ForestEcology and Managemem259 (2010) 660-684

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ForestEcology and Management
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journal homepage: www.elsevier.comllocate/foreco
Aglobal overviewofdroughtand heat-induced tree mortality reveals
emergingclimate change risks for forests
Craig D. Allena.*, Alison K. Macaladyb, Haroun Chenchounic, Dominique Bacheletd, Nate McDowelle,
Michel Vennetierr, Thomas Kitzbergerg, Andreas Rigling1\ David D. Breshearsi, E.H. (Ted) Hoggj,
Patrick Gonzalez1<, Rod Fenshamt, Zhen Zhangm, Jorge Castro11, Natalia Demidova0,
Jong-Hwan Limp. GillianAllardq, Steven W. Runningr. Akkin Semerci5, Neil Cobbt
U,S Geological Survey, Fort Col/ins SCience Cepner,Jemez MounlOlns Field 510 non, Los Alamos, NM 87544, USA
bSchool of Geography ond Development and Laboratory of Tree-Rmg Reseorch, University of Arizona, Tucson, AZ 85721, USA
'Deporrmentof Biology, University of 80rno, 05000 Borna, Algerio
d Deparrmenr of BiolOgical and ecological engineering. OregonState UniverSIl)', CorvoWis, OR 97330, USA
Eareh ond Environmental SCiences, MS J495, Los Alamos Notiollol Lobomrory. Los Alamos, NM 87544,USA
rCEMAGREF, ECCOREV FR 3098, Aix-MarseilJe University. Aix-ea-Pravence, France
gLobaratorioEcotono, 1N/8IOMA-CONICET and Univ. Naaanal del Comalwe, Quinrra/ 1250. 8400 Boriloche, Argennna
h Swiss Federa/lnstitute for Forest, Snow and Landscape Research VV'iL. larellentr. //r, CH-890J Birmensdorf. Switzerland
'School of Natural Resources and rhe Environment, and Deporrment of Ecolagy and Eva/urianary Bia/agy, Universil)' af Arizona, Tucson, Al 85721. USA
J Northel7l Farestry Centre. Canadian Farest Service, 5320-/22 Street. Edmonton, A/berta T6H 355, Conoda
kCenter for Forestry, Univenity of California, Berkeley, CA 94720, USA
IQueellSland Herbarium, Envlronmenral Protection Agency, MI Caot-tha Raad, Taowallg. Queenslond 4066, AustralJO
on Research Illstitute af Fores[ Ecolagy, Ellvironment alld Pra/eetian. Chinese Academy of Forestry.
Key Labaratory of Faresr ProtectIOn af SlOte Faresrry Adminlstraliall, Berjmg 100091, China
nGrupo de Ecolagra Terresrre, Departo menro de Eealogio, Universidad de Granada, Granada E-1807f, Spa in
NortheTil Research Insnrute ofFaresny, NikilOV 5r., 13, Arkhangelsk 1G3062, Russion Federanon
PDivision of Fore.<t ecology. Departmenr Of ForesrConservation. Karea Forest Research Inmwre #57, Hoegl-rG, Dongdaemun-gu, Seoul 130-712, RepUblic Of Korea
q Forestry Deparrmellr. Food alld Agriculrure Organizanon(FAG), Viole delle Tenne di Caraeal/a. 00100 Rame, Uaiy
''''umenco) Terradynamlcs Silnu!onon Group, UniverSity of Montana, Mi"ouJo, MT 59812, USA
'Ceatral Anoralia Forestry Research }nsnlu[e, P.K 24, 06501 /JahceJiev!er-Ankara, Turkey
'Depamnenr Of BiolOgical Sciences and Men-iam Powell Cenler for Ellvlronlllenro! Research. "'orrhernArIZona Universll)', Flagstaff, AZ 860lJ. USA
ARTICLE INFO ABSTRACT
Article hisrory:
Ree", ved 4March 2009
Received in revised rorm 13 Augusr 2009
Accepted 1September2009
KeywordS"
Climate (hange
Droughr eflecr<
porest dIe-orr
Forest mortality
Global pattems
Tree mortality
Greenhousegas emissions have significantly altered global climate.and will continue to do so in the
future. Increasesin thefrequency, duranon,and/orseverityofdroughtand heatstressassociatedwith
climate change could rundamentally alter the composition, structure, and biogeography offorests in
many regions. Of particularconcern are potential increases in tree mortalityassociated with c1imate-
induced physiological stress and interactions with otherclimate-mediated processes such as insect
ourbreaksand wildfire. Despiterhis risk,eXisring projectionsofcceemortalityarebasedon Inodels that
lackfunctionally realisticmorralitymechanisms,and therehas been no attempttotrackobservationsor
cllmare-driven tree mortality globally. Here we present the first global assessment or recent tree
mortality attributed to droughtand heatstress. Alrhough episodic mortality occurs in the absence of
climate change. studies compiled here suggest that at least some ofthe world's forested ecosystems
alreadymayberesponding to climatechangeand raise concern thatforests may become increasingly
vulnerableto higher background tree mortality rates and die-off in response to future warmingand
drought, even in environments thatare not normallyconsidered water-limited.This further suggests
risks to ecosystemservices, includingthe lossofsequesteredforest carbonandassociated atmospheric
feedbacks, Ourreviewalso identifies key information gaps and scientific uncertainties that currently
hinderourability to predicttreemortalityin responsetoclimatechangeandemphasizestheneed fora
globally coordinated observarion system. Overall, our review reveals rhe potential for amplified tree
mortalitydue to droughtand heatin forests worldwide.
Published by Elsevier B.V.
Correspolldl1\g auchor. Tel:+1 505672 3861x541: r"x: +1 5056729607,
Emall address: eraig...allen@u5gs.gov (CD.Allen).
0378-1127/$ - see front mateer, Published by Elsevier B.V.
doi:10.1016/J.foreco.2009.09.001
661 CD. Allen et aL/Fore;t Ecology ond Management 259 (2010) 660-684
1. Introduction
Forested ecosystems are being rapi dly and directly transformed
by the land uses of our expanding human populations and
economies. Currently less evident are the impacts of ongoing
climate change on the world's forests. Increasing emissions of
greenhouse gases are now widely acknowledged by the scientific
community as a major cause of recent increases in global mean
temperature (about 0.5 cC since 1970) and changes in the world's
hydrological cycle (lPCC, 2007a), ind uding a widening of the
Earth's tropical belt (Seidel et aI., 2008; Lu et a!., 2009). Even under
conservative scenarios, future climate changes are likely to include
fnrther increases in mean temperature (about 2-4"C globally)
with significant drying in some regions (Chrisrensen et al.. 2007:
Seageret aI., 2007), as well as increases in frequency and severity of
extreme drollgllts, hot extremes, and heat waves (IPCC, 2007a:
Sterl et aI., 2008).
Understand1l1g and predicting the consequences of these
climatic cllanges on ecosystems is emerging as one of the grand
challenges for global change scientists, and forecasting the impacts
on forests is of particular importance (Boisvenue and Running,
2006: Bonan, 2008). Forests, here hroadly defined to include
woodlands and savannas, cover 30% of the world's land surface
(FAD, 2006). Around the globe societies rely on forests for essential
services such as timber and watershed protection. and less tangible
but equally important recreational, aesthetic, and spiritual
benefits. The effects of climate change on forests include both
positive (e.g. increases in forest vigor and growth from CO
2
fertilization. increased water use efficiency. and longer growing
seasons) and negative effects (e.g. reduced growth and increases in
stress and mortality due to the combined impacts of climate
change and climate-driven changes in the dynamics of forest
insects and pathogens) (Ayres and Lombardero, 2000: Bachelet
et aI., 2003: Lucht ec aI., 2006; Scholze et aI., 2006: Lloyd and Bunn,
2007). Furthermore, forests are subject to many other human
influences such as increased ground-level ozone and deposition
(Fowler et a!., 1999; Karnosl{j' et aI., 2005; Ollinger ec aI., 2008).
Considerable uncertainty remains in modeling how these and
other relevant processes will affect the risk of future tree die-off
events, referred to hereafter as 'forest mortality', under a changing
climate (LoehJe and LeBlanc, 1996; Hanson and Weltzin, 2000:
Bugmann et aI., 2001). Alchough a range of responses can and
should be expected, recent cases of increased tree mortality and
die-offs triggered by drought and/or high temperatures raise the
pOSSibility that amplified forest mortality may already be occurring
in some locations in response to global climate change. Examples of
recent die-offs are particularly well documented for southern parts
ofEurope (Peiiuelas et al., 2001; Breda et aI., 2006; Bigleretal., 2006)
and for temperate and boreal forests of western North America,
where background mortality rates have increased rapidly in recent
decades (van Mantgem et aI., 2009) and widespread death of many
tree species in multiple forest types has affected well over 10 million
ha since 1997 (Raffa et aI., 2008). The common implicated causal
factor in these examples is elevated temperatures and/or water
stress. raising the possibility that the world's forests are increasingly
responding to ong01l1g warming and drying.
This paper provides an overview of recent tree mortality due to
climatic water stress and warm temperatures in forests around the
globe. We idemify 88 well-documented episodes of increased
mortality due to drought and heat and summarize recem literature
on forest mortality and decline. From this review we examine the
possibility of emerging mortality risks due to increasing tempera-
tures and drought. Climate as a driver of tree mortality is also
reviewed, summarizing our scientific understanding of mortality
processes as context for assessing possible relationships between
changing climate and forest conditions. Note that while climatic
events can damage forests in many ways ranging from ice storms
to tornadoes and hurricanes, our emphasis is on climate-induced
physiological stress driven by drought and warm temperatures.
The ecological effects of increased mortality in forests and the
associated consequences for human society remain largely
unassessed. We conclude by outlining key information gaps and
scientific uncertainties that currently limit our ability to determine
trends in forest mortality and predict future climate-induced forest
die-off. Addressing these gaps would provide improved informa-
tion to support policy decisions and forest management world-
wide.
2. Methods
This paper emerged in part from collaborations and presenta-
tions developed in special sessions on climate-related forest
mortality at two international meetings: the 2007 annual meeting
of the Ecological Society of America in San Jose, California (Allen
and Breshears, 2007), and the 2008 international conference
entitled "Conference on Adaptation of Forests and Forest Manage-
ment to Changi ng Climate with Emphasis on Forest Health" in
Umea, Sweden (Allen, 2009). In addition to citing contributions
from these sessions, we conducted a systematic search for
published accounts of climate-induced tree morrality since 1970
using the ISrWeb of Science and Google Scholar. We used different
combinations of the key words "tree," "forest," "mortality," "die-
off," "dieback:' "decline," and "drought" in the searches. We also
consulted regional forestry experts to find examples recorded in
government documents and other sources outside the scientific
literature.
From the extensive set of documents uncovered during these
searches, we used two specific criteria to determine whether
the reference was appropriate for this review. Criteria for
inclusion were that the study included: (1) an estimate of area
affected or amount of adult tree mortality at the stand or
population level. based on ground measurements, aerial photo-
grap hy. or remote sensing, and (2) documentation of a strong
correspondence between increases in mortality and increased
water stress or lligh temperatures. We included examples where
biotic agents were involved in the mortality, but eXcluded
examples of fire-driven death. Studies of forest decline or partial
canopy dieback without significanr increases in mortality
were also excluded, as were studies that documented only
seedling mortality. To simplify presentation, we standardized
study descriptors and combined references that describe
impacts of the same event on the same tree species but used
slightly different methods or were conducted at different spatial
scales.
To estimate trends in the literature related to climate-induced
forest mortality, we searched the lSI Web of Science using the topic
words "forest AND mortality AND drought" over the available
interval from 1985 to 2009. We then controlled for increases in the
general scientific literature related to forests by srandardizing the
number of target articles by the number of citations uncovered by a
search using only the topic word "forest."
For each mortality event (listed as rows in Appendix Tables A1-
A6) we tested the association between the forest type affected by
mortality and the categorized duration of the mortality-triggering
drought (seasonal event vs. multi-year drought) with a Chi-square
analysis, comparing number of observed triggering drouglltS (by
drought and foresr type) versus expected number of triggering
droughts. Forest types were grouped into four major biome types
considering similar water limitations: (1) savannas, (2) conifer
forests and Mediterranean woodlands, (3) temperate evergreen
and deciduous forests, and (4) evergreen broadleaved tropical
forests.
662 CD. Allen et al./Forest Ecology and Management 259 (2010) 660-684
Fig. 1. White dots indicate documented localities with rarest mortality related to climatic stress from drought and high temperatures. Background map shows potential
environmentallimirs co vegetation net primary production (Boisvenue and Running, 2005). Only the general areas documented in ehe tables are shown-many addinonal
localities are mapped more precisely on the continental-5cale maps. Droughe and heat-driven [otest mortality often is documented in relatively dry regions (-red/orangel
pink). bU( also occurs outside ehese regions.
3. Results
3.1. Examples of recent climate-induced forest mortality
More than 150 references that document 88 examples of forest
mortality met our criteria of events that were driven by climatic
water/heat stress since 1970. The examples range from modest but
significant local increases in background tree mortality rates to
climate-driven episodes of regional-scale forest die-off. We found
examples from each of the wooded continents that collectively
span diverse forest types and climatic zones (Figs. 1-8 and
Tables AI-A6). Despite our collective efforts to secure references
from non-English language sources, this review is clearly more
comprehensive for North America, Europe, and Australia, and
obviously incomplete particularly for some regions, including
mainland Asia and Russia.
Our searches also reveal that published reports of climate-
related forest mortality in the scientific literature have increased
marl<edly in recent decades. For example, a search ofthe lSI Web of
Science (23 July 2009) using the topic words "forest AND mortality
AND drought" showed 546 references for the period 1985 through
2009, with a steep increase in articles published since 2003 (Fig. 9).
even when standardized for general increases in the forest-related
scientific literature. The years of elevated mortality documented in
the references that met our criteria also show a clear increase in
m011ality events with ajump in 1998 and marked accumulation of
events in the 2000s, particularly the years 2003-2004, Although
these trends could be coincidental or a reflection of greater
scientific imerest in the topic of tree mortality, recent increases in
reported events also mirror warming global temperarures.
3.1.1, Cancinencal-scale summal1es
3.1.1.1. Africa. Increased tree mortality linked to drought and heat
in Africa (Fig, 2; Table A1) includes examples from tropical moist
forest in Uganda (Lwanga, 2003), mountain acacia (Brachystegia
glaucescens) in Zimbabwe (Tafangenyasha, 2001), mesic savanna
trees in South Africa's Kruger National Park (Viljoen, 1995), and
centuries-oldAfae dichatama in Namibia (Faden et aI., 2007). In the
Sahe!,long-term decreases in precipitation linked to anthopogenic
climate change (Biasutti and Giannini, 2006) have caused a die-off
of mesic tree species in parts of Senegal (Gonzalez, 2001),
especially following the severe drought of 1968-1973 (Poupon,
1980). Recent extreme drought in North Africa (Touchan et aI.,
2008) is linked to severe morcality of Atlas cedar (Cedrus atlantica)
from Morocco to Algeria (El Abidine, 2003; Bentouati, 2008: Box 1,
see also Fig, 3).
3.1.1.2. Asia. Reports of forest mortality in Asia (Fig. 4: Table A2)
include death triggered by severe El Nino droughts in 1982/1983
and 1997/1998 in the tropical moist forests of both Malaysian and
Indonesian Borneo (Leighton and Wirawan, 1986; Woods, 1989:
Nakagawa et aI., 2000; van Nieuwstadc and Sheil. 2005). Severe
droughts are also associated with increased mortality among many
tree species from tropical dry forests in northwest and southwest
India (Khan et aI., 1994l, Abies koreana in South [(orea (Lim et aI.,
2008), Juniperus procera from Saudi a Arabia (Fisher, 1997), and
pine and fir species in central Turkey (Semerci et al.. 2008). Recent
droughts have triggered mortality of Pinus tabufaefarmia across 0.5
million ha in east-central China (Wang et aI., 2007), and across
extensive areas of Pinus yunnanensis in southwest China (Li, 2003).
The Russian Federal Forest Agency has mapped zones of forest
health risk ("threat") across the Russian Federation, shOWing 338
million ha as "low threat", 260 million ha as "medium" threat. and
76 million ha of "high" threat, predominantly in southerly portions
of the country (Kobelkov, 2008), where forest health problems due
(0 drought appear to be concentrated (Ermolenko, 2008).
3.1.1.3. Auscralasia. ln the sub-humid environments of nortl1east
Australia (Fig. 5: Table A3), multi-year droughts have repeatedly
triggered widespread Eucalyptus and Corymbia mortality (Fensllam
and Holman. 1999; Rice et aI., 2004; Fensham and Fairfax, 2007),
and have also caused tree death in Acacia woodlands (Fensham and
Fairfax, 2005). There is also documentation of drought-induced
mortality in temperate Nothofagus forests in New Zealand
(Hosking and Hutcheson, 1988).
3,1.1.4. Europe, In Europe (Fig. 6: Table A4), forest mortality due to
dry and warm conditions in the 1990s and 2000s arcs across the
Mediterranean regions, including increased death among many
663 CD Allen et at/ForestEcology and Management 259(2010) 660-684
Fig.2.SatellitemaporArrica.withdocumenteddrought-inducedmortalityareasindicatedwithnumbers.tIedtoTableAI references.Upperphoto:Cedrus atlanlicadie-ofTlIl
BelezmaNational Park.Algeria;2007.byHarounChenchouni.Lowerphoto:quivertree(Aloe dichatoma) morTalityinTirasbergMountains.Namibia; 2005.byWendy Faden.
woodyspeciesinSpain(Penuelasetal.. 2001 ;Maninez-Vilaltaand (Dobbertin and RigJing, 2006; Bigler et al.. 2006; Verrui and
Pinol.2002),increasedmortalityofoak.fir,spruce,beech,andpine Tagliaferro, 1998). A severe drought in 2000 killed many Abies
species in franceaftertIleextremeheatwaveanddroughtduring cephalonica in mainland Greece (Tsopelas etal., 2004) and Pinus
the summerof2003 (Breda et aI., 2006; Landmann et aI., 2006; IlQlapensis sub. bruda-the most drought tolerantofthe Mediter-
Vennetieretal.. 2007).and jncreasesinmortalityofPinus sylvestris raneanpines-ineasternGreece([(orneretal..2005).Farrhernorth.
near the species' southern range limits in SWitzerland and Italy summerdrought paired with biotic stressors has been linked to
Mediterranean Sea
Codru tIa11llCa chlnbulion
Codnis 6taM
_ ""Oeaa
o ""Live
Fig. 3. Map ofnorthern Algeria climate zones and mortalitydistribution ofcedrus arlantica "Box I-AtlasCedarDie-ofTin Algeria" serves asIhe rull caption.
664 CD. Allen et aLI forest Ecology and Management 259 (2010) 660-684
Box 1. Atlas Cedar Die-off in Algeria
Atlas cedar (Cedrus atlantica) occurs in northern Algeria, dis-
tributed in scattered montane populations near the limits of its
bioclimatic tolerance between the Sahara Desert and the Med-
iterranean Sea (Fig. 3). Since the onset of severe drought from
1999 to 2002 cedar forests have undergone mass mortality,
affecting all age classes (Bentouati, 2008). While all Algerian
cedar forests are affected, the magnitude of mortality varies
along a steep moisture gradient (Fig. 3), with die-off greatest
(up to 100%) in the drier mountains nearest [he Sahara, drop-
ping to much lower mortality levels in the moister coastal
mountains (Chenchouni et a!., 2008), Prolonged soil moisture
deficits lead to decline and progressive death of cedar trees
over a period of 1-3 years; a variety of insects and fungi have
continued to kill weakened cedar trees since the drought eased
after 2002 {Chenchouni et aI., 2008), The Cedrus mortality
began assmall patches on drier aspects in the arid near-Sahara
mountains, eventually coalescing into large patches affecting
all ages on all exposures. In contrast, only small patches of old
trees on dry aspects have died in more mesic regions near the
coast. This recent drought also triggered substantial mortality
in other Algerian tree species, including Pinus halapensis,
Quercus ilex, Quercus suber, and Juniperus thurifera. Dendro-
chronological reconstructions of drought in Algeria show that
this early 2000s dry period was the most severe drought since
at least the middle of the 15th century (Touchan et aI., 2008),
consistent with climate change projections for a trend of
increasing aridity in this region (Seager et aI., 2007).
mortality of Quercus robur in Poland (Siwecki and Ufnall<si, 1998),
Piceo abies in southeast Norway (Solberg, 2004), and with a severe
die-off of Picea obovaw in northwest Russia (Kauhanen et al.. 2008;
Ogibin and Demidova, in press).
3.1.1.5. NorthAl71erica. Climate-induced tree mortality and forest
die-off is relatively well documented for North America (Fig. 7;
Table AS). Drought and warmth across western North America in
the last decade have led to extensive insect outbreal<s and
mortality in many forest types throughout the region, affecting
~ million ha and many tree species since 1997 from Alaska to
Mexico (Raffa et aI., 2008; Bentz et aJ., 2009). Examples of forest
die-off range from> 1 million ha of multiple spruce species in
Alaska (Berg et aI., 2006) and>10 million ha of Pinus contorta in
British Columbia (Kurz et aI., 2008a), [0 drought-induced
Populus tremuloides mortality across a million hectares in
Saskatchewan and Alberta (Hogg et al.. 2008), In [he south-
western U.S., die-off of Pinus edulis all over a million hectares
was specifically linked to "global-change-type drought" (Bre-
shears et aI., 2005). In the eastern portion of the continent,
declines and increased mortality among oaks, particularly in the
red oak family, have been reported from Missouri (Voelker et aI.,
2008) to South Carolina (Clinton et aI., 1993) in relation to
multi-year and seasonal droughts in the 1980s-2000s. Drought
during the 1980s followed by an unusual spring thaw in eastern
North America also contributed to decline and mortality of
maples in Quebec (Hendershot and Jones, 1989). In addition,
recent increases in background rates of tree mortality across the
Fig. 4. Satellite map or Asia, with documented drought-induced mon:ality localities indicated with numbers, tied to Table A2 references. LowerR Photo: Dead Abie5 koreana,
Mount Halla, South Korea; 2008, byJong-Hwan Um. Upper Rpharo: Pinu5 tabulaeformis mortality in Shanxi Province, China; 2001, by Yugang Wang. Center photo: Dying Pinus
YUllnanensls in Yunnan Province, China; 2005, by Youqing Luo. Upper Lphoto: Abies cilicicia martalily in the Bozkir-I<onya region, Anatoli., Turkey; 2002, by Orphan (elil,.
Lower Lphoto: Dying Pinus nigra near Kastamonu, AnatoHa, Turkey; 2008, by Akkin Semerci.
665 CD. Allen er all Forest Ecology and Management 2S9 (2010) 660-6&4
Fig. 5. Sarelhre map of Australasia, with documenred drought-induced mon:aliry areas Indicated wirh numbers, tied to Table A3 references. R photo: Die-off of mulga, AcaCIa
aneUIll, the dominant tree across large areas of semI-arid Australia: 2007, by Rod Fensham. l photo: Eucalyptus xanthoclada mortaliry in Queensland, northeastern Australia:
1996. by Rod Fenshaln.
western U.S. have been attributed to elevated temperatures (van warm sea surface temperatures in the North Atlantic, has also
Mantgem et aI., 2009). recently been tied to regionally extensive increases in tree
mortality rates and subsequent aboveground biomass loss,
3.1.1.6. Sourh and Central America. [n Latin America (Fig. 8; indicating vulnerability of Amazonian forests to moisture suess
Table A6), ENSO-relaced seasonal droughcs have amplified back- (Phillips et aI., 2009) (Fig. 9).
ground uee mortality rates in tropical foresTs of Costa Rica
(Chazdon et aI., 2005), Panama (Condit et al., 1995), northwest 3.1.2. Spatial and temporal patterns of mortality
Brazil (Williamson et a1.. 2000). and southeast Brazil (Rolim et aI., Climate-induced mortality events in this review include
2005). and caused extensive mortality of Nothofagus dombeyi in examples that span a broad gradient of woody ecosystems. from
Patagonian South America (Suarez et aI., 2004). A hot and severe monsoonal savannas with mean precipitation <400 mm/year, ro
drought across the Amazon basin in 2005, linked to anomalously subalpine conifer forests with a Mediterranean climate, to tropical
Fig. 6. Satellite map of Europe. with documented drought-indu<Ed morlality areas mdicated with numbers. tied to Table A4 references. R pharo: Pinus sylvestns mon:ality,
Valais. Switzerland; 1999, by Beat Wermelinger. l photo: Pinus sylvestris die-off, Sietra de los Filabres, Spain: 2006. by Rafael Navarro-Cerrillo.
666 CD. Allen et o/./Forest Ecology l1lld Monogement 259 (2010) 660-684
Fig. 7. Satellite map of North Amenca. with documented drought-induced mortali(}' localities indIcated with numbers. tied to Table AS references. Top pharo: Aenal view
showIng severe moftality of aspen (PopulllS cremu/oides) m the parkland zone of Alberta. Canada; 2004. by Michael MichaeJian.lower photo: Pinus ponderosa die-off. Jemez
Moumalns, New Mexico, USA; 2006. by Craig D. Allen.
Fig. 8. SatellIte map of South and Central America. with documented drought-induced mortality localities indicated with numbers, tied to Table A6 references. Phoco:
NDtJiojagus dombeyi mortality at Rio Manso Inferior, norrhern PaL'lgonia. Argemina; 2004. by Thomas Kitzberger.
667 CD. Allen er a!.fForest Ecology and Managemenr 259(2010) 660-684
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Fig.9.151 Webof Science searchofthetrendinpublishedreportsof climate-relatedforestmortaliryin thescientificliterature,fortheyears 1985-2009.Plonedbarsshowthe
percentofreferences usmgthetopicwords "forestAND mortaliryAND drought". relativetoall "forest" references. Line representsthelinearregressionmodelfitled to lhe
data [R'=0.61: F=35.73; p <0.001).
rainforests with mean precipitation >3000mm/year.Thesecases
reveal a complexsetofmortality patternsin response to drought
and heat stress, ranging from modest and short"lived local
increases in background mortality rates to episodes of acute,
regional-scale forest die-off, whichoften(but notalways)involve
biotic agents like insectoutbreaks. At broad spa tial scales.drought-
relatedforestmortalityhasbeenreportednearspeciesgeographic
orelevational range margins where climatic factors (particularly
water stress) are often presumed to be limiting (Allen and
Breshears, 1998; Foden et aI., 2007; Jump et al., 2009; Fig. 1;
Fig. 3andlinkedBox 1).Spatiallyextensivedie-offs arecommonly
associated withprolonged waterdeficits. such as in savanna and
temperate conifer forest vegetation types during multi-year
droughts (Fensham et al., 2009; Fig. 10). Notably, however.
drought-induced mortality is not restricted to forests typically
thought to be water-limited, as highlighted by events in tropical
rainforestsofBorneowherestand-level mortalityreachedas high
as26% afterthesevereEI Nino in 1997/1998(van Nieuwstadtand
Sheil, 2005). or the Amazon basin in 2005 (Phillips etal .. 2009).
Mortality in ever-wet and seasonally dl)' tropical rainforests
appearstoberelativelydiffuseandincitedmostoftenbyshortbut
extreme seasonal droughts (Fig. 10). In temperate forests. short
(seasonal) droughts may be more likely to induce dieback of
broadleaved (deciduous angiosperm) trees (Fig. 10) than conifer
(evergreen needleleaf) trees because of their increased vulner-
ability toxylem cavitation (Maherali etaJ.. 2004).
Patterns of tree death are often quite patchy at finer spatial
scalesacross the synoptic region wheredroughtoccurs. Although
mortalityis sometimesgreatestin locallydl)'landscape positions
(Oberhuber, 2001; Dobbertin et aI., 2005; Worrall et al., 2008),
ecosite variability (soils, elevation, aspect, slope, topographic
position) may interactwitb density-dependentprocesses such as
insect outbreal(s, competition, orfacilitation co produce complex
spatialpatternsofmortalityatthestandandforestscale(Fensham
andHolman,1999;L10retetaI.,2004).Greatermortalitycanoccur,
for example, on more favorable sites within the middle of
geographicand landscapedistributionswherehigher treedensiry
drives increasedcompetition for waterorelevated insectactivity
(Guarin and Taylor, 2005; Greenwood and Weisberg, 2008;
Fensham et aI., 2009; Horner et al.. 2009; Klos et aI., 2009).
25
20'
,..
'5
" c
" ::>
0-
~
10
U-
S .
Savanna Woodlllloc!fConife7 l empeulile Tropical Savanna WoodlilndlComre( Tcmperele 7"ropiCal
Broadleavad BrolJdleoved Broad!-a 8vcd arordleOl't'dd
MuIIi-yeard roug hI Seesonaldroughl
Fig. 10. Dlfferences between observed and expected frequenCies ofreponedforesl mortalitycases listed inTablesAI-A6,sortedbyduration ofassociateddrought eWllls
(seasonalVS. muln-year).with forestsgrouped intofour majorbiomes.Monalltydiscriminatedbyforesttypeisdependentondroughtdnratioll.withmoredrought-adapted
forest rypesshowlIlgmortalitydurillglongdroughtsdndlessdrought-adaptedforest rypesshOWing moremortalirycasesduringshort-termseasonaldroughts.Pearson(111-
square;23.46, df 3.p ~ 0.000012.
668 CO. Allen et al./Foresr Ecology and Managemenc259 (2010) 660-684
However, high severity drought can drive extensive forest
mortality independenc of tree density (Floyd et a!., 2009). Higher
mortality rates can also occur on favorable sites where trees do not
invest in adequate root systems or where they otherwise become
hydraulically overextended (Ogle et aI., 2000; Fensham and
Fairfax, 2007: Nepstad et aI., 2007).
Spatial patterns of mortality at the stand and forest scale are
also heavily influenced by life-history traits and tolerances of
individual species within forests, with drought commonly
triggering differential mortality rates between co-occurring tree
species (Suarez et aI., 2004; Gitlin et a!., 2006; Fensham and Fairfax,
2007: Newbery and Ungenfelder. 2009: Phillips et al.. 2009). Larger
andfor older trees often appear more prone to drought-induced
mortality (Mueller et al.. 2005; Nepstad et aI., 2007: Floyd et aI.,
2009), al though this relationship is species-dependent, and in
cases where stands are undergoing intense self-thinning, smaller
sub-dominant trees and saplings are often more affected (Kloeppel
et aI., 2003: Elliott and Swank, 1994; Hanson and Weltzin. 2000).
Temporal patterns of drought-related tree mortality also can be
difficult to interpret due to lagged responses in some species, in
which mortality has been shown to occur years or even decades after
drought stress (Pedersen, 1998, 1999; Bigler et al.. 2007).
Furthermore, the long-lived nature of trees and their ability to shift
allocation of resources and change their hydraulic architecture
throughout their lives can result in non-linear responses to drought
stress in both space and time. Different sequences of climate events
may also affect the risk of mortality (Miao et aI., 2009).
4. Discussion
4.1. Climate-induced forest mortality-are new trends emerging?
The diverse instances of mortality reported here clearly
illustrate that drought and heat can impact trees in mauy forest
types. However awareness of. and interest in, climate-induced
forest mortality and diebacl< is not new (Auclair, 1993; Ciesla and
Donaubauer, 1994). Past die-offs have been extensively docu-
mented. Historic examples include: Widespread death of Euca-
lyptus, Acacia, and Callirris species in the early 1900s triggered by
the worst drought of the instrumental record in northeastern
Austral1a (Fensham and Holman, 1999); Not/lOfagu.s mortality
during 1914-1915 in New Zealand (Grant, 1984); Picea meyeri
mortality during the 1920s in northern China (Liang et al.. 2003):
extensive tree mortality in the southern Appalachian Mountains
and the Great Plains during the dust-bowl droughts of the 1920s-
1930s (Hursh and Haasis, 1931; Albertson and Weaver. 1945);
Pinus sylvestris death during 1940-1955 in Switzerland (Dobbertin
ec a!., 2007): oak mortality in many European countries following
severe droughts episodes in 1892-1897, 1910-1917, 1922-1927,
1946-1949, 1955-1961 (Delatour, 1983); extensive tree mortality
of Austrocedrus cl1ilens;s during El Nino droughts in the 1910s,
1942-1943, and the 1950s in Argentina (Villalba and Veblen,
1998); and die-off of multiple pine species during the 1950s
drought in the southwestern USA (Swetnam and Betancourt, 1998;
Allen and Breshears, 1998). Furthermore, the overwrought
perception of unprecedented forest decline and impending death
due to air pollution in central Europe (where it was referred to as
'Waldsterben') and eastern North America that received much
attention in the 1980s provides a cautionary example of
exaggerated claims of widespread forest health risk in the absence
of adequate evidence (Skelly and [nnes, 1994).
So are recent occurrences of die-off simply well-documented
examples ofa natural phenomenon linked to climate variability, oris
global climate change driving increases in forest mortality? We
recognize that the available data on climate-induced forest mortality
have many limitations: our examples represent a compilation of
idiosyncratic case studies with uneven geographic coverage. The
studies differed greatly in their goals, methods. and definitions of
mortality, and inconsistently report mortality rates, spatial scale and
panerns of mortality, and severity parameters of climate stress. The
recent increase in forest mortality reports that we document could
merely be an artifact of more scientific attention on climate change,
perhaps in concert with a few high profile cases of climate-related
forest die-off. These limitations. and the lack of any systematic global
monitoring program, currently constrain our ability to determine if
global changes in forest mortality are emerging.
Even though our review is insufficient to make unequivocal
causal attributions, our data are consistent with the possibility that
climate change is contributing to an increase in reported mortality.
Documentation of climate-related forest mortality in association
with recent warming and droughts is rising rapidly (fig. 9), and in
some of these cases the droughts have been the most severe of the
last few centuries. Furthermore, recent research indicates that
warmer temperatures alone can increase foresr water stress
independent of precipitation amount (Barber er al.. 2000). In
addition, new experimental results show that warmer temperatures
can greatly accelerate drought-induced mortality (Adams et aI.,
2009, and associated correspondence). If the recent increase in
mortality reports is indeed driven in part by global climate change.
far greater chronic forest stress and mortality risk should be
expected in coming decades due to tbe large increases in mean
temperature and significant long-term regional drying projected in
some places by 2100, in addition to projected increases in the
frequency of extreme events such as severe drougbts, hot extremes.
and heat waves (IPCC. 2007a; Jentsch et aI., 2007: Stel'l et aI., 2008).
4.2. Climate and plant physiological interactions that drive
forest mortality
Understanding complex spatial and temporal patterns of
climate-induced tree death and forest die-off requires knowledge
of the physiological drivers of tree mortality. The fundamental
mechanisms underlying tree survival and mortality during
drought remain poorly understood despite decades of research
within the fields of forestry, pathology, entomology, and ecology
(Waring, 1987; Manion, 1991; Mueller-Dombois, 1986,1988;
Breda et aI., 2006; Ogaya and Penuelas. 2007; McDowell et al..
2008). Part of the challenge is that tree mortality commonly
involves multiple, interacting factors, ranging from parricular
sequences of climate stress and stand Ii fe histories to insect pests
and diseases (Franklin et a!., 1987; Miao et aI., 2009). Based on the
decline spiral model (Manion, 1991: Manion and Lachance, 1992),
drougbt can operate as a trigger ("inciting factor") that may
ultimately lead to mortality in trees that are already under stress
(by "predisposing factors" such as old age. poor site conditions and
air pollution) and succumb to subsequent stem and root damage
by biotic agents ("contributing factors" such as wood-boring
insects and fungal pathogens). McDowell et al. (2008) build upon
Manion's framework to postulate three mutually non-exclusive
mechanisms by which drought could lead to broad-scale forest
mortaliry: (1) extreme drought and beat kill trees tbrough
cavitation of water columns within the xylem (Rennenberg
et al., 2006; Zweifel and Zeugin, 2008); (2) protracted water
stress drives plant carbon deficits and metabolic limitations that
lead to carbon starvation and reduced ability to defend against
attack by biotic agents such as insects or fungi (McDowell et aI.,
2008: Breshears et aI., 2009; Adams et aI., 2009): and (3) extended
warmth during droughts can drive increased population abun-
dance in these biotic agents. allowing them to overwhelm their
already stressed tree hosts (Desprez-Loustau et aI., 2006: Raffa
et at., 2008; Wermelinger et al.. 2008). Although tbese hypmheses
have groWIng support, our physiological knowledge remains
669 CD. Allen et al./ Fares! Ecology and Management 259 (2010) 660-684
inadequate for confidently predicting patterns of regional die-off,
as well as variation in survival for trees within the same stand.
The degree to which trees regulate water loss during drought may
explain pattems of carbohydrate (and resin) production and
subsequent susceptibility to drought or biotic attack (McDowell
et aI., 2008; Zweifel et al.. 2009). Acontinuum of stomatal responses
to drought exist from drought avoidance (isohydlY), in which
stomata close at a threshold water potential to minimize further
transpiration, to drought tolerance (anisohydry), in which stomatal
closure is less severe and transpiration continues at relatively high
rates (McDowell et aI., 2008). The isohydric response protects xylem
from cavitation through avoidance of severe low water potentials,
but can cause eventual carbon starvation as stomatal closure shuts
down photosynthesis While respiration costs continue to deplete
carbon stores. The anisohydric response can allow continued carbon
gain through maintaining open stomata but at greater risk of
cavitation, which might kill trees directly or could increase the
likelihood of future carbon deficits. Plants that typify eacl1 response
have associated traits consIstent with (heir mode of stomatal
regulation. such as deep rootIng access to more reliable soil water
and cavitation-resistant xylem for drought-tolerant species.
In addition to hydraulic failure and carbon starvation. a third
physiological mechanism predisposing plants to mortality may
exist-cellular metabolism limitation. This hypothesis suggests that
low tissue water potentials during drought may constrain cell
metabolism (WUrth et al.. 2005; Ryan et aI., 2006; Sala and Hoch,
2009), thereby preventing the production and translocation of
carbohydrates, resins, and other secondary metabolites necessary for
plant defense against biotic attack. The common observation that
trees which succumb to insect attacks have weak resin flow and are
unable to pitch out attacl<ing insects is consistentwith constraints on
photosynthetic carbon uptake, cellular carbon metabolism. and/or
treewaterrelations.A likely sequence for many isohydric species that
is consistem with Manion's cascade (Manion, 1991) is that c1imate-
stressed trees starve forcarbon, perhaps due to poor edaphic position
combi ned with drought, which causes poor resin flow and aninabili ty
to defend against insect attack, which subsequently allows fungi that
are symbiotic with the beetles to colonize and occlude the sapwood,
causing transpiration to cease, drying of the canopy, and eventual
mortality (McDowell et at., 2008, 2009).
The observation that climate-induced tree mortality is happening
not only in semi-arid regions but also in mesic forests suggests that
the global rise in temperature may be a common d river (van
Mantgem et al., 2009; Adams et aI., 2009). The mechanisms by which
rising temperature in the absence ofsevere precipitation deficits may
result in increased tree mortality inClude impacts on both host
physiology and biotic agents. Increasing temperature raises the
vapor pressure deficit and evaporation to the atmosphere. This
results in increased water loss through transpiration and either
stomatal closure in the case of isohydric species, or decreased margi n
of safety from hydraulic failure in the case of anisohydric species.
Rising temperatures may impact the carbon storage of trees in a
particularly negative way because the rate of carbohydrate
consumption required to maintain cellular metabolism (respiration)
is strongly linked to temperature (Am thor, 2000). The first
experiment under controlled climate to isolate the effect of
temperature on drought-induced tree mortality, conducted on Pinus
edulis, indicates a high degree of sensitivity to elevated temperature
and indirectly implicates carbon starvation (Adams et al.. 2009).
Warmer temperatures may also be important where cold
winters are usual. in that abnormally warm winter temperatures
maintain significant physiological actiVity a fter the growth
season, with tree respiration costs wasting stored carbohydrates
(Damesin, 2003). Even though CO
2
uptake can occur during mild
winters and partially compensate for carbon loss during summer
droughts (Holst et aI., 2008), the annual Cbalance often remains in
deficit under these conditions. Therefore under climatic warming
scenarios, drought-avoiding tree species may move closer to
carbon starvation, and drought-tolerant species may come closer
to hydraulic failure (McDowell et aI., 2008).
Presumably, surviving individuals after a severe climate event
would have some degree of genetic drought resistance that would
be inherited by the next generation (Gutschick and BassiriRaci.
2003; Parmesan. 2006; Millar et al., 2007a). But the adaptation of a
tree species to a marI<edly different local climate, with only one or a
few generations per century, may be too slow to successfully
respond to the rapid present rate of climate change.
Warming temperawres also have direct effects on insect
population dynamics-in particular, outbreaks of some aggressive
bark beetle species are closely tied to temperature (Logan et al..
2003: Berg er aI., 2006: Hicke et aI., 2006; Rouault et al.. 2006).
Higher temperatures can accelerate insect development and
reproduction, i ncreasi ng infestation pressure directly (e.g., Wer-
melinger and Seifen, 1999; Bale et aJ.. 2002; Caldeira et aI., 2002;
Gan, 2004), while at the same time heat-induced drought stress
may reduce tree vigor and increase host susceptibility to insect
attack (Mattson and Haack. 1987; Rouault et aI., 2006). Warming
temperatures and drougllt-stressed trees also may foster increased
mortality from non-insect pathogens, particularly fungi (Ayres and
Lombardero. 2000; Desprez-Loustau et aI., 2006: Garrett et aI.,
2006). However, fungal responses to climatic factors are complex
and uncertain because of interactions with tree host susceptibility
and insect vectors, and some fungi-tree relationships are difficult
to assess because important belowground interactions between
fungi and tree roots are not well studied.
4.3. ConsequE'nces of broad-scale forest mortality
Due to the increasingly tight coupling of human and environ-
mental systems, the consequences of broad-scale forest mortality
are important to contemplate. Trees grow relatively slowly but can
die quickly: a 200-year-old tree may be killed by severe drought
within a few months to a few years. Therefore, mortality of adult
trees can result in ecosystem changes far more rapidly than a
gradual transition driven by tree regeneration and growth (Fig. 11).
If forests are forced to adjust abruptly to new climate conditions
through forest die-ofT. many pervasive and persistent ecological
and social effects will result. Major changes in understory species
may occur (Rich et aI., 2008), as well as the possible development
of novel ecosystems due to new combinations of native and
invasive exotic trees that. depending on the climatic tolerances of
seedlings. eventually repopulate the overstory (Walther et aI.,
2005; Millar et aI., 2007h; Suarez and Kitzberger, 2008).
Abiotic ecosystem impacts may include changes in solar energy
fluxes reaching ground level and reflecting back to the atmosphere,
with potentially large feedbacks to regional climate in some areas
(Bonan. 2008; Chapin et aI., 2008), as well as alterations in hydrology
and ecosystem water budgets due to increases in evaporation and
reductions in transpiration (e.g.. Huxman et al., 2005). and changes
in groundwater recharge. Potential effects of extensive forest
mortality on water resource availability could have large effects
on human societies (Millennium Ecosystem Assessment. 2005).
In addition, broad-scale forest mortality could change local.
regional, and global carbon budgets (Breshears and Allen, 2002;
Jones et aI., 2009). Forests store considerably more carbon than the
atmosphere, and forest die-off could redistribute within-ecosystem
carbon pools and release pulses of carbon back to the atmosphere. A
recent modeling study simulated this type of transformation in
managed forests of Canada, where climate-related increases in fire
and insect disturbance are forecast to turn these forests into a net
carbon source (Kun et 31., 2008b). Meanwhile, climate-related
increases in the spatial extent of mass tree mortality by insects,
670 CD. Allen etal./Foresl Ecology ond Management259 (2070) 660-684
I
, I
I
Hala'!!! I
8!"'\d
: i
,
Gro\A;th
Time ---+
Fig.11.AbruptreductionsIn forest biomass(orecasysremcarbon)canresultfrom
drought-induced forest die-afT and occur more rapidly than the relatively slow
countervailing biomass incrementsfrom tree natality and growth.Trajectone5 of
change vary wnh ecosystem, as do minimuIII biomass and carbonvalues, and are
notto scale In this conceptual figure.
notably mountain pine beetle. have recently transformed some
forestsofinteriorBritishColumbia(Canada)from anetcarbonsink
intoanetcarbonsource(KurzetaI.,2008a).Similarly,itis possible
that"widespread forest collapsevia drought" could transform the
world'stropicalforestsfromanetcarbonsinkintoalargenetsource
duringthiscentury(Lewis.2006,p. 195;cr. Phillipsetal.,2009:Jones
etaI.,2009).Land-useimpactssuchasanthropogenicfiresandforest
fragmentation, interacting with climate-induced forest stress, are
likelytoamplifytheseeffectsinsomeregions.includingtheAmazon
Basin (Nepstad et aI., 2008). Overall, climate-induced forest
mortalityand related disturbanceswill increaseglobal carbonflux
rates at least temporarily, potentiallyunderminingthe capacityof
theworld's forests to act ascarbon sinks in thecomingcenturies.
Pastforest management may have exacerbated recent mortal-
ity in some regions. [n portions ofwestern NorthAmerica, overa
centulyoffire suppression has fostered thebuildup ofunusually
hig!) tree densities. Trees in these unnaturally dense forests can
have decreased vigor, which can increase their vulnerability to
multiplemortalityfactors(Savage.1997).Extensive reforestations
with pine plantations in regions such as China and the
MediterraneanBasin(e.g., -3.5millionhareforestedwithconifers
since 1940 in Spain alone; J. Castro-from agency statistical
sources)may be particularlyvulnerable,especially becausesome
of these plantations are on marginal sites given the excessive
densitiesand unknown genetic provenances of the trees.
In summary,given thepotentialrisksofclimate-inducedforest
die-off, forest managers need to develop adaptation strategies to
improve the resistance and resilience of forests to projected
increases in climate stress (Seppala et aI., 2009). Options might
include thinning stands to reduce competition, selection of
appropriate genotypes (e.g., improved drought resiscance), and
even translocation ofspecies to match expected climate changes
(e.g.Millareta1.. 2007b;Joyceeta1., 2008; RichardsonetaI., 2009).
4.4. Key informarion gaps ond scientific uncertainties
The conclusions thatcan bedrawn about recent trendsin tree
morrality and the predictions that can be made about future
climate-induced forest die-off are limited by a number of key
information gaps and scientific uncertainties.
(1) Accurate documentation ojglobal Jorest mortality patterns and
uends requires the establishment oj a worldwide monitoring
program. Despitemanynationaland regional forest-monitoring
efforts (e.g.. the European Union's intensive forest health
monitoringEU!lCP-ForestsLevell!network),thereis anabsence
ofadequate data on forest health status globally (FAO, 2006,
2007).Existingpermanentsampleplotnetworkscandetectlarge
scaleeventsorageneralized background mortalityincrease,but
are notdesigned to detectand assess patchymortality.even at
rather high rates, as is common when forest landscapes are
heterogeneousandin mostofthecasesofbioticagentoutbreaks.
Reliable, long-term,global-scaleforest healtbmonitoring,likely
combiningremote-sensingandground-based measurementsin
a mechodologically coordinated and consistent manner, is
needed toaccurately determine che scatus and trends offorest
stressandmortalityonplanecEarth,Regionalandglobalmapsof
aceualpatternsofclimate-inducedcree mortalityarealsovically
important for the development and validation of models for
predictingforest die-offin response to climatechange.
(2) Undersranding tile mechanisms by which climare change may
afJectJorests requires quantitativeknowledge oJthe physiological
thresholds ojindividual uee mortality under chronic or acute
warerstress (Fig, 12). With theexception ofinformation for a
few tree species (McDowell et aI., 2008; Zweifel etal.. 2009),
there is surprisingly little species-specific Imowledge on
regulationof xylemwaterpotentials; therefore,placingvarious
species on the continuum ofisohydly-anisohydry is difficult,
and predicting how diverse species differentially experience
carbon starvation or hydraulic failure is curremly impossible.
Similarly, there is almost no knowledge on the patterns or
mechanisms ofcarbohydrate storage in response to droughc
andheat.Thepotentialeffectsofothercomponentsofchanging
atmospheric chemistry (e.g.. elevated levels of nitrogen
deposition and ground-level ozone) on the sensitivityoftrees
co drought remain inadequately known (Grulke et a1., 2009),
Research is also needed on howtree phenologieswill respond
to climate warming, because increasing winter temperatures
maycontribucetodepletionofcarbohydratereserves relevant
tocarbon starvation thresholds.In addition. betterknowledge
is neededonWithin-speciesgeneticvariabilityandselectionof
trees related to drought and heatstress,
(3) More accurate global vegetation maps are needed as essenrial
inputsto calibrateandvalidaredynamicgfobalvegetationmodels.
The extent of foresc mortality can only be documented or
modeled if there is precise informacion on the locations and
extentof pre-die-offforests.
DroughtDuration
ShOl'l
,'-- 'f'
.c
MortaJlly MORTAUTY
jJ r hord

:;,

<IJ
Q.
E
I! , i'
..!!! NO MORTAliTY
o
Uo _
6
...l
Wetter Drier
Precipitation
Fig. 12. Conceplual diagram. showing range of vanability of "Curreut Climate"
parameters for precipitationand temperature, oralteroanvelyfor drought duration
and Illtensity, WIth onlyasmall portionoftheclimate"space"currentlyexceedinga
species-specifictreemorralltythreshold."FutureClimate"showsincreasesInextreme
draught and temperarure events associaled witl) praJecred global climate change.
indicatingheightened risks ofdrought-lnduced die-ofTfor current tree populations
671 C.D AI/en el all Forest Ecology and Monogemerrr 259 (2010)
(4) Spatially documentotion oj environmental conditions in
areas oj Joresr die-off is necessary to IlJlk mortality to callsal
climare dlivers. including precipitation, temperature, and vapor
pressure deficir. Given the difficulties in measuring precipita-
tion and the absence of reliable soil datasets at adequate
resolutions for continental-scale studies. a robust water
availability index. possibly derived from remote sensing. is
neededtohelpmodelerssimulatewaterstressin trees.Inorder
to disentangle moisture deficit from temperature effects on
tree mortality. more research is also needed to relate spatial
gradients of mortality to variation in temperature. This
research might utilize historical and dendrochronological
recordsacrossspatialandtemporalgradientswherevariations
in rainfall deficit and temperature increase are expressed.
(5) Mechanistic understanding oj climare-induced tree mortality
requires unproved knowledge oj belowground processes and soil
moisture conditiolls (e.g. Brunner et al.. 2009). Models often
include detailed algorithms describing aboveground physiolo-
gicalprocessesbuttreatbelowgroundprocessesasa"blact<box".
Understanding of the impacts of increasing atmospheric CO
2

nitrogen deposition. ground-level ozone, and drought on root

dynamics, productivity, exudation fluxes, and mycorrhizal
interactionswouldparticularlyimprovebelowgroundmodeling.
(6) TIledirecteJJecrs oj climate onrhe population dynamics ojalmost all
Joresrinsect pescs and other biotic disturbance agents remainpoorly
understood but are importallrto modeling climare-induced Joresr
mortaliry (Wermelinger and Seiffert. 1999; Logan et aI., 2003:
Desprez-Loustauetal..2006;BredaetaI.,2006;Bentzetal..2009).
Generalization through synthesis ofcurrent knowledge on (he
dynamICSofdamagingbioticagentsand treeresponsetoattacks
could improveexisting mortalityfunctions in forest models.
(7) Feedbacks between physiological stress (and rreemorrolity) driven
by climore and orller Joresr disturbance processes (e.g., insece
outbreaks,jire) are poorly undersrood (Allen,2007).Thesemajor
disturbance processes may increasingly drive the mortality
dynamicsofforestsinarapidlychangingclimate,necessitating
improved modeling oftheircumulativeand collective effects
(Nepstad eta1.. 2008).
Current models of vegetation response to climate change share
weaknesses associated with the knowledge gaps idenrified here.
including individual tree-based process models (Keane et a!.. 2001),
species-specific empirical models (climate envelope models, e.g..
Hamann and Wang, 2005; Thuiller et aI., 2008), climate envelope
threshold models linked to plant functional types in dynamic global
vegetationmodels(Scholzeetal.,2006).andearthsystemmodels(Ciais
et aI., 2005; Huntingford et aI., 2008). The significant uncertainties
associated with modeling tree mortality are reflected in ongoing
debatesaboutthemagnitudeof die-offrisktoAmazon rainforestsand
boreal forests from climatechangethiscentury, thepotentialfor die-
ofTs inforestsmoregenerally(LoehleandLeBlanc, 1996;PhillipsetaI.,
2008;SqjaetaI., 2007),and thedegreetowhichforests worldwideare
likely to become a netcarbonsourceorsink(e.g.. Kurz etal.. 2008b).
5. Conclusions
Thls overVlew illustrates the complex impacts ofdrought and
heatstress on patterns oftree mortality. and hints at the myriad
ways in which changes in droughtand/or heat severity, duration,
and frequency maylead to gradually increasing background tree
mortality rates and even rapid die-off events. Many recent
examplesofdroughtand heat-related tree mortality from around
the world suggest that no forest type or climate zone is
invulnerable to anthropogenic climate change. even in environ-
ments not normally considered water-limited. Current obseTVa-
tionsofforest mortalityareinsufficienttodetermineifworldwide
trendsareemergingin partduetothelackofa reliable,consistent.
global monitoringsystem. Although theeffects ofclimatechange
cannotbeisolated in these studies andclearlyepisodicfQrest rree
mortality occurs in the absence of climate change. the globally
extensivestudiesidentifiedhereareconsistentwithprojectionsof
increased forest mortality and suggest that some forested
ecosystems may already be shifting in response toclimate.
Therearemajorscientific uncertaintiesin ourunderstandingof
climate-induced treemortality.particularlyregardingthemechan-
ismsthatdrive mortality,includingphysiologicalthresholdsoftree
death and interacrions with biotic agents. Recent advances 111 the
understanding oftree mortality mechanisms suggest that forests
could be particularly sensitive to increases in temperature ill
addition to drought alone, especially in cases where carbon
starvation rather thanhydraulic failure is the plimarymechanism
oftree mortality. However, wecurrentlylack the abilityto predict
mortality and die-off of tree species and forest types based on
specificcombinationsofclimaticeventsandtheirinteractionswi til
bioticstressors and place-specificsiteconditions.The potential for
broad-scaleclimate-inducedtreemortalitycanbeconsideredanon-
linear"tippingelement"in theEarth'sclimatesystem(Lentonetal..
2008).becauseforestd ie-offsfromdroughtcanemergeabruptlyata
regional scale when climate exceedsspecies-specific physiological
thresholds.orifclimatetriggersassociatedirrupDonsofinsectpests
inweakenedforests.Suchcross-scalemortalityprocessesin forests
remain poorlyunderstood.
Collectively. rhese uncertainties currently prevent reliable
determination of actual mortality trends in forests worldwide.
and also hinder model projections of future forest mortality In
responsetoclimatechange.As one consequence,the potencialfor
climatechange to triggerwidespreadforest die-offmaybeunder-
represented in importantassessments to dare, norably including
thelatestmajor(PCe report(2007b). Ifextensiveclimate-induced
tree mortaliry occurs, thell substantial negative ecological and
societal consequences can beexpected.Detenniningthe potential
for broad-scale. climate-induced tree mortality is therefore a key
research priorityforecologists andglobal changescientists.and is
essential for informingand supportingpolicydecisions andforest
management practices.
AcJ<nowledgements
We thank Rebecca Oertel,Andrew Goumas.Angeles G. Mayor,
Russell Fairfax. and Megan Eberhardt Franl< for literature review
assistance; Jennifer Shoemaker for graphics support; and Julio
Be(ancourt, Adrian Das. Dan Fagre. BrianJacobs. Francisco Lloret.
Cynthia Melcher, Catherine Parks. Tom Veblen. and Connie
Woodhouse.and twoanonymousreviewersforcommentson this
paper. Support was provided by the U.s. Geological $uTVey.
BiologicalResourcesDiscipline,Global ChangeProgram(CDA); the
National Science Foundation and Science Foundation Arizona
(AKM); US DOE NICCR DE-FC02-06ER64159 and Biosphere 2-
Philecology (DDB); and Chinese Special Research Program for
Public-Welfare Forestry2007BAC03A02 and 200804001 (ZZ).This
workis a con[ribution oftheWesternMountain Initiative, a USGS
global change research project.
Appendix A
Theseappendix tables (TablesA1-A6)accompany the continen-
tal-scale maps and associated text descriptions. and are the core
compilation ofdocumented examples ofdroughtand heat-induced
tree mortality. Organized by continemand yearofmortality event.
concisely listing key information for each documemed example,
includinganidemificationnumberallowingeasyvisuallinkagetothe
continental-scale map locations.
en
....
'"
Table Al
Documented of dronght and/or heat-induced forest mortal>ty from Afnc.l, 1 10 numbers refer lu m"pped in FIg. 2,
10 Wc.l !I On Year(s) of Forest type/mean precip. ' Domlu"nl tree l.lXa Spatial concentration Climate anumaly Stand/ Scale of Biotic agents Reference(s)d
mortalitY of mortality Within Huked to mort"litY population- impact/"rea associated with
geogr"phic or level affected morta Iity?'
elevational rauge mortalitY
(%)"
Senegal 1972-1973 Sav"nna (300) Acacia Guiero Middle-lower edges of Multi-year drought 50 Regional None Poupon (1980)
senegolensis elevational range:
arid edge of geogra phic
range
2 Sou th Africa 1988-1992 Savanna (366) Caiophospermum mopane Patchy within r"nge Multi-year drought 1'1-87 Not reported Not reported MacGregor and
(Northern
Province)
(basal area) O'Connur (2002)
n
p
3 Zimbabwe 1970-1982, $.av,anna IJrachysteg;o glalicescens; Not repo rted Multi-year Nor reported Subregional; Elephants, Tafangenyasha
).
::::
"
(Southeast) 1991-1992 other savanna species scale insecls (2001. 1998, 1997)
::I

affected
::;.
4 Senegal 1945-1993 Savanna. deciduous Anocoruium aCCldenrale, Arid edges of geographic Multi-year drought 23 Regional None Gonzalez (2001)
.....

btoadleaf woodland Cordylo pinnow, Ficus ingens, range

[240-560) many orhers

5 Sou th Africa
[Nonhero
Province)
1991-1993 Woodland, decidUOUS
broad leaf (500-600)
cinerea,
I'teracarplls angoieflsis.
Strychnos modagoscoriensis,
Terminatia sericea. C. mopane,
Patchy willun range Multi-year drought 1-78
(species-
dependent)
NOt reported None Vlfjoen (1995)
<>
g
Q
g,

many others ::I

6 Sourh Africa 1982-1997 Savanna (24D-500) C mopone, Combrecum Patchy within range Multi-year drought 7 Not reported None O'Connor (1999)
3
(Northern
Province)
opiallarum, Grewia spp.,
X/menta americana
E
...,
Ln
'" 7 Uganda 1999 TropIcal Rainforest (1492) Uvonap,is .'pp.. Celtis spp. Not reported Seasonal drought 19 No r reported Not reported (/.003)
>::;
(Western)
-3
8 Namibia,
South Africa
1904-2002 (10D-200) Aloe dlchoroma Arid edge of geographic
range
Multi-year drought.
high temperatures
2-71 Subregional None Faden et a!. (2007) CT\
0\
c
I
0\
9 Algeria 2000-2008 Med. conifer (348-3S6) [eunJS arlanrica Arid edge of geographic Mulri-year drought 40-80 SubregIOnal Insects Bentouati (2008):
range Benlouatl and
(2006);
Chenchouni
et al. (2008)
10 Morocco 2002-2008 Med. montane COnifer Cedl1J.S atlonrica Arid edge of geographic Multi-year drought 10-40 Subregional Not reported 1 Ahidine (2003);
(300-600) Adil (2008)
forest types are abbreviated as Med. In Ih" column. precipitation IS in mmlyr in parentheses If reported.
h Severily of mortality IS reported 1he stand or population level as percentage of lrees (depending on study design). unless otherwise noted In the entry. Other common Ufll IS are annual mortality during drought (%/
year). percent dead area, and dead wood volume ,n mete,.,'.
< If biotIC agents arc thought to have played a role in tree morlalitY. thiS IS noted in bold type. Ifbiotic agents were involved 10 mortality bUllheir role was nOl evaluated or IS secondary to Climate, Ihe agents are SImply
lisled.
d CitatIOns from which reporred mortality dala is deflved are written In bold lype. Other citarions provide corroborating or secondary eVIdence. If there are multiple citatIOns without no bold type, reported data rellects number<
compiled rrom all CItations.
Table A2
Oocllmeoted casesofdroughtaod!or heat-Iodllred foresr morrality frolll Asia. 1970-I,resent. lD numbers refer to locations ""ppedin 4.
10 Location YeaJ{s) of
moruhty
Fnresttype/me,m
preclp!
Domin.:lnt tree Spatial concentration
ofmortality within
geograplnc or
elevarional range
Climate
anollMly
linked to
mortal,ry
Stancl/
populatlOn-
level mnrtality
(%)"
Scale of
impact!
area affected
BiOtic agents
associated
with
mortality?'
Reference(s)d
Sri Lanka 1976-1980 Monlane tropical
rainforest
Calaphyllum spp..
Sy"",gium <pp.
Upper-mid
elevational range
Seasonal
drought
50-100 NOl reported Not reported Werner(1988J
2 Malaysia
(Borneo)
19B2-1983 Tropical ralllforest

Dipreroca'1Ju$ spp..
Shorea spp,
Nor reported Seasoll"1
drought
12-28 Not repnrted Not reported Woods(1989); Beckerelal.
(199B); leighlonand
Wlrawan (1986)
3 India (Cujaral) 1987 Tropical dry deciduous Acacia senegal.
Halarrhena
nnlidysearerica,
Helieteres isora,
Not reported Seasonal or
single-year
droughl
37-82(spCCles-
dependen,)

affected
Ungulates
(Cervu.< trnirolor)
Khan eta1. (1994)
Terminalin crenu/ara,
others
'i
'='
4
5
Russia (Far
East)
Saudi Arabia
and Oman
1987-1988
E.1r1y 1990s
Montane mixedconifer
Woodland (55!'!)
Picea jezoensi5. Abies
nephmlepis
Jun/penlS procem.
j. excelsa
Mountain slopes
and plareaus.
variable aspects
Lower edges of
elevational range
Drought
Possibly
drought
1qMm' timber
lost
30 U exce/sa)
ha
affecled
Landscape-
subregional
Fungi
NOlle
Man'koand Gladkova (2001)
FisherandGMdner(1995): Fisher
(1997): Gardnerdnd Fisher
:..

"
'"

g.
-..


6 Indonesia
(Snmarra)
1997-1998 Tropical rainforest Nor reported Not reported Seasonal
drought
9.8 Not reporreu Not reported Kinnaird'IldO'Brien. 1 '"
"

7 Indonesia and
Malaysia
(Borneo)
1997-1998 Tropical ralnlorest
(-2100-3000)
Dipterocorpus spP..
Lauroceae
Not reported Seasonal
drooght
0.6-26.3 Not reported Not reported vanNleuwsladtand51,eil (200S):
Potts(2003); Aiba and Kitay,lnl,'
(2002); 5hk(2004)
c
"
Q.

'"

B
9
10
Indonesia
(Borneo)
MalaySia
(Boroeo)
China (Shanxi.
Hebei, Henan)
1997-1998
1997-19!'!8
1998-2001
Tropical lowland
,wamp(2800)
TropICal l',lnforest
(-2700)
Temperate coniferous
plantation
Anacardiacea,
Oipterocarpaceue,
Sapatace"e.Rutaceae
Diprerocarpoceae,
Euphorbioceae.
BurseraceQt:',
Myristicaceae
Pinus IObulaeformia
Not reported
NOI reported
Not reporred
Seasonal
drought
Seasonal
drought
Seasonal
drought
4.2-6.1
0-6.4
-30
Nor reported
Nor reponed
Subregional:
500.000ha
affected
Not
Not reported
Bark beetles
(Dendroctonus
valens)
Ni,himuaeral. (2007)
Nakagawa etal. (2000);
Lingenfelderand
Newbery(2009)
Wang et al. (2007)
O<l
3
"
"

'"
\.n
ID
'':;
:::


'"
1
&l
...
11 China
(Yunnan)
1986-1988.
1998-2000:
2003-2005
Subtropical coniferous
plantauon
Pinus Nor reporled Seasonal
drought
Vaned in
different
planlations
lAndscape:
113.000ha affected
Bark beelIes
(Tomicus
yunnancnsLs,
L. (2003)
T. minor)
12 Turkey
(Centr.1
Anatolia)
2002-2007 Temperate coniferand
mixed (400-600)
Qercus .. Juniper
spp.. Pinus nigra.
p, sylvesrr;s.
Abies cilicicio
Soothern edge of
geographiC range
for P. sylvesrris
DrouKht Not reported Nor repurted Insects Semerci el al. (2008)
13 South Korea 2003-2008 Temperate montane
mixed(1400-2000)
Abies koreana Not reported Warm winters!
sprinKs. pOSSIbly
drought
20-50 lAndscape NOI reported Lim et al. (2008):
Wooe1 al. (2007)
1q Ru>sia 2005-2008
Footnotes.sin Table A1.
Boreal alld temperate Picea spp., Pinus spp. Southern portlOIlS of
Russian foresl xones
DrouKht Not reported >400.000ha
Jfross the na [ioll
Nor reported Ermolenkn (2008)
en
'"
'"
Table A3 '"
-.J
Documented(a,esofdrought andlorheat-induced forest mnrlallty from Aus,ralasia, 1970-present. ID numbers refer to locationsmapped III Fig. 5. "
ID Location Year(s) of Forest type/ Dornlnanl lree SpatialConcentration Climate St.lnd/ Scale of,mpact/ Biotic agents assOCiated Reference(s)"
mortality mean precip.' [axa ofmortality within anomaly linked population- area aflected with mortahty?'
geographic or to mortality level
elevational range mortality (%)b
New Zealand 1978-1980 MOlltane Nothofagw;fusm Not reported Spring droughts 75 1.1Odscape; Beech scale(lnglisiafagi); Hoskingand Kersh,'w
(WestCoast) broadJeaf -50UOh,1 fungi (Hypocre/lo duplex); (1985)
.1ffected Wood borer(Platypus spp.,
Psepholax spp.)
2 New Zealand 1984-1987 Montane Nothofogu.s solandri Not reported Springdroughts 24-52 Not reported Lcafminer(Neomycta Hoskingand Hutcheson
(Hawkes Bay) broad/ear pulienris); FungI I ~ B B
(Nodulisporium spp.)
3 Australia 1992-1996 Tropical savanna Eucalyptu.s spp.. Patchy within Multi-year drought 29(basal area) Regional; None Fensh<lm and Holman
(Queensland) (480-2600) Corymbia spp. ranges ~ 5 . s M ha (1999); Fensham (1998);
affe([ed Rice et.11. (2004) r;
<:J
4 Australia 1990-2002 Tropical savanna Eucolypt.us sPp.. P.lIchy withlll MultI-yeardrought 78 sland level; Regional; None Fensham etal.
~
(Quf"ensland) (500-850) CorymblO spp. ranges 17.7 across -5.5M ha (2003, 2009) ..
region affected "

'"
c
~
5 Australia 2004 Tropical savanna Eucalyptus spp., Patchywithin MultI-year drought 15.0(basal area; Not reported None Fensham and Fairfax -....
(Queensland) (500-850) Corymbia spp. ranges unpublished data) (2007) 6'
~
6 Australia 2005 Tropical savanna AcaCia spp Widespread MultI-yeardrought Not recorded -60Uh" None Fensham and Fairfax
(Queensland) (500-850) affected (2005) [
Footnotes areasgiven In Tahle AI. ~
c
c.
;;:
TableA4 "g
Documented cases ofdrought and/or heac-induced forest morlalityfrom Ellrope. 1970-present. ID numhe" refer to10G.)[ions mapped III Fig. 6. ~
~
"
ID Location Year(s)of Forest typel Domin.ant tree Sp,1tial concentration Climate Standi Scale ofimpact/ Biotic agemsassociated Reference(s)"
"-
mortality me.ln precip.d taxa ofmortality within <lnomaly Itnked population- area affected w](h monality?< v,
'"
geographic or to mort.>lily level
<0
t:;
elevacional range mortality (%)b 0
-3
Switzerland 1960-1976 Temperate PiTlUS sylvestl15 Lowerlsouthern edges Multi-year 5-100 Landscape- Not reported Kienast etal. (1981)
U1
(Valais) conifer (572) ofranges drought suhregional 0
'"
I
U1
2 Europe 1970-1985 Tempecate Abies spp.. Picea Lower edges of Repeated 1-20 Regional: Barkbeetles(Sco(yrus. Schuttand Cowling(1985) ....
'"
(Western, cOlllfer and spp.. Pinus spp.. elevation range droughts patchy across Ips, PiryogeTlrs, Tomicus.
Central) broadleaf Fagus sylvotjca <IMha DeTldrochtonus,
(600-1500) f'yriokreines); Fungi
France 19801985 Temperate Q!Jercus spp.. Patchy across ranges Seasonal or 10-50 SubregIonal: Fllngi; b.lfk beetles Nageleisen (1994);
broadleaf mainly Qrobur single-year patchy across (Agriles. Sco/yrus) Nagelelsen etal. (1~ ~ I):
(b50-850) drought -SOO,OOOha Delatour(1983)
4 Poiand 1979-1987 Temperate Quercus robut Not reported Seasonal 111,000m' Landscape Moths ("rortrix viridiana); Siwecki and Ufnalksl
llroadleaf drought timber loS[ pathoKens (199B)
(500-550) (Opilioswma spp.)
5 Greece 1987-1989 Mediterranean Ab,es albo Mill. Middleofelevation Multi-year 1.8/yr >n Landscape- Rark heetlcsand other Markalas ("1992); Kailidis
mixed conifer xA cepha/onrca ranges drought drought SUbregional insects and Markalas(1990)
(1622) Loud. years
6 Italy(Sourh 1992 Temperate Pmu.s .<ylvrsrris Lower/southern edges Multi-ye..1I' Not reported Landscape- V(uiolls insecrs MlI1erbl (1993)
Tyrol) mixed conifer ofranges drought subregional
(650)
7 Austria (lower 1990-1996 Temperate mixed PInus sylvestris, lower edge of Seasonal 27.6-49.2 Stand-Ia ndscape Various insects Cech and Tomlczek (19%)
Austna) conifer (-G50) Pmus nigra elevational rauge draugbts
8 Austria (TyrOl) 1991-1997 Temperate mixed I'inus sylvestr'is Lower edge of Seasonal 10.0-70.0 l.lndscape Various insecl5 Cech and Perny (2000)
conifer (-840) e1evatlonal range drooghts
9 Italy (AoSta) 1985-1998 Temperate mixed fJllWS o;ylvestris lower/southern Multi-year Nor reported wllc1scape- Fungi (Arm iI/ana spp.); Verrui and TaghJferro
con,fel and edges of range> drought subregional wood borers (1998)
broadleaf (-550)
10 Spain 1994, 1998 Mediterranean Quercus spp.. Patchy w;thi n Multi-year 0.0-19.4 wndscape- Not reported Peiiuelas et al. (2001);
{Nartheast, mixed coni fer /'lnus spp., elevalional range: drought, (spec.es- subregional L10ret and S,scar! (1995);
central, South) and broadleaf Juniperus spp. southern edge of recorrent dependent) L10ret et al. (2004);
(537-605) geographic range summer Martlnez-Vilalt. and
(I'. sylvestris) droughts Pinal (2002)
11 France lY98 Mont.llle mixed Fagus sylvonco Middle of ranges Ueep frost 5-30 Subregional: None French Forest Health
(Ardennes. conifer and after an palchy across Department (1998-1999)
Vosges) broadleaf abnoma/ly hal _200,000 ha
(800-1200) penod
"
12 Norway 1992-2000 Temperate Piceo ubles Patchy across ranges Multi-year 2-6.6 wndscape- Bark beetles Solberg (2004)
0
'" conifer summer
droughts. higb
subregIonal ( Polygraphus
pa/igraplrus)
"'
"

summer
~
temperatures ....
~
13 Greece
(Samos)
2000 Mediterranean
mixed conifer
~ 700-BOO)
Pmus bnJlia Lower edge of
elevationa I range
Multi-year
drougbt
Not reported No l repa rted Not reported Korner et al. (200S):
Sams er aJ. (2007)

::;'
0
0-
14 Auslria (Tyrol) 2001 Temperate
mixed coni reI'
PinllS sylvest.ris lower edge of
elevatianal range
Seasonal
droughts
Not reported ldndsc.Jpe-
subregional
Not reported Oberhul'er (2001)
~
Q
Q.
(710) ;:
!2
IS Greece 2000-2002 Mediterranean Auie. Not reported Multi-year 5-1 O/yr in landscape Primary role, bark Tsopelas et al. (2004):
Q
o.
(Soulh, mixed conifer cepha/oniea drought drought ye,rs beetles (Phaenops Raftayanm> el al. (20U8)
"' 3
Central) (-700-1100) vs. 0.17-0.50/ {moreki, I',cyokteincs
~
16 Switzerland 2003 Temperate Picea auies Not reported Drought. high
yr in non-
dronght years
-2.0Mm' wlldscapc-
sp,mdens) mistletoe
Bark beetles ForSler et al. (2008)
""
'" 'J:>
-.:;
'"
conifer and
broadleaf
temperatures timber lost subregional (Ips cypographus)
~
Ol
Ol
'"
I
17 Switzerland
(Val,.s)
1973-1976.
1987-1993,
Temperate
mixed conifer
Pinus sylvescris lower/southern
edges of I',lOKes
Seasonal ;lOd
molti-year
7-59 h,"dscape-
subregion,1
Primary role, bark
beetles (Phaenops
Wennelinger er al. (2008);
Dabbertin el al. (2007).
CO)
'"
...
1996-2000. and braadleaf droughts. high cyanea, Ips ocum;natus); BiKler et al. (2006);
2000-2004 (500-600) temperarures nematodes; mIstletoe DobbertlO and Righng
(2006); Itigli Ilg et al. (2006);
Dabber( in er al. (2005);
Rigling alld Cherubini (1999)
18 Germany 2003-2U06 TemperMe fagus sylvatica Not reported Draughl, high -98.000m' landscape- Bark, ambros.a beetles Petercord (2008)
(Baden- braadleaf tempera tures timber lnst subregiollal ('faphrmychlls blcalor.
Wurttemberg) 'frypadendrotl domesr;cull1);
wood borer
19 Spain 2004-2006 Temperate conifer Pinus sylvestris, Not reported Multi-year Not reported Patchy across Nut reported Navarra-Ccrrillo et al.
plantations Pinus nigra drougbl 13,404ha (2007)
20 Russia 2004-2006 Boreal conifer Picco obavaw Patchy Drought, higb 208 Mm' -1.9 Mha Bark beetles Krorov (2007); Tsvetkav
(Nortbwe.st) temperatures limber lost ,ffecte<! (Ips cypagroplras). fungi '11<1 Tsvetkov (2007): (huprav
(2007): Shlrakhov (200B):
Kauhdnen et al. (2008) 0>
-..0
V>
Table A4. (Continued) ~
Ol
ID location YeM(S) of forest Iypel Dominant tree Spatial concentration Climate Standi Scale of imp.et/ 810tic agents associated Reference{S)d
mortality mea n precip:' tdxa of mortality wIthin anomaly linked population- area arlected with mortality?'
geographic Or to mortality level
elevalional range mortality (X)"
21 SWItzerland 2003-2007 Temperate mixed Pinus sylvesr,;s lower edge of Orollght, high 6.3-16.0 landscape- Not reported Schill' el al. (in press)
(Grisons) conifer (750) elevational range temperatures sublPgiunal
22 ~ r n e 2003-2008 Mediterranean Pinus sylvestris lowerisollihern edges MUlti-year 20-80 Subregional; Bark beetles Vennet'er el al. (2007);
(Provence, conifer of ranges drought, h.igb patchy across (Tom;eus, Ips, Pissodes) Thabeet et al. (2009)
Southern Alps) (750-950) temperatures _1 00,000 ha
23 France 2003-2008 Temperate Quercus sp p. lower and middle of Spring and 1-3/yr Regional Bark beetles; fungi Breda et al. (2006);
mixed conifer Fogu.< sylvorico, eleva tiona I range summer wne"nann et al. (2006);
and bruadleaf Abies spp., druught, Rouault er al. (2006);
(650-1100) Piceo abies, scorchong heat french Foresr Healrh
Pinus spp DepalT.ll1ent (2003-2008)
24 France 2003-2008 Temperate Abies albo lower edge 10 middle Recurrent 10-30 subregional; Bark beetles French Forest Health
n
>='
(Eastern mIxed conifer of ranges drnllght, high palchy across (Ips, fusodes) Departmenr (2003-2008) :to
Pyrenees) (800-1000) tempefJtures -150,000ha
~
;:>
"
25 France 2006-2008 Mediterranea n Quercus SUber Northern edge to MultI-year 10-70 Subregion.ll; Insects (Platypus spp.. Vennet,er el al. (2008)
~
(Provence.
Maures
broadleaf middle of geographic
range
drought patchy across
-120,000ha
Coroebus spp.) --...
ti'
Mount.:>ins) ~
Footnotes as In Table A1.
8
'f
c
" Q.
~
Table AS
;:>
Q
Documented cases of drollgbt andlor heal-induced forest mortality from North Amenc.,. I970-present. ID numbers rerer to lootlol\< mapped in r.g. 7
ID location Ye.,(s) of Forest type' Dominant tree spatial Climate anomaly Standi Scale 01 Bio tic agents Reference(s)"
~
ii
"-
mortality mean precip.' taxa co I1centralion
of mortality
linked to mortality population-
level
impact/area
affected
associa red wllh
monaliry?L
tv
'"
'"
wHniu
geographic or
mortality (%)" ~
2
USA (50lltheast, late 1970s- Upland Quercus sPp..
elevational range
NOl reported Mufti-year droughts; 16.6 in stands Regional Wood borers Stringer et al. (1989); Starkey

I
~
Northeast, Midwest) 1980s temperate Cayro spp. high remperatures across Southeast; (Agrilus and Oak (1989); Starkey et aL
....
mixed preceded hy severe 1.2-6.3 in MIssouri bilineacus); (1989): Clinton et al. (1993);
winters fungi; insecr Millers et al. (1989); Tainter
defnhalors er al. (1983); law Jnd Gort
(1987); Ke.<sler (1989); Jenkins
and Pallardy (1995)
2 USA (Midwest) 1984 Temper.lte AcP.r spp. Not reported Drought Nut reported landscape- Wood burers Millers et al. (1989)
deciduous subregional (Agrilus spp.)
3 USA (MIdwest) 1979-1986 Temperate Belilia spp. Not reported Multi-year drought Not reported landscape- leafminers; MIllers et at. (1989)
deciduous subregional wood borers;
bitch
skeletonlzers
4 USA (North Carolma) 1984-1989 Temperate Acer \Uc.dlOrum, No t re ported Multi-yea, drought 1.0-3.25/yr. in Not repol reel Not reported Olano ami Palmer (2003)
deciduous Fugus grondifolro, drought year<
( 1270-1520) TUu] americuna,
Aesnllus ./lava
Table AS (Continued)
ID Location Yearts) of
mortality
Foresr type!
mean prccip.'"
Dominan{ lfee
IrtXa
Spatial
concentration
ofmort.>lity
wirhin
geographic ur
elevarional range
Chmate anomaly
linked to morLllily
Stand!
popularion-
level
mortality (X)
Scale of
Impact/;"ea
affected
Biotic agems
assocl;lted with
mottality?'
Referencer S)d
5 USA (Minnesola) 1987-1989 Savanna (726) QlJercus ellipsoidalis,
Q mocrocarpa
Not reported Mulll-yeal druught 18,2 Not reported Not reported Faber-Iansendocn and
Tester
6 East ern North Amenca 1980s Temperate
deciduous
(900-1200)
Ace' saccharum Patchy within
ranges
Drought. high
lemperarures
preceded by wlntel
thaw
10-15 Subregional:
parchyacross
>1 Mba
Insect defoliator
(Malocosoma
disstria)
Hendershot and Jones (1989);
Payette et al. (1996): Auclair
el at (1996): Kay et al (2004):
Robilaille el at (1982)
7 USA and Mexico
(C.lhfomia and Baja
California)
8 USA (Califorma)
1985-early
1990s
1986-1992
Montane
mixed conifer
(-600-800)
Montane
mixed cunifer
(945)
Pinus jeffreyl,
Abies co ncOlor
Pinus ponderosa,
Caiocedf1Js decurrens,
Abies cOllc%r
Not reported
Not repOited
Multi-year drollght
Multi-year drought.
high spring and
summer remperatures
4-15
23,3-69,2
Landscape-
subregional
Landscape
Bark beetles
(Dendroceonus
spp,)
Bark beetles
(Dendroceonus
spp.)
Savage (1
Guarin and Taylor (2005) "
"
).

"

9 USA (Califonlla) 1986-1992 Montane

mixed conifer
Not reporred Not repotted Multi-year droughl 13 (basal area) Landsc.,pe-
subregional:
56,OOOha
affected
Engraver
beetles (Scolycus
spp.)
Macomber and Woodcock
(1994)
"
r-
....
;;>


10 USA (Califorma) 1986-1992 Montane
mixed cunifer
Pinus spp., Ah,es spp. Drier edge of
local range:
lower edges
of elevariona'
ranges
Multi-year drought Nor reporled Landscape-
subregional
Primary role,
bark beetles
( Dendrarlonus
spp.): engraver
beetles (Scolytus
spp.)
Ferrell et "I (1994):
Ferrell (1996)
0
0-

"

Q.
;::
0
:J

11 USA (Cahforn,a) Montane
mixed conifer
PinllS flexills Lower edges
of elevational
range
Multi-year drollght,
high temperatures
50-75 Stand-
landscape
Mistletoe
(ArceutllObium)
bark beetles
(Dendrocronus
panderosne)
Millar et (200ia)

;;;
"
'"
t.n
I<l
to
'"
2
12 USA 1996 Woodland
(-370)
Pinus edulis,
jumpPf1Js
monospermQ
Patchy within
range
Single-year dronghl 2.3-2S.9 Landscape-
subregional
Not reported Mueller et al. (2005):
Ogle et al. (2000): Trotter (2004)

'"
I
&l
...
13 Canada (Alberta) Boreal rarest,
prairie ecotone
(450)
Populus rrenwloide> Patchy withlO
ranges
Drought preced IllS
warm winter and
spring
18-47 Subreglunal:
patchy across
-1 Mha
Insect defolialor
( Ma/ocosama
disstria)
Hogg et aL (2002)
14 USA (Midwest.
Southeast)
1990-2002 Upland
temperate
mixed
Quercus spp. Patchy within
rang'"
Multi-year dluuglll 1S-50 basal
area reduction
Regluna):
-1.8 M ha
affecred
Wood borers
(fnaph%des
rututus. Agrilus
,pp.): fungi
Starkey et at (2004):
O.lk et al. (2004): VnelkPl
et at. (200H): Heilzman
et ai, (2004): Lawrence
et aL (2002)
15 USA (California) 1983-2004 Monlane mixed
cOlllfer
(1100-1400)
Pinus spp.,
Abies spp.
Lnwer edges
of elevational
range
Drought, high
tern peratures
63% increase
in annual
mortality rotc
Landscape-
sUbreglUlILlI
Insects,
pathogens
van Mantgcm and
Stephenson (2007)
16 USA and Canada
(Alaska, Yukon)
1989-2004 Coasral rainforest,
boreal (4HS)
Picco spp. Palchy within
ranges
Oruught, high summer
temperatures
Not reported Subregional:
> 1.2 M ha
Primary role,
bark beetle
(Dcndroctonus
rujipenni., )
Derg et aL (2006)
a>
...,
...,
AS (Collflllued)
."
....
co
10 Location of forest lype/ Dominant tree Spatial Climate anomaly Stand/ Scale of Biotic agents Reference(s)"
mortality mean precip.' taxa concentration linked to mortahty population- Impact/are> associated wi th
of mortality level affected mortality?C
within mortality (%}b
geographic or
e1evational rilnge
17 USA (Southwe<lj 2000-2004 Woodland, Pinus ponderoso, Patchy within Multi-year drought 3.3-41.4 (spedes ' ... ndscape- Not reported Gitlin e[ al. (200G):
cOllifer Pinus edulls, elevational range dependant) subregional Burkett el al. (200S)
,/ul1lperus
monospermo,
Populus spp
18 Somhwest. USA 2000-2004 Woodland Pinus eduli.,. Pinus Patchy within M1111 i-year drought, region-wide, Regional; Primary role, Breshears et al. (lOOs):
(New MexIco, (200-450) monophyllo, geographIC high sprlllg and 0-90 stand-level -1.2Mha ba rk beetles Shaw et al. (2005); Swaty
Arizona, Color.1do. ./uniperus elevational range summer for Pinus spp.; afli"cted (Ips conrusus); et al. (2004); Mueller et al.
Utah, Nevada) monosperma, 4.5 stand-level for twig beetles; (2005); Allen (2007).
./uu,perus J. /TIonospermo pitch mothS; Greenwood .111d Weisberg
n
scopulorum root fungus; (2008) r:o
mistletoe
..-
" 19 USA (Amona) 2001-2004 Coniferous (lS0) Pinus ponderosa Lower edges of Multi-year drought, 7 -21 Landscape- Primary role, Negron et aI. (2009)

elevational range high temperatures subregional bark and
e.
engraver
beetles (Ips spp.)
'--
ti'


20 Canaela
(Saskatchewan
2002-2004 80real forest,
prairie eCOlone
Popllius
ere'" uloides
Southem edge
of geographiC
MUlti-year drought 3,6/yr vs. 1.6/yr
III non-drought yrs.
SUbcOO[lIlemal;
patchy across
Insect defoliators Hogg et al (200S)
'"
,.,
"0'
and Alberta) (360-4GO) range -10Mha
21 Canada (Bntlsh Montane mixed Pinus contorta Middle of Droughr. high spring >435Mm' Regional - Prim.ary role, Kutz er al.
e
ii
Columbia) conifer geographic range and S\l/llmer (timber lost) contmenr.11: bark beetle
" {empera tures 13 M ha affected (DendroCConus
"
"
22 USII (Color.1do) 2005-2006 Montane mixed PopUlUS lTemuloides Patchy but Multi-year drought, 32 (stand scale); Landscape-
ponderosoe)
Wood borers; Worrall et al. (2008)
'" [3
..
"-
(380-1100) concentrated at high sptlng and 5.G2 (landsc.3pe subregional: cytospora
'"
In
lower edges of
elevational range
summer temperatures. s(ille) 58,374 ha affected canker: bark
beetle.1
'"
"
0
23 USII 1955-2007 All western Ma ny species Not reported High temperatures 3.9-rold increase Regional Not reported van Mantgem et al. (2009)
-2
."
(Western States] forest types in annual mortality
rate
0)
0
I
'"
0:>
....
24 Wester" North 1997-2007 Conirerous PinUS spp., Picea Not reported Drought, high NOl repo rted Regional; Primary role, Bentz et al. (2009)
America spp., Ables spp., temperatures 60.7Mha bark and
Pseudotsugo affected engraver beetles
menzies;; (DendroctotlUs.
Ips. Dryocoeres,
Scolytus spp.)
25 USII (Mmnesola) Boreal and l'opulus Lower edges Drought Not reported Nol reported Insect defoliators Minnesota Dept. Na r.
temperate mixed eremuloldes, and mir1dle of Resou rreS (2007)
(480-900) fraxillus spp. ranges
26 USII (California) 1998-2001, NOl reported Not re ported No t leported Droughl preceded 423,000 dead Iress Landscape- Primary role. GMletl el al (2006)
2005-2008 by wet, warm episodes in northern California s\lbregional pathogen
(Phytophrlloro
ranorum)
27 Canada and USA Long-term Temperate coastal Cilamuecypal;s Middle Warmer winters 70% of basal area lost Snbregional: None Beier et al. (2008); Hennon
(Alaska. 1880-2008 rainrotest nootkatenSlS and springs -200.000 ha and Shaw (1997); Hennon
Btltish Columbia) (1300-4000) allecled el al. (2005)
foolnotes as ,n rabl. AI.
679 CD. Allel! et al./Forest Ecology and Management 259(2010) 660-684
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