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Postglacial
dynamics of
Olympic Peninsula
forests: comparing
simulations and
observations
Thanks to
Dan Gavin
Bart Bartlein
Jed Kaplan

A Master’s Thesis
presented to the

Department of Geography
University of Oregon
by

David Fisher
Olympic Mountains, WA
Research Questions

•

How does knowledge of the life history of tree species -- regeneration,
growth, competition, mortality -- as well as the relationship between
species and climate at important life stages, improve our explanation of
the dynamic changes in forest composition since the Late Glacial period,
15,000 years ago?
Outline of this talk

• Paleoecology & Biogeography of Olympic Peninsula
forests

• Methods -- quantifying important bioclimatic

variables and simulating paleoecological records with
a process-based vegetation model

• Results -- Simulations compared to present and
paleo records

• What did we learn?
Paleoecological Records
Cold, Pollen Percentage Diagram
Yahoo Lake -Dry? Warm, Dry Cool, Wet

Figure by D. G. Gavin

(Gavin, D.G. 2013)
Limiting factors for tree growth:

On the west side: energy
On the east side: moisture

sretemoliK 08

04

02

0

¯
50km

PRISM
(Daly et al. 2008)

4
3
2

Potential
Natural
Forest Zones

¯

-4

0

-2

02

0

6 meters

04

PRISM
(Daly et al. 2008)

2 deg C

Mean Annual Precip

sretemoliK 08

Minimum Jan Temp

(Henderson et al. 2011)

Alpine
mountain hemlock
subalpine fir
Pacific silver fir
Douglas-fir
western hemlock
Sitka spruce
mountain hemlock
Tsuga mertensiana

western redcedar
Thuja plicata

subalpine fir
Abies lasciocarpa

western hemlock
Tsuga heterophylla

lodgepole pine
Pinus contorta

Sitka spruce
Picea sitchensis

Pacific silver fir
Abies amabilis

red alder
Alnus rubra

Douglas-fir

Pseudotsuga menziesii

grass
Methods: Considering the seedling, how should we measure
energy and moisture requirements?
Methods: Considering the seedlings, how do we measure
energy?
Growing Degree Days

Average Daily
Temperature
(Deg C)

Post-Snow
Growing Degree Days

15

2000

Size of Snowpack
(Snow-Water equivalent)
(mm)

10

1000

5
-5

J F M A M J J A S O N D

0
Post-Snow GDD Measurements
Local Measurements of Post-Snow GDD
(two SNOTEL stations)

Year

2006
2007
2008
2009
2010
2011
2012

SW

487
373
650

NE

649
432
493
670
473
384
563

SW = Buckinghorse
NE = Waterhole
http://www.wcc.nrcs.usda.gov/snotel/SNOTEL-brochure.pdf
Post-Snow GDD Simulations

annual PS-GDD required
for each species to establish

# Post-Snow
GDDs
1600
1400

S. spruce
w. redcedar w. hemlock
r. alder

800

Doug-fir

300

m. hemlock l. pine
subalpine fir

Five Locations
Low - Wet
Mid - Wet
High - Wet
High - Dry
Low - Dry

−60

1890

−30

1920

0

1950

Year

30

1980

60

2010

P. silver fir
LPJ-GUESS: a process-based vegetation model

Cohorts of trees grow on patches of land (15m x 15m).
Each species is unique in its ability to establish, grow,
and die.
Continuous Climate (temperature, precipitation,
cloudiness) is the main driver.

http://www.nateko.lu.se/lpj-guess/education/html/guess.pdf
Process Parameters
title
nyear_spinup
vegmode
ifdailynpp
ifdailydecomp
ifbgestab
ifsme
ifstochmort
ifstochestab
estinterval
distinterval
iffire
ifdisturb
ifcalcsla
ifcdebt
npatch
patcharea
outputdirectory
ifsmoothgreffmort
ifdroughtlimitedestab
ifrainonwetdaysonly
ifspeciesspecificwateruptake

Shade Group Parameters (shadevintol, shadeintol, shadetol, shadevtol):
Title for run
Number of simulation years to spinup for
Vegetation mode ("INDIVIDUAL", "COHORT", "POPULATION")
Whether photosynthesis calculated daily (alt monthly)
Whether soil decomposition calculated daily (alt monthly)
Whether background establishment enabled (0,1)
Whether spatial mass effect enabled for establishment (0,1)
Whether mortality stochastic (0,1)
Whether establishment stochastic (0,1)
Interval for establishment of new cohorts (years)
Generic patch-destroying disturbance interval (years)
Whether fire enabled (0,1)
Whether generic patch-destroying disturbance enabled (0,1)
Whether SLA calculated from leaf longevity
Whether to allow C storage
Number of patches simulated
Patch area (m2)
Directory for the output files
Whether to vary mort_greff smoothly with growth efficiency (0,
Whether establishment drought limited (0,1)
Whether it rains on wet days only (1), or a little every day (0)
Whether or not there is species specific soil water uptake (0,1)

Global PFT Parameters:
lambda_max
emax
reprfrac
wscal_min

Non-water-stressed ratio of intercellular to ambient CO2 pp
Maximum evapotranspiration rate (mm/day)
Fraction of NPP allocated to reproduction
Water stress threshold for leaf abscission (raingreen PFTs)

Lifeform Parameters (Tree, Grass):
crownarea_max
ltor_max
k_allom2
k_allom3
k_rp
cton_leaf
cton_root
cton_sap
pathway
kest_repr
kest_bg
kest_pres
litterme
respcoeff
k_chilla
k_chillb
k_chillk

Maximum tree crown area (m2)
Non-water-stressed leaf:fine root mass ratio
Constant in allometry equations
Constant in allometry equations
Constant in allometry equations
Leaf C:N mass ratio
Fine root C:N mass ratio
Sapwood C:N mass ratio
Biochemical pathway ("C3" or "C4")
Constant in equation for tree estab rate
Constant in equation for tree estab rate
Constant in equation for tree estab rate
Litter moisture flammability threshold (fraction of AWC)
Respiration coefficient (0-1)
Constant in equation for budburst chilling time requirement
Coefficient in equation for budburst chilling time requirement
Exponent in equation for budburst chilling time requirement

turnover_sap
greff_min
est_max
alphar
parff_min

Sapwood turnover (fraction/year)
Threshold for growth suppression mortality (kgC/m2 leaf/yr)
Max sapling establishment rate (indiv/m2/year)
Shape parameter for recruitment-juv growth rate relationship
Min forest floor PAR for grass growth/tree estab (J/m2/day)

PFT (or species) specific parameters:
include
! Include PFT in analysis
phengdd5ramp
! GDD on 5 deg C base to attain full leaf cover
rootdist
! Fraction of roots in each soil layer (first value=upper layer)
turnover_leaf ! Leaf turnover (fraction/year)
wooddens
! Sapwood and heartwood density (kgC/m3)
k_allom1
! Constant in allometry equations
k_latosa
! Tree leaf to sapwood xs area ratio
sla
! Specific leaf area (m2/kgC)
fireresist
! Fire resistance (0-1)
tcmin_surv
! Min 20-year coldest month mean temp for survival (deg C)
tcmin_est
! Min 20-year coldest month mean temp for establishment (deg C)
tcmax_est
! Max 20-year coldest month mean temp for establishment (deg C)
twmin_est
! Min warmest month mean temp for establishment (deg C)
twminusc
! Stupid larch parameter
gdd5min_est
! Min GDD on 5 deg C base for establishment
longevity
! Expected longevity under lifetime non-stressed conditions (yr)
leaflong
! Leaf longevity (years)
drought_tolerance!Drought tolerance level (0 = very -> 1 = not at all) (unitless)

Output Files:
file_cmass
! !
file_anpp
! !
file_lai
! !
file_cflux
! !
file_dens
! !
file_cpool
! !
file_runoff
! !
file_firert
! !
file_mnpp
! !
file_mlai
! !
file_mgpp
! !
file_mra
! !
file_maet
! !
file_mpet
! !
file_mevap
! !
file_mrunoff !
!
file_mintercep ! !
file_mrh
! !
file_mnee
! !
file_mwcont_upper
file_mwcont_lower
file_speciesheights

C biomass output file
Annual NPP output file
LAI output file
C fluxes output file
Tree density output file
Soil C output file
Runoff output file
Fire retrun time output file
Monthly NPP output file
Monthly LAI output file
Monthly GPP-LeafResp output file
Monthly autotrophic respiration output file
Monthly AET output file
Monthly PET output file
Monthly Evap output file
Monthly runoff output file
Monthly intercep output file
Monthly heterotrphic respiration output
Monthly NEE output file
Monthly wcont_upper output file
Monthly wcont_lower output file
Mean species heights in 2000

Climate Group Parameters (Boreal, Temperate):
pstemp_min
pstemp_low
pstemp_high
pstemp_max

Approximate low temp limit for photosynthesis (deg C)
Approx lower range of temp optimum for photosynthesis (deg C)
Approx higher range of temp optimum for photosynthesis (deg C)
Maximum temperature limit for photosynthesis (deg C)

Leaf Group Parameters (Broadleaf, Needleleaf):
gmin
phenology
turnover_root
intc

Canopy conductance not assoc with photosynthesis (mm/s)
Phenology ("EVERGREEN", "SUMMERGREEN", "RAINGREEN" or "ANY")
Fine root turnover (fraction/year)
Interception coefficient

Parameters of LPJ-GUESS
(Smith et al. 2001)
Parameterizing LPJ-GUESS, 9 different tree species

Shade&Tolerance

Min&Annual&Post4Snow&
GDD

%&available&
soil4water

Maximum&
Age

Very&Intolerant

300

0.15

250

subalpine'fir

Very&Tolerant

300

0.5

250

mountain'hemlock

Very&Tolerant

300

0.68

400

Pacific'silver'fir

Very&Tolerant

800

0.7

400

Intolerant

800

0.2

750

western'hemlock

Very&Tolerant

1400

0.6

400

western'redcedar

Tolerant

1400

0.5

1000

Very&Intolerant

1400

0.5

80

Tolerant

1600

0.7

500

lodgepole'pine

Douglas8fir

red'alder
Sitka'spruce
14
12
10

JJA

8
6
4

SON

2

Temperature (C)

0

MAM

-2
-4
-6
-8

DJF

-10
-12
-14
-16

[decadal averages every 10 years]
-18
-20
2

Temperature (C)

0

Annual

-2
-4
-6

[-120.0, 46.389]

-8
-22

-20

-18

-16

-14

-12

-10

Age (-ka)

-8

-6

-4

-2

0

Figure by P.J. Bartlein
Results: present-day
Percent cover of tree species in
639 ecological plots in the
Olympic National Forest
(downloaded from Ecoshare.info)

East Side

West Side
1500

●

●

lodgepole pine

●

0

●

●

●

●

subalpine fir

●
●

●

●

●

mountain hemlock

●

elevation (meters)

Elevation (meters)

Simulated biomass of tree
species averaged over
the last 1000 years at each
site

●

●

●

●

Pacific silver fir

●
●

●

●

●

Douglas-fir

●
●

●
●

●

western hemlock

●
●

●
●

●

western redcedar

●
●

●
●

●

Sitka spruce

●
●

●

1500

●

●

red alder

●

0

●

●

0

25

50

75

0

25

50

% Cover in field plots (gray); Simulated biomass x 10 (black)

75
Percent of Total
< 0.5

0.5-2

2-10

10-50

Low Elevation, Wet Side

M = Model Results
D = Pollen Data
50-100

Low Elevation, Dry Side
M
D

grass

Grass

lodgepole pine

lodgepole pine

subalpine fir

subalpine fir

mountain hemlock

mountain hemlock

Pacific silver fir

Pacific silver fir

Douglas-fir

Douglas-fir

western hemlock

western hemlock

western redcedar

western redcedar

Sitka spruce

Sitka spruce

M
D

red alder
−15000

15,000

−10000

10,000

−5000

5000

0

0

red alder
−15000

15,000

Years before present

−10000

10,000

−5000

5000

0

0
Percent of Total
< 0.5

0.5-2

2-10

10-50

High Elevation, Wet Side

M = Model Results
D = Pollen Data
50-100

High Elevation, Dry Side
M
D

grass

grass

lodgepole pine

lodgepole pine

subalpine fir

subalpine fir

mountain hemlock

mountain hemlock

Pacific silver fir

Douglas-fir

Douglas-fir

western hemlock

western hemlock

western redcedar

M
D

Sitka spruce
−15000

15,000

−10000

10,000

−5000

5000

0

0

Sitka spruce
−15000

15,000

Years before present

−10000

10,000

−5000

5000

0

0
Mid Elevation, Wet Side
Late Glacial to late Holocene
transition is captured.....

M
D

grass
lodgepole pine
subalpine fir
mountain hemlock

But what about the abundant Douglas-fir and
alder pollen in the early Holocene

Pacific silver fir
Douglas-fir
western hemlock
western redcedar

M = Model Results
D = Pollen Data

Sitka spruce

M
D

red alder
−15000

−10000

15,000

10,000

−5000

0

5000

0

Years before present
At the Mid elevation, Wet Side site

Jan & July Temperature

Deg C

Jan & July Precipitation

22
20
18
16
14
12
10
8
6
4
2
0
−2
−4
−6
−8
−10
−15000

600

mm

400

200

0
−10000

−5000

Years before present

0

−15000

−10000

−5000

Years before present

0
Fraction
available
soil-water

Post-snow
GDD
1600

S. spruce

w. redcedar r. alder
1400 w. hemlock

S. spruce P. silver fir m. hemlock

0.7

w. hemlock
800

Doug-fir

P. silver fir

300 m. hemlock
l. pine

subalpine fir

subalpine fir

Simulated
Biomass
Kg C per
square
meter

−15000

−10000

w. redcedar

r. alder

0.5

Low - Wet
Mid - Wet

−20000

0.6

−5000

0

20

High - Wet

Doug-fir
−20000

0.2
0.15

l. pine
−15000

−10000

−5000

0

Fire Return
Time (years)

High - Dry
Low - Dry

15

900

10

600

5

300

0
−20000

−15000

20,000

15,000

−10000

10,000

−5000

5000

0

0

−20000

20,000

−15000

15,000

−10000

10,000

−5000

5000

0

0
The influence of snow

• The two climate variables that are controlling species composition
across the Peninsula are both influenced by snowpack

Rain-Snow Temp

More Snow

+2
+1
0
-1

Less Snow

-2
Fraction
available
soil-water

1600

Post-snow
GDD
1400

0.7
800

0.6
0.5

300
0.2
0.15
−20000

20,000

−15000

−10000

15,000

−5000

10,000

5000

0

−20000

−15000

−10000

−5000

0

0
Fire Return
Time (years)

Scenarios with varying Rain-Snow temp
threshold at the high elevation wet site
More Snow

900

+2
600

+1
0 °C

300

-1
Less Snow

-2
−20000

−15000

−10000

20,000

15,000

10,000

−5000

5000

0

0
Scenarios with varying Rain-Snow temp
threshold at the high elevation wet site

Thresholds crossed
0°C to 1°C
enough snow to favor mountain
hemlock (by limiting others)

10.0

7.5
variable

Simulated
Biomass

Pacific silver fir
subalpine fir
lodgepole pine
Douglas-fir
mountain hemlock
grass

5.0

Kg C per
square
meter

2.5

0.0
−2
Less Snow

−1
0
1
Rain-Snow Temp
°C

2
More Snow

1°C to 2°C
too much snow for consistent forest
cover, open canopy plants thrive (grass
and pine)
What did we learn?

•

Supported the theory that energy requirements control species composition on the west
side, and moisture controls composition on the east side

•

Quantified the climatic tolerance of seedlings in terms of one specific energy-related
climate variable (Post-snow GDD) and one moisture-related variable (fraction available
soil-water)

•

Provided the first species-level test against the paleoecological record of the influence
of these climate variables
Thank You!
References
Daly, C., M. Halbleib, J. I. Smith, W. P. Gibson, M. K. Doggett, G. H. Taylor, J. Curtis, and P. P. Pasteris. 2008. “Physiographically Sensitive Mapping of
Climatological Temperature and Precipitation Across the Conterminous United States.” International Journal of Climatology 28 (15) (December): 2031–
2064. doi:10.1002/joc.1688.
Gavin, D. G., L. B. Brubaker, and D. N. Greenwald. 2013. “Postglacial Climate and Fire- Mediated Vegetation Change on the Western Olympic Peninsula,
Washington.” Ecological Monographs (April 18). doi:10.1890/12-1742.1.
Henderson, J.A., R.D. Lesher, D.H. Peter, and C.D. Ringo. 2011. “A Landscape Model for Predicting Potential Natural Vegetation of the Olympic Peninsula
USA Using Boundary Equations and Newly Developed Environmental Variables”. USDA General Technical Report: PNW-GTR-941. USDA Forest Service.
Liu, Z., B. L. Otto-Bliesner, F. He, E. C. Brady, R. Tomas, P. U. Clark, A. E. Carlson, et al. 2009. “Transient Simulation of Last Deglaciation with a New
Mechanism for Bolling-Allerod Warming.” Science 325 (5938) (July 16): 310–314. doi:10.1126/science.1171041.
Smith, B., Prentice, I.C. & Sykes, M.T. 2001. Representation of vegetation dynamics in modelling of terrestrial ecosystems: comparing two contrasting
approaches within European climate space. Global Ecology and Biogeography 10: 621-637.

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Postglacial dynamics of Olympic Peninsula forests

  • 1. Postglacial dynamics of Olympic Peninsula forests: comparing simulations and observations Thanks to Dan Gavin Bart Bartlein Jed Kaplan A Master’s Thesis presented to the Department of Geography University of Oregon by David Fisher
  • 3. Research Questions • How does knowledge of the life history of tree species -- regeneration, growth, competition, mortality -- as well as the relationship between species and climate at important life stages, improve our explanation of the dynamic changes in forest composition since the Late Glacial period, 15,000 years ago?
  • 4. Outline of this talk • Paleoecology & Biogeography of Olympic Peninsula forests • Methods -- quantifying important bioclimatic variables and simulating paleoecological records with a process-based vegetation model • Results -- Simulations compared to present and paleo records • What did we learn?
  • 5. Paleoecological Records Cold, Pollen Percentage Diagram Yahoo Lake -Dry? Warm, Dry Cool, Wet Figure by D. G. Gavin (Gavin, D.G. 2013)
  • 6. Limiting factors for tree growth: On the west side: energy On the east side: moisture sretemoliK 08 04 02 0 ¯ 50km PRISM (Daly et al. 2008) 4 3 2 Potential Natural Forest Zones ¯ -4 0 -2 02 0 6 meters 04 PRISM (Daly et al. 2008) 2 deg C Mean Annual Precip sretemoliK 08 Minimum Jan Temp (Henderson et al. 2011) Alpine mountain hemlock subalpine fir Pacific silver fir Douglas-fir western hemlock Sitka spruce
  • 7. mountain hemlock Tsuga mertensiana western redcedar Thuja plicata subalpine fir Abies lasciocarpa western hemlock Tsuga heterophylla lodgepole pine Pinus contorta Sitka spruce Picea sitchensis Pacific silver fir Abies amabilis red alder Alnus rubra Douglas-fir Pseudotsuga menziesii grass
  • 8. Methods: Considering the seedling, how should we measure energy and moisture requirements?
  • 9. Methods: Considering the seedlings, how do we measure energy? Growing Degree Days Average Daily Temperature (Deg C) Post-Snow Growing Degree Days 15 2000 Size of Snowpack (Snow-Water equivalent) (mm) 10 1000 5 -5 J F M A M J J A S O N D 0
  • 10. Post-Snow GDD Measurements Local Measurements of Post-Snow GDD (two SNOTEL stations) Year 2006 2007 2008 2009 2010 2011 2012 SW 487 373 650 NE 649 432 493 670 473 384 563 SW = Buckinghorse NE = Waterhole http://www.wcc.nrcs.usda.gov/snotel/SNOTEL-brochure.pdf
  • 11. Post-Snow GDD Simulations annual PS-GDD required for each species to establish # Post-Snow GDDs 1600 1400 S. spruce w. redcedar w. hemlock r. alder 800 Doug-fir 300 m. hemlock l. pine subalpine fir Five Locations Low - Wet Mid - Wet High - Wet High - Dry Low - Dry −60 1890 −30 1920 0 1950 Year 30 1980 60 2010 P. silver fir
  • 12. LPJ-GUESS: a process-based vegetation model Cohorts of trees grow on patches of land (15m x 15m). Each species is unique in its ability to establish, grow, and die. Continuous Climate (temperature, precipitation, cloudiness) is the main driver. http://www.nateko.lu.se/lpj-guess/education/html/guess.pdf
  • 13. Process Parameters title nyear_spinup vegmode ifdailynpp ifdailydecomp ifbgestab ifsme ifstochmort ifstochestab estinterval distinterval iffire ifdisturb ifcalcsla ifcdebt npatch patcharea outputdirectory ifsmoothgreffmort ifdroughtlimitedestab ifrainonwetdaysonly ifspeciesspecificwateruptake Shade Group Parameters (shadevintol, shadeintol, shadetol, shadevtol): Title for run Number of simulation years to spinup for Vegetation mode ("INDIVIDUAL", "COHORT", "POPULATION") Whether photosynthesis calculated daily (alt monthly) Whether soil decomposition calculated daily (alt monthly) Whether background establishment enabled (0,1) Whether spatial mass effect enabled for establishment (0,1) Whether mortality stochastic (0,1) Whether establishment stochastic (0,1) Interval for establishment of new cohorts (years) Generic patch-destroying disturbance interval (years) Whether fire enabled (0,1) Whether generic patch-destroying disturbance enabled (0,1) Whether SLA calculated from leaf longevity Whether to allow C storage Number of patches simulated Patch area (m2) Directory for the output files Whether to vary mort_greff smoothly with growth efficiency (0, Whether establishment drought limited (0,1) Whether it rains on wet days only (1), or a little every day (0) Whether or not there is species specific soil water uptake (0,1) Global PFT Parameters: lambda_max emax reprfrac wscal_min Non-water-stressed ratio of intercellular to ambient CO2 pp Maximum evapotranspiration rate (mm/day) Fraction of NPP allocated to reproduction Water stress threshold for leaf abscission (raingreen PFTs) Lifeform Parameters (Tree, Grass): crownarea_max ltor_max k_allom2 k_allom3 k_rp cton_leaf cton_root cton_sap pathway kest_repr kest_bg kest_pres litterme respcoeff k_chilla k_chillb k_chillk Maximum tree crown area (m2) Non-water-stressed leaf:fine root mass ratio Constant in allometry equations Constant in allometry equations Constant in allometry equations Leaf C:N mass ratio Fine root C:N mass ratio Sapwood C:N mass ratio Biochemical pathway ("C3" or "C4") Constant in equation for tree estab rate Constant in equation for tree estab rate Constant in equation for tree estab rate Litter moisture flammability threshold (fraction of AWC) Respiration coefficient (0-1) Constant in equation for budburst chilling time requirement Coefficient in equation for budburst chilling time requirement Exponent in equation for budburst chilling time requirement turnover_sap greff_min est_max alphar parff_min Sapwood turnover (fraction/year) Threshold for growth suppression mortality (kgC/m2 leaf/yr) Max sapling establishment rate (indiv/m2/year) Shape parameter for recruitment-juv growth rate relationship Min forest floor PAR for grass growth/tree estab (J/m2/day) PFT (or species) specific parameters: include ! Include PFT in analysis phengdd5ramp ! GDD on 5 deg C base to attain full leaf cover rootdist ! Fraction of roots in each soil layer (first value=upper layer) turnover_leaf ! Leaf turnover (fraction/year) wooddens ! Sapwood and heartwood density (kgC/m3) k_allom1 ! Constant in allometry equations k_latosa ! Tree leaf to sapwood xs area ratio sla ! Specific leaf area (m2/kgC) fireresist ! Fire resistance (0-1) tcmin_surv ! Min 20-year coldest month mean temp for survival (deg C) tcmin_est ! Min 20-year coldest month mean temp for establishment (deg C) tcmax_est ! Max 20-year coldest month mean temp for establishment (deg C) twmin_est ! Min warmest month mean temp for establishment (deg C) twminusc ! Stupid larch parameter gdd5min_est ! Min GDD on 5 deg C base for establishment longevity ! Expected longevity under lifetime non-stressed conditions (yr) leaflong ! Leaf longevity (years) drought_tolerance!Drought tolerance level (0 = very -> 1 = not at all) (unitless) Output Files: file_cmass ! ! file_anpp ! ! file_lai ! ! file_cflux ! ! file_dens ! ! file_cpool ! ! file_runoff ! ! file_firert ! ! file_mnpp ! ! file_mlai ! ! file_mgpp ! ! file_mra ! ! file_maet ! ! file_mpet ! ! file_mevap ! ! file_mrunoff ! ! file_mintercep ! ! file_mrh ! ! file_mnee ! ! file_mwcont_upper file_mwcont_lower file_speciesheights C biomass output file Annual NPP output file LAI output file C fluxes output file Tree density output file Soil C output file Runoff output file Fire retrun time output file Monthly NPP output file Monthly LAI output file Monthly GPP-LeafResp output file Monthly autotrophic respiration output file Monthly AET output file Monthly PET output file Monthly Evap output file Monthly runoff output file Monthly intercep output file Monthly heterotrphic respiration output Monthly NEE output file Monthly wcont_upper output file Monthly wcont_lower output file Mean species heights in 2000 Climate Group Parameters (Boreal, Temperate): pstemp_min pstemp_low pstemp_high pstemp_max Approximate low temp limit for photosynthesis (deg C) Approx lower range of temp optimum for photosynthesis (deg C) Approx higher range of temp optimum for photosynthesis (deg C) Maximum temperature limit for photosynthesis (deg C) Leaf Group Parameters (Broadleaf, Needleleaf): gmin phenology turnover_root intc Canopy conductance not assoc with photosynthesis (mm/s) Phenology ("EVERGREEN", "SUMMERGREEN", "RAINGREEN" or "ANY") Fine root turnover (fraction/year) Interception coefficient Parameters of LPJ-GUESS (Smith et al. 2001)
  • 14. Parameterizing LPJ-GUESS, 9 different tree species Shade&Tolerance Min&Annual&Post4Snow& GDD %&available& soil4water Maximum& Age Very&Intolerant 300 0.15 250 subalpine'fir Very&Tolerant 300 0.5 250 mountain'hemlock Very&Tolerant 300 0.68 400 Pacific'silver'fir Very&Tolerant 800 0.7 400 Intolerant 800 0.2 750 western'hemlock Very&Tolerant 1400 0.6 400 western'redcedar Tolerant 1400 0.5 1000 Very&Intolerant 1400 0.5 80 Tolerant 1600 0.7 500 lodgepole'pine Douglas8fir red'alder Sitka'spruce
  • 15. 14 12 10 JJA 8 6 4 SON 2 Temperature (C) 0 MAM -2 -4 -6 -8 DJF -10 -12 -14 -16 [decadal averages every 10 years] -18 -20 2 Temperature (C) 0 Annual -2 -4 -6 [-120.0, 46.389] -8 -22 -20 -18 -16 -14 -12 -10 Age (-ka) -8 -6 -4 -2 0 Figure by P.J. Bartlein
  • 16. Results: present-day Percent cover of tree species in 639 ecological plots in the Olympic National Forest (downloaded from Ecoshare.info) East Side West Side 1500 ● ● lodgepole pine ● 0 ● ● ● ● subalpine fir ● ● ● ● ● mountain hemlock ● elevation (meters) Elevation (meters) Simulated biomass of tree species averaged over the last 1000 years at each site ● ● ● ● Pacific silver fir ● ● ● ● ● Douglas-fir ● ● ● ● ● western hemlock ● ● ● ● ● western redcedar ● ● ● ● ● Sitka spruce ● ● ● 1500 ● ● red alder ● 0 ● ● 0 25 50 75 0 25 50 % Cover in field plots (gray); Simulated biomass x 10 (black) 75
  • 17. Percent of Total < 0.5 0.5-2 2-10 10-50 Low Elevation, Wet Side M = Model Results D = Pollen Data 50-100 Low Elevation, Dry Side M D grass Grass lodgepole pine lodgepole pine subalpine fir subalpine fir mountain hemlock mountain hemlock Pacific silver fir Pacific silver fir Douglas-fir Douglas-fir western hemlock western hemlock western redcedar western redcedar Sitka spruce Sitka spruce M D red alder −15000 15,000 −10000 10,000 −5000 5000 0 0 red alder −15000 15,000 Years before present −10000 10,000 −5000 5000 0 0
  • 18. Percent of Total < 0.5 0.5-2 2-10 10-50 High Elevation, Wet Side M = Model Results D = Pollen Data 50-100 High Elevation, Dry Side M D grass grass lodgepole pine lodgepole pine subalpine fir subalpine fir mountain hemlock mountain hemlock Pacific silver fir Douglas-fir Douglas-fir western hemlock western hemlock western redcedar M D Sitka spruce −15000 15,000 −10000 10,000 −5000 5000 0 0 Sitka spruce −15000 15,000 Years before present −10000 10,000 −5000 5000 0 0
  • 19. Mid Elevation, Wet Side Late Glacial to late Holocene transition is captured..... M D grass lodgepole pine subalpine fir mountain hemlock But what about the abundant Douglas-fir and alder pollen in the early Holocene Pacific silver fir Douglas-fir western hemlock western redcedar M = Model Results D = Pollen Data Sitka spruce M D red alder −15000 −10000 15,000 10,000 −5000 0 5000 0 Years before present
  • 20. At the Mid elevation, Wet Side site Jan & July Temperature Deg C Jan & July Precipitation 22 20 18 16 14 12 10 8 6 4 2 0 −2 −4 −6 −8 −10 −15000 600 mm 400 200 0 −10000 −5000 Years before present 0 −15000 −10000 −5000 Years before present 0
  • 21. Fraction available soil-water Post-snow GDD 1600 S. spruce w. redcedar r. alder 1400 w. hemlock S. spruce P. silver fir m. hemlock 0.7 w. hemlock 800 Doug-fir P. silver fir 300 m. hemlock l. pine subalpine fir subalpine fir Simulated Biomass Kg C per square meter −15000 −10000 w. redcedar r. alder 0.5 Low - Wet Mid - Wet −20000 0.6 −5000 0 20 High - Wet Doug-fir −20000 0.2 0.15 l. pine −15000 −10000 −5000 0 Fire Return Time (years) High - Dry Low - Dry 15 900 10 600 5 300 0 −20000 −15000 20,000 15,000 −10000 10,000 −5000 5000 0 0 −20000 20,000 −15000 15,000 −10000 10,000 −5000 5000 0 0
  • 22. The influence of snow • The two climate variables that are controlling species composition across the Peninsula are both influenced by snowpack Rain-Snow Temp More Snow +2 +1 0 -1 Less Snow -2
  • 23. Fraction available soil-water 1600 Post-snow GDD 1400 0.7 800 0.6 0.5 300 0.2 0.15 −20000 20,000 −15000 −10000 15,000 −5000 10,000 5000 0 −20000 −15000 −10000 −5000 0 0 Fire Return Time (years) Scenarios with varying Rain-Snow temp threshold at the high elevation wet site More Snow 900 +2 600 +1 0 °C 300 -1 Less Snow -2 −20000 −15000 −10000 20,000 15,000 10,000 −5000 5000 0 0
  • 24. Scenarios with varying Rain-Snow temp threshold at the high elevation wet site Thresholds crossed 0°C to 1°C enough snow to favor mountain hemlock (by limiting others) 10.0 7.5 variable Simulated Biomass Pacific silver fir subalpine fir lodgepole pine Douglas-fir mountain hemlock grass 5.0 Kg C per square meter 2.5 0.0 −2 Less Snow −1 0 1 Rain-Snow Temp °C 2 More Snow 1°C to 2°C too much snow for consistent forest cover, open canopy plants thrive (grass and pine)
  • 25. What did we learn? • Supported the theory that energy requirements control species composition on the west side, and moisture controls composition on the east side • Quantified the climatic tolerance of seedlings in terms of one specific energy-related climate variable (Post-snow GDD) and one moisture-related variable (fraction available soil-water) • Provided the first species-level test against the paleoecological record of the influence of these climate variables
  • 26. Thank You! References Daly, C., M. Halbleib, J. I. Smith, W. P. Gibson, M. K. Doggett, G. H. Taylor, J. Curtis, and P. P. Pasteris. 2008. “Physiographically Sensitive Mapping of Climatological Temperature and Precipitation Across the Conterminous United States.” International Journal of Climatology 28 (15) (December): 2031– 2064. doi:10.1002/joc.1688. Gavin, D. G., L. B. Brubaker, and D. N. Greenwald. 2013. “Postglacial Climate and Fire- Mediated Vegetation Change on the Western Olympic Peninsula, Washington.” Ecological Monographs (April 18). doi:10.1890/12-1742.1. Henderson, J.A., R.D. Lesher, D.H. Peter, and C.D. Ringo. 2011. “A Landscape Model for Predicting Potential Natural Vegetation of the Olympic Peninsula USA Using Boundary Equations and Newly Developed Environmental Variables”. USDA General Technical Report: PNW-GTR-941. USDA Forest Service. Liu, Z., B. L. Otto-Bliesner, F. He, E. C. Brady, R. Tomas, P. U. Clark, A. E. Carlson, et al. 2009. “Transient Simulation of Last Deglaciation with a New Mechanism for Bolling-Allerod Warming.” Science 325 (5938) (July 16): 310–314. doi:10.1126/science.1171041. Smith, B., Prentice, I.C. & Sykes, M.T. 2001. Representation of vegetation dynamics in modelling of terrestrial ecosystems: comparing two contrasting approaches within European climate space. Global Ecology and Biogeography 10: 621-637.