Presentation of the fall and rise and fall again of eastern hemlock, a foundation tree species of eastern North American forests. Testing the hypothesis that it is a foundation species, modeling its future given climate change scenarios, and validating model results with eddy covariance data
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Things fall apart
1. THINGS FALL APART
Land-use history, non-native insects, climatic change, and
the decline of a forest foundation species
Aaron M. Ellison
Harvard University, Harvard Forest
2.
3. Frontiers in Ecology and the Environment (2005)
Benjamin Baiser
Audrey Barker Plotkin
Peter Bettman-Kerson
Hannah Buckley
Brad Carlin
Brad Case
Aimeé Classen
Eric Davidson
Ally DeGrassi
Joe Elkinton
Elizabeth Farnsworth
Matthew Fitzpatrick
David Foster
Joe Kendrick
Michael Lavine
Nicky Lustenhouwer
Heidi Lux
Liza Nicoll
Dave Orwig
Annie Paradis
Mainsha Patel
Adam Porter
Evan Preisser
Sydne Record
Tara Sackett
Kathleen Savage
Jonathan Thompson
Mark Van Scoy
Lance Waller
Collaborators
5. Yale University Press (2014)
The way a crow
Shook down on me
The dust of snow
From a hemlock tree
Has given my heart
A change of mood
And saved some part
Of a day I had rued.
― Robert Frost
6. Flux(mgCm-2hr-1)
0
100
200
300
400
SoilTemperature(oC)
at10cm
0
5
10
15
20
25
VolumetricSoilMoisture
(gH2Ocm
-3
soil)
0.0
0.2
0.4
0.6
1995 1996 1997 1998 1999 2000
Soil respiration, temperature, and water content in a hardwood stand, a young hemlock
stand, and the old hemlock stand at Harvard Forest Unpublished data of E.A.
Davidson, S.E. Trumbore, and J.L. Hadley
[Data: HF072, HF103, HF153]
Hemlock basal area (percent of total)
0 20 40 60 80 100
Numberofantspecies
2
4
6
8
10
12
14
CT sites
MA sites
r2
= 0.50, P = 0.002
Hemlock basal area (percent of total)
0 20 40 60 80 100
IncidenceofFormicaspp.
(probabilityofoccurrence)
0.0
0.2
0.4
0.6
0.8
1.0 P = 0.045
Ellison, Chen, Díaz, Kammerer-Burnham, and Lau,
Proc. 3rd Symp. HWA in eastern N. Am. (2005).
[Data: HF065]
7. Kathy Shields USDA Forest Service
Photos: David Orwig/Harvard Forest
SEM: USFS
10. Long-term research questions
• What is the process by which the system reorganizes following loss
of hemlock, and how is it related to the biology of hemlock and the
adelgid?
• What new endpoints will the system reach following this
reorganization?
• How does logging vs. the adelgid alter these transitions and
endpoints?
• Has this happened before, and what can we learn from history?
11. Research approach
Retrospective studies Decades to centuries
Patterns of insect dis-
persal and logging
Years to decades
Whole-tree physiology Days to years
Insect physiology and
effect on trees
Days to years
Soil ecosystem pro-
cesses
Months to decades
Atmospheric CO2 and
H2O flux
Instants to years
Forest structure and
environmental varia-
bles
Months to years
Structure of associat-
ed biota
Years to decades
ED and PnET models Decades to centuries
12. Paleo-hemlock decline
Raw data interpolated to 50-year time intervals by Julia Jones ; LOI data and pollen data
courtesy of Ed Faison and Wyatt Oswald; GISP2 data from http://www.gisp2.sr.unh.edu/
[Data: HF184, 185, 186]
13. Post-agriculture hemlock recovery
• Hemlock has steadily increased
in abundance in the last
century, especially following the
loss of American chestnut
(Castanea dentata), another
forest dominant (foundation
species?) due to chestnut
blight, and after the 1938
hurricane.
• In the last 100 years, hemlock
has not been systematically
logged.
• The adelgid arrived in the
northeast ~1980.
20. • Eight 90 × 90 m (0.81
ha) plots established
2003 (Audrey Barker Plotkin,
Aaron Ellison, David Foster,
Dave Orwig, Eric Davidson)
• Blocked BACI design
with 4 treatments/block:
• Hemlock control
• Girdle all hemlocks
• “Commercial” cut of
hemlocks and other
merchantable timber
• Hardwood control
• 2 years pre-treatment
data
• Treatments applied
winter/spring 2005
Project description: Ellison et al. (2010) Methods in Ecology & Evolution
21. Ellison et al. (2014) Rhodora; [Data: HF000, HF001, HF086, HF126, HFDA 1935-01, 1944-10, 1950-17, HFDA 1980-13, 1933-05, HFAdm 137, HF Stand Records Simes Tract, MASS-GIS]
23. • All trees tagged, measured, and mapped
• Stand reconstruction (deed research and
dendrochronology)
• Sap flow during girdling
• Seed bank composition
• Seed dispersal and recruitment
• Understory vegetation composition
• Animal diversity and abundance
• Ants
• Beetles
• Amphibians
• Birds
• Mammals: from rodents to moose
• Adelgid colonization
Ongoing projects
28. Vegetation structure and
compositional change driven by seed
bank, seed rain, and understory
responses.
Farnsworth et al. (2012) Canadian Journal of Forest Research
[Data: HF105, HF106, HF120]
35. Historic data from VEMAP2 grid cell elements of New England. Projections using Canadian and
Hadley GCMs; published by New England Regional Assessment Group (2001) Preparing for a
changing climate: the potential consequences of climate variability and change. New England
regional overview. University of New Hampshire and U.S. Global Change Research Program.
Regional climate
models project
warmer and wetter
conditions in the
northeast USA
36. 1850
1997
USDA, US Census and FAO data
and projections
Historical and contemporary land-use
Model: Albani et al. 2010 Canadian Journal of Forest Research 40: 119
37. Integrated model of adelgid spread and impact
Rate of hemlock decline from Dave Orwig
AFLP data from Annie Paradis and Adam Porter
Average year
of HWA
infestation
from
stochastic
model
Albani et al. (2010) Canadian Journal of Forest Research
38. Modeled impact of HWA infestation at the site scale on above-ground biomass trajectories
(left) for grasses (G), early-, mid- (MSD), and late- (LSD) successional deciduous trees,
pines (NP, SP), hemlock (LSE); and carbon fluxes (right: NPP and heterotrophic respiration)
for central Massachusetts (top) and central New Hampshire (bottom).
Model: Albani et al. (2010) Canadian Journal of Forest Research
39. Impact of the adelgid
on net ecosystem
productivity (NEP),
expressed as difference
between the infestation
and the no infestation
scenario. Note the
characteristic pattern of
decreased and then
increased NEP as the
infestation moves from
east to west.
Model: Albani et al. (2010) Canadian Journal of Forest Research
41. Turning and turning in the widening gyre
The falcon cannot hear the falconer;
Things fall apart; the centre cannot hold;
Mere anarchy is loosed upon the world,
The blood-dimmed tide is loosed, and everywhere
The ceremony of innocence is drowned;
The best lack all conviction, while the worst
Are full of passionate intensity.
—from The Second Coming by W. B. Yeats (1920)