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Hemlock Woolly Adelgid in New England Forests: Canopy Impacts Transforming Ecosystem Processes and Landscapes

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Abstract

Exotic insect pests may strongly disrupt forest ecosystems and trigger major shifts in nutrient cycling, structure, and composition. We examined the relationship between these diverse effects for the hemlock woolly adelgid (HWA, Adelges tsugae Annand) in New England forests by studying its impacts on local canopy processes in stands differing in infestation levels and linking these impacts to shifts in canopy nutrient cycling and stand and landscape effects. HWA initiated major changes in canopy biomass and distribution. Whereas uninfested trees exhibit a significant decline in canopy biomass from the center to the periphery and a positive correlation between total needle litter and estimated biomass, infested trees have significantly less total canopy biomass, produce less new foliage, shed relatively more needles, and exhibit no correlation between litter and canopy biomass. Foliar N content of infested trees was 20%–40% higher than reference trees, with the strongest increase in young foliage supporting the highest densities of HWA. Foliar %C was unaffected by HWA or foliar age. Epiphytic microorganisms on hemlock needles exhibited little variation in abundance within canopies, but colony-forming units of bacteria, yeast, and filamentous fungi were 2–3 orders of magnitude more abundant on medium and heavily infested than uninfested trees. Throughfall chemistry, quantity, and spatial pattern were strongly altered by HWA. Throughfall exhibits a strong gradient beneath uninfested trees, decreasing in volumes from the canopy periphery to the trunk by more than 45%. The amount of throughfall beneath infested trees exhibits no spatial pattern, reaches 80%–90% of the bulk precipitation, and is characterized by significantly higher concentrations of nitrogen compounds, dissolved organic carbon, and cations.

Across the southern New England landscape there is a strong south-to-north gradient of decreasing hemlock tree and sapling mortality and understory compositional change that corresponds to the duration of infestation. Regionally, black birch (Betula lenta L.) is profiting most from hemlock decline by significantly increasing in density and cover. These findings suggest that it is necessary to study the connections between fast/small-scale processes such as changes in nutrient cycling in tree canopies and slow/integrative processes like shifts in biogeochemieal cycling and compositional changes at forest stands and landscapes to better understand the effects of an exotic pest species like HWA on forest ecosystem structure and function.

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Acknowledgments

We thank John Aber and David Foster for providing valuable comments on a previous version of the manuscript and two anonymous reviewers for constructive criticism. Bettina Popp and Petra Dietrich helped with the chemical analyses. Generous access to field sites was provided by the The Nature Conservancy and the Connecticut Department of Environmental Protection. Financial support was given from the German Ministry for Research and Technology (Fördernummer: BMBF No. PT BEO 51-0339476D) and the Harvard Forest Long Term Ecological Research program (NSF 94-11764). BS was supported at the Harvard Forest by a Charles Bullard Fellowship from Harvard University.

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Authors and Affiliations

  1. Bayreuth Institute for Terrestrial Ecosystem Research, University of Bayreuth, Dr. Hans-Frischstr. 1−3, 95440, Bayreuth, Germany

    Bernhard Stadler

  2. Centre for Agricultural Landscape and Land Use Research Müncheberg, Institute of Primary Production and Microbial Ecology, Gutshof 7, Paulinenaue, D-14641, Germany

    Thomas Müller

  3. Harvard Forest, Harvard University, Petersham, Massachusetts, 01366, USA

    Bernhard Stadler, David Orwig & Richard Cobb

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  1. Bernhard Stadler
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Correspondence to Bernhard Stadler.

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Stadler, B., Müller, T., Orwig, D. et al. Hemlock Woolly Adelgid in New England Forests: Canopy Impacts Transforming Ecosystem Processes and Landscapes. Ecosystems 8, 233–247 (2005). https://doi.org/10.1007/s10021-003-0092-5

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