harvesting in Central Massachusetts 2003-2009
The recent unimpeded infestation of the hemlock woolly adelgid (HWA; Adelges tsugae) across the northeastern U.S. is driving large-scale hemlock decline and mortality. HWA has already infested over 40% of the towns in Massachusetts and, as a result, many landowners are considering pre-emptively harvesting their eastern hemlock (Tsuga canadensis) stands. To examine the local impacts of this cutting activity, we examined stand and ecosystem dynamics in 10 hemlock-dominated sites on public and private lands where hemlocks were harvested between 1 to12 years ago with nearby uncut forests. Forty-two to ninety-seven percent of hemlock stems and basal area were removed from the harvested sites. Sapling densities of 10,500 to 24,000 stems ha-1 dominated the vegetation at older cuts and consisted primarily of black birch (Betula lenta), red maple (Acer rubrum), and white pine (Pinus strobus). Seedling densities averaged 5 m-2 across sites, peaked at 20 m-2 at 6 to 9 year old sites, and consisted of hemlock, white pine, black birch, and red maple. Additional understory species that were common in cuts sites included various raspberry (Rubus) species, often averaging 30% cover, hay-scented fern (Dennstaedtia punctilobula), and various sedge (Carex) species. Harvesting resulted in soils that were 3 to 5 (deg C) warmer and tended to be drier in recent vs. older cuts. Net nitrogen mineralization rates in organic soils averaged just over 10 kg N ha-1yr-1 in recent cuts and approx. 5 kg N ha-1yr-1 in older cuts. Mineral soil net mineralization rates were lower among all cuts, ranging from 2.1 to 3.5 kg N ha-1yr-1. Nitrification rates were low in both soil horizons at all harvest ages. Recent cuts had much higher nitrogen capture (NH4- + NO3+) on resin bags averaging around 1200 μg N g resin-1 vs. 300 to 600 μg N g resin-1 captured in older cuts. Nitrification rates and nitrate capture on resin bags were lower in Massachusetts sites than those reported on sites with a long history o
Detection Histories for Hemlock Woolly Adelgid Infestations at Cadwell Forest in Pelham MA 2008
Monitoring programs increasingly are used to document the spread of invasive species in the hope of detecting and eradicating low-density infestations before they become established. However, interobserver variation in the detection and correct identification of low-density populations of invasive species remains largely unexplored. In this study, we compare the abilities of volunteer and experienced individuals to detect low-density populations of an actively spreading invasive species and we explore how interobserver variation can bias estimates of the proportion of sites infested derived from occupancy models that allow for both false negative and false positive (misclassification) errors. We found that experienced individuals detected small infestations at sites where volunteers failed to find infestations. However, occupancy models erroneously suggested that experienced observers had a higher probability of falsely detecting the species as present than did volunteers. This unexpected finding is an artifact of the modeling framework and results from a failure of volunteers to detect low-density infestations rather than from false positive errors by experienced observers. Our findings reveal a potential issue with site occupancy models that can arise when volunteer and experienced observers are used together in surveys.
Tree Growth in Hemlock and Deciduous Forests at Harvard Forest HEM and LPH Towers 2000-2005
Tree growth was measured to determine tree ages and growth rates and to quantify carbon storage in these forests, and to detect changes in growth and carbon storage that could be associated with climate changes or historic disturbances.
Litterfall in Hemlock Removal Experiment at Harvard Forest since 2005
Hemlock decline in New England is caused by direct and indirect effects of invasion of the hemlock woolly adelgid. Direct damage from the insect is causing gradual mortality of hemlock, and widespread harvesting of hemlock in advance of mortality, in contrast, causes immediate mortality and removal of biomass from the site. Although both processes affect thousands of acres of forest annually we have only a limited understanding of their effects on forest ecosystem function and productivity and the nature of the subsequent forest community. We anticipate that harvesting will yield different consequences than gradual mortality from the insect. Therefore we designed an experiment to simulate these contrasting impacts, by logging or girdling hemlock stands. Results from the experimental treatments will be compared to the changes observed in forests that are being infested by the adelgid, and can also be included in integrated analyses of a suite of large experiments that form a core component of the Harvard Forest LTER program.
Soil Arthropods in Hemlock Removal Experiment at Harvard Forest 2008
In eastern North American forests, eastern hemlock (Tsuga canadensis) is a foundation species. As hemlock is lost from forests due to the invasive hemlock woolly adelgid (Adelges tsugae) and pre-emptive salvage logging, the structure of assemblages of species associated with hemlock is expected to change. We manipulated hemlock canopy structure at hectare scales to investigate the effects of hemlock death on assemblages of ants, beetles, and spiders in a New England forest. Relative to reference hemlock stands, both in situ death of hemlock and logging and removal of hemlock altered composition and diversity of beetles and spiders, and logging increased the species richness and evenness of ant assemblages. Species composition of ant assemblages in disturbed habitats was non-random relative to the regional species pool, but we found no evidence that interspecific competition shaped the structure of ant, beetle or spider assemblages, in either manipulated or intact forest stands. Environmental filtering by hemlock appears to maintain low levels of species richness and evenness in forest stands, suggesting that the loss of hemlock due to the hemlock woolly adelgid or human activities will not likely lead to extirpations of ant, beetle, or spider species at local scales.
Modeling Range Expansion of Hemlock Woolly Adelgid in Eastern North America 1951-2009
Range expansion by native and non-native species will continue to be a major component of global change. Anticipating the potential effects of changes in species distributions requires models capable of forecasting population spread across realistic, heterogeneous landscapes and subject to spatiotemporal variability in habitat suitability. Several decades of theory and model development, as well as increased computing power and availability of fine-resolution GIS data, now make such models possible. This R code allows simulation of the spread of the hemlock woolly adelgid (HWA, Adelges tsugae) under climatic conditions experienced from December of 1951 until March of 2009. The code describes a spatially explicit stochastic model that combines dynamic dispersal and population processes with fine-resolution maps characterizing spatiotemporal heterogeneity in winter temperature and hemlock abundance to model range expansion of HWA. The model is parameterized using multi-year datasets describing population and dispersal dynamics of HWA and is applied to eastern North America.
Regional Distribution and Abundance of Eastern Hemlock in Eastern North America 2010
We developed comprehensive maps on the distribution and abundance of hemlock for the purposes of mapping the host distribution, and modeling the spread, of the hemlock woolly adelgid. Multiple statistical models were used to map the distribution of hemlock. Hemlock occurrence data were taken from the FIA database and multiple environmental predictors were gathered from various databases as described in methods. The raster map depicts the predicted abundance of hemlock m2 basal area per hectare) across its range in eastern North America.
Simulating Intrastand Movement of Hemlock Woolly Adelgid at Harvard Forest 2009
The hemlock woolly adelgid (HWA), Adelges tsugae Annand (Hemiptera: Adelgidae), has spread rapidly across the eastern USA since its introduction from Japan 60 years ago, causing widespread mortality of both eastern hemlock [Tsuga canadensis (L.) Carriere] and Carolina hemlock [Tsuga caroliniana Engelm. (Pinaceae)]. Although HWA spread patterns have been repeatedly analyzed at regional scales, comparatively little is known about its dispersal potential within and between hemlock stands. As the small size and clonal nature of HWA make it nearly impossible to identify the source populations of dispersing individuals, we simulated intrastand HWA movement in the field by monitoring the movement of clumps of fluorescent powder that are slightly larger than HWA, but much easier to detect in the forest understory. Using three hemlock trees with three colors of fluorescent powder as source populations, we detected dispersal events at the farthest distances within our trapping array (400 m). However, more than 90% of dispersal events were less than 25 m. Dispersal patterns were similar from all three source trees and the distribution of dispersal distances in all cases could be described by lognormal probability density functions with mean dispersal distance of 12-14 m, suggesting that dispersal was relatively independent of location of source trees. In general, we documented tens of thousands of passive dispersal events in the forest understory despite the presence of a dense forest canopy. Thus, even under relatively light-wind conditions, particles of similar dimensions to HWA are capable of intra-stand movement, suggesting that a large population of HWA could rapidly infest other trees within several hundred meter radius, or beyond.
Ungulate-Disturbance Interactions in Hemlock Ecosystems at Harvard Forest 2012-2013
Densities of ungulates are often associated with recent forest disturbances such as fire, logging and insect outbreaks, as increased resources stimulate tree regeneration, leading to abundant available browse. Despite the often significant role that ungulates play in disturbed forests, surprisingly little is known about ungulate-disturbance interactions, and ungulate herbivory is often excluded from examinations of forest response to disturbance. In the Northeastern United States, a large-scale insect outbreak, the hemlock wooly adelgid (HWA), has begun to have important effects on hemlock forests both directly and indirectly (i.e., by preemptive salvage logging). No studies have examined the interactions of both moose and deer activity with these associated canopy disturbances. Our objectives are twofold: to determine (1) the response of ungulates to varying disturbance types and intensities and (2) the influence of ungulate interactions with canopy disturbance on vegetation, other biota, and ecosystem processes. We will examine the relative abundance of moose and deer in four treatments (2 replicates) at the Hemlock Removal Experiment using three indices of ungulate activity: pellet group density, evidence of past browsing on seedlings and saplings, and the occurrence of animals using game cameras. We will also monitor the response of vegetation to ungulate browsing by sampling woody and herbaceous vegetation in fenced exclosures and paired controls in disturbed and undisturbed plots.
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