Physioecological mechanism of the alpine treeline dynamics under global climate change
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Physioecological mechanism Physioecological mechanism of the alpine treeline dynamics of the alpine treeline dynamics
under global climate changeunder global climate change
Swiss Federal Research Institute WSL
Dr. Mai-He Li
E-mail: [email protected]
The 2nd Third Pole Environment WorkshopKathmandu, Oct. 26 – 28, 2010
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1. Definition of the alpine treeline
2. Driving forces of treeline formation
3. Upword shifts of the alpine treelines, corresponding to global warming, in the Himalayas and worldwide
4. Physioecological machanisms of the alpine treeline shifts
5. Conclusions
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Treelines across the globe ---a virtual site visitation
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Davos, Swiss Alps The Himalayas
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Ch Koerner, J Paulsen 2004
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Miehe et al. 2007. Mountain Res. & Develop. 27, 169-173
Kyi Chu, North of Lhasa29°42‘ N, 96°45 ‘ E
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Holtmeier & Broll, 2009. Polarforschung 79, 139-153
Factors affecting trees at the alpine treelines
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The alpine treeline position is very closely correlated to the 10°C
isotherm for the warmest month
The temperature at the alpine treelines varied from 6 to 13°C (±500 m in treeline elevation)
Koeppen 1923; Aulitzky 1961Wu 1983; Oshawa 1990
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n=2n=211 1212
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Sea
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l mea
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°C)
Sea
sona
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mpe
ratu
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T (
°C)
7070 6060 5050 4040 3030 2020 1010 00 1010 2020 3030 4040Latitude (°Latitude (°CC))
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100100
200200
300300
Gro
win
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riod
G (
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6.7 ± 0.8 °C 6.7 ± 0.8 °C
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Ch Körner, J Paulsen (2004) J Biogeogr 31:713-732Ch Körner, J Paulsen (2004) J Biogeogr 31:713-732
A global mean of 6.7 °C soil temperature (-10 cm depth) for growing season at treeline
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Source: Massachusetts Ave, Cambridge MA
Predicting changes in temperature
It is predicted that the temperatures in the Indian sub-continent will rise between 3.5 and 5.5°C by 2010, and on the Tibetan Plateau by 2.5°Cby 2050, and 5°C by 2010
Kumar et al. 2006
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Location Authors, year Species Upword shift (m)
Period (year)
Rate (m/yr)
South Island, New Zealand
Wardle & Coleman, 1992
Nothofagus menziesii, N. solandri
7-9 1930-90 0.12-0.15
Italian Alps Leonelli et al. 2010 115 1901-2000 1.15
Glacier N.P. Montana
Bekker, 2005 Picea engelmannii, Pinus contorta, Abies lasciocarpa
7-16 1800-1980 0.28-0.62
Sunwapta Pass, Alberta
Luckman & Kavanagh, 2000
P. engelmannii, A. lasciocarpa
145 1700-1994 0.5
Uinta Mts. UT Munroe, 2003 P. engelmannii, A. lasciocarpa
61-183 1870-2001 0.5-1.4
SW Yukon Danby & Hik, 2007 Picea glauca 65-85 1920-2005 0.8-1.0
Scands Mts, Sweden
Kullman, 2001 Betula pubescens, Pinus sylvestris, Picea abies
100-165 1915-2000 1.2-1.9
Kootenay NP, B.C.
Roush, 2009 Larix lyallii, P. engelmannii, A. lasciocarpa
149 1909-1976 2.2-5.7
W. Himalayas of India
Dubey et al. 2003 14-19 Over 10 yrs
1.4-1-9
Baima Snow Mt, Yunnan
Baker & Moseley, 2007
67 Since 1923
Nanda Devi in C. Himalaya
Panigrahy et al., 2010
300 Since 1960
Himalayas, Nepal
Vijayaprakash & Ansari, 2009
Abies spectabilis N-aspect South-asp.
1.7 (N-s)2.3 (S-s)
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Species No. Elevation (1970) Elevation (1992) Elevational shift (m)
Vaccinium uliginosum 11 2216 2216 0
Vaccinium vitis-idaea 30 1898 1885 -13
Vaccinium myrtillus 31 1972 2002 30
Rhododendron ferrugineum 30 2098 2098 0
Pinus cembra 31 2064 2005 -59
Hieracium pilosella 13 1851 1930 79
Scleropodium purum 26 1425 1630 205
Dupouey et al. 1997
Each species is likely to respond to climate change in its own way:
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Treeline formationTreeline formation
Global driversGlobal drivers'general principle''general principle'
Regional driversRegional drivers'modulation''modulation'
General physioecologicalexplanation
Local environmentalexplanations
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(1)(1) The The stressstress hypothesis hypothesis
(2) The (2) The maturationmaturation time hypothesis time hypothesis
(3)(3) The The disturbancedisturbance hypothesis hypothesis
(4)(4) The The reproduction/germinationreproduction/germination
hypothesishypothesis
Environmental explanation of treelineEnvironmental explanation of treeline
Körner Ch (1998) Oecologia 115:445Li MH, Krauchi N (2005) J.S.For.Tech.26, 36-42Li MH et al. 2008. Tree Physiology 28, 1287-1296 Li MH et al. 2008. Plant, Cell & Environment 31, 1377-1387
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(5)(5)The The growthgrowth limitation hypothesis limitation hypothesis
(6)(6)The The carboncarbon balance hypothesis balance hypothesis
Biological explanation of treelineBiological explanation of treeline
Körner Ch (1998) Oecologia 115:445Li MH, Krauchi N (2005) J.S.For.Tech.26, 36-42Li MH et al. 2008. Tree Physiology 28, 1287-1296 Li MH et al. 2008. Plant, Cell & Environment 31, 1377-1387
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1. The stress hypothesis
• Repeated damage
by freezing, frost desiccation or phototoxic effects after frost impair
tree growth
Tranquillini W.1979. Physiological ecology of the alpine timerlineKörner Ch. 1998. Oecologia 115, 445-459Li MH, Kräuchi N. 2005. J.S.For.Tech. 26, 36-42
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Maturation of leaves,shoots, fruits, and buds
e.g. seed maturation of Pinus sylvestris needs at least 600 – 890 GDD (growing degree-days >5°C)
Odum 1979
2.2. The The maturationmaturation time hypothesis time hypothesis
Short gro
wing seaso
n
Tranquillini W.1979. Physiological ecology of the alpine timerlineKörner Ch. 1998. Oecologia 115, 445-459Li MH, Kräuchi N. 2005. J.S.For.Tech. 26, 36-42
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Mechanic damage by wind, ice blasting, snow break and avalanches, fire……
Animal disturbances such as insectFungal pathogensMan-made impacts such as logging, grazing etc.
3.3. The The disturbancedisturbance
hypothesishypothesis
Tranquillini W.1979. Physiological ecology of the alpine timerlineKörner Ch. 1998. Oecologia 115, 445-459Li MH, Kräuchi N. 2005. J.S.For.Tech. 26, 36-42
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Pollination, pollen tube growth, seed development, seed dispersal, germination and seedling establishment
4.4. The The reproductionreproduction hypothesis hypothesis
Tranquillini W.1979. Physiological ecology of the alpine timerlineKörner Ch. 1998. Oecologia 115, 445-459Li MH, Kräuchi N. 2005. J.S.For.Tech. 26, 36-42
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5. Growth limitation5. Growth limitation
PhotosynthesisPhotosynthesis
GrowthGrowth
““Sink“ drivenSink“ driven
““Demand“ drivenDemand“ driven
PhotosynthesisPhotosynthesis
GrowthGrowth
““Source“ drivenSource“ driven
““Supply“ drivenSupply“ driven
6. Carbon limitation 6. Carbon limitation
Körner Ch (1998) Oecologia 115:445Li MH et al. 2008. Tree Physiology 28, 1287-1296 Li MH et al. 2008. Plant, Cell & Environment 31, 1377-1387
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Source limitation hypothesis:Tree growth is considered source limited when carbon assimilation through photo-synthesis is insufficient to meet growth requirements.
Sink limitation hypothesis:
Trees are considered carbon sink limited when there is an abundant supply of the resources necessary to support growth, but growth itself is directly limited by environ-mental conditions.
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Mito
tic t
ime
(h)
Mito
tic t
ime
(h)
Temperature (°C)Temperature (°C)
00
1010
2020
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Cel
l dou
blin
g tim
e (h
)C
ell d
oubl
ing
time
(h)
00
100100
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Mitotic timeMitotic timeCell doubling timeCell doubling time
Ch Körner (2003) Ch Körner (2003) Alpine Plant LifeAlpine Plant Life. Springer, Berlin. Springer, Berlin
5050
100100
Ne
t-ph
oto
syn
thes
is (
%)
Ne
t-ph
oto
syn
thes
is (
%)
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Late seasonLate season
End of winterEnd of winter
Mid seasonMid season
Non
-str
uctu
ral c
arbo
hydr
ates
+ li
pids
N
on-s
truc
tura
l car
bohy
drat
es +
lipi
ds
in s
tem
sap
woo
d (%
d.m
.)in
ste
m s
apw
ood
(% d
.m.)
AltitudeAltitude
00
22
44
00
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00
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NSCNSC
LipidsLipids****
****
******
HighHighLowLowHighHighLowLowHighHighLowLow
Pinus hartwegiiPinus hartwegiiMexico (19° N)Mexico (19° N)Pinus cembraPinus cembraAlps (46° N)Alps (46° N) Pinus sylvestrisPinus sylvestrisSweden (68° N)Sweden (68° N)
G Hoch & C Körner (2003) Oecologia 135:10-21
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00
1010
2020
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4040
5050
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1010
2020
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NS
C c
once
ntr
atio
ns (
mg
cm
NS
C c
once
ntr
atio
ns (
mg
cm
33 ))
43604360 45504550 4810481043604360 45504550 48104810
Elevation (m a.s.l.)Elevation (m a.s.l.)
BranchwoodBranchwood StemwoodStemwood
00
4040
8080
120120
160160
200200
G Hoch & Ch Körner (2005) Funct Ecol 19, 941-951
43604360 45504550 4810481043604360 45504550 4810481043604360 45504550 4810481043604360 45504550 48104810
LeavesLeaves
abab bb
aa
aaaa
a a aaaa aa
Polylepis tarapacana, Polylepis tarapacana, Volcano Sajama, BoliviaVolcano Sajama, Bolivia
00
1010
2020
3030
4040
5050
00
1010
2020
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4040
5050
Sug
ars
Sug
ars
Sta
rch
Sta
rch
NSC=Non-structural carbohydrates = solube sugars + starch
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Gas exchange with altitude
Cabrera HM et al. 1998, Oecologia 114, 145-152
Acaena cylindrostachya
Senecio formosus
Maximum CO2 assimilation rates4200 m: 3.9 µ mol/m2 s3550 m: 5.2 µ mol/m2 s2900 m: 9.0 µ mol/m2 s
4200 m: 3.6 µ mol/m2 s3550 m: 5.8 µ mol/m2 s2900 m: 7.5 µ mol/m2 s
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0
5
10
15
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25
30
3800
m
3400
m
3800
m
3400
m
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m
3400
m
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m
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m
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m
April July April July April July April July April July
Elevations of trees and sampling time
Sta
rch
/ S
ug
ars
/ NS
C c
on
cen
trat
ion
(%
d.m
.) Sugars
Starch
1-yr-old needles
Stem wood
Fine roots
a b
*
a a a a a ab b b b bb
AB
* ***
*
*
2-yr-oldneedles
3-yr-oldneedles
Li MH et al. 2008. Tree Physiology 28, 1287-1296 Li MH et al. 2008. Plant, Cell & Environment 31, 1377-1387
Picea balfouriana var. hirtella
Carbon shortage? - Yes!
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Soluble sugars Starch NSC
April July April July April July
Three treeline cases combined (Lower E trees = lower elevation trees)
Treeline trees 11.09 10.51 3.61 4.19 14.70 14.70
Lower E trees 10.66 10.37 4.77 4.55 15.43 14.91
F1,89 3.53 0.24 31.00 1.35 6.80 0.31
p 0.07 0.63 <0.001 0.25 0.011 0.58
An overall trend in NSC – 3 treelines data pooledAn overall trend in NSC – 3 treelines data pooled
A winter C-shortageA winter C-shortage
Li MH et al. 2008. Tree Physiology 28, 1287-1296 Li MH et al. 2008. Plant, Cell & Environment 31, 1377-1387
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A winter carbon shortage?orAn effect of phenological phase-shift?
Hoch G. 2003PhD thesisUni. Basel
??????
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Camarero & Gutierrez (2002) Plant ecology 162: 247
Krummholz
SeedlingsDead individuals
Tree species line
Timberline
Adults
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General conclusionGeneral conclusion
• The treeline trees may suffer from a The treeline trees may suffer from a winter winter carbon shortagecarbon shortage leading to treeline formation leading to treeline formation
• Global warming leads to Global warming leads to increase in treeline increase in treeline elevationelevation to keep pace with climate change to keep pace with climate change
• PPlants may also respond to climate change in lants may also respond to climate change in Himalayas viaHimalayas via
• Re-adaptationRe-adaptation• InvasionsInvasions• ExtinctionExtinction
Li MH et al. 2006. J Integrat Plant Biology 48, 255 - 259Li MH et al. 2008. Tree Physiology 28, 1287-1296 Li MH et al. 2008. Plant, Cell & Environment 31, 1377-1387
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30
Modeled Climate-Induced Glacier and Vegetation Change in Glacier National Park, 1850-2100
http://www.nrmsc.usgs.gov/images/glacier_animation_slow.gif
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Cavieres et al. 2000. Acta Oecologia 21, 203-211
Podocarpus oleifoliusPodocarpus oleifolius
2550 m a.s.l.2550 m a.s.l.
3200 m a.s.l.3200 m a.s.l.2400 m a.s.l.2400 m a.s.l.
3200 m a.s.l.3200 m a.s.l.
Espeletia neriifoliaEspeletia neriifolia
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Richardson AD. 2004Plant & Soil 260, 291-299
P<0.05P<0.05P<0.05P<0.05
P<0.05P<0.05 P<0.05P<0.05
Abies balsamea Picea rubens
Shade needlesShade needles
Sun needlesSun needles