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Altered dynamics of forest recovery under a changing climate


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Forest regeneration following disturbance is a key ecological process, influencing forest structure and function, species assemblages, and ecosystem-climate interactions. Climate change may alter forest recovery dynamics or even prevent recovery, triggering feedbacks to the climate system, altering regional biodiversity, and affecting the ecosystem services provided by forests. Multiple lines of evidence-including global-scale patterns in forest recovery dynamics; forest responses to experimental manipulation of CO2 , temperature, and precipitation; forest responses to the climate change that has already occurred; ecological theory; and ecosystem and earth system models-all indicate that the dynamics of forest recovery are sensitive to climate. However, synthetic understanding of how atmospheric CO2 and climate shape trajectories of forest recovery is lacking. Here, we review these separate lines of evidence, which together demonstrate that the dynamics of forest recovery are being impacted by increasing atmospheric CO2 and changing climate. Rates of forest recovery rates generally increase with CO2 , temperature, and water availability. Drought reduces growth and live biomass in forests of all ages, having a particularly strong effect on seedling recruitment and survival. Responses of individual trees and whole-forest ecosystems to CO2 and climate manipulations often vary by age, implying that forests of different ages will respond differently to climate change. Furthermore, species within a community typically exhibit differential responses to CO2 and climate, and altered community dynamics can have important consequences for ecosystem function. Age- and species-dependent responses provide a mechanism by which climate change may push some forests past critical thresholds such that they fail to recover to their previous state following disturbance. Altered dynamics of forest recovery will result in positive and negative feedbacks to climate change. Future research on this topic and corresponding improvements to earth system models will be key to understanding the future of forests and their feedbacks to the climate system. © 2013 Blackwell Publishing Ltd.
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Altered dynamics of forest recovery under a changing
*Conservation Ecology Center, Smithsonian Conservation Biology Institute, National Zoological Park, Front Royal, VA, USA
Smithsonian Institution Global Earth Observatory-Center for Tropical Forest Science, Smithsonian Tropical Research Institute,
Panama, Repu
´blica de Panama
´Institute for Genomic Biology, University of Illinois, §Odum School of Ecology, University of
Georgia, Athens, GA, USA Energy Biosciences Institute, University of Illinois, at Urbana-Champaign, Urbana, IL, USA **Plant
Biology, University of Illinois, at Urbana-Champaign, Urbana, IL, USA ††US Dairy Forage Research Center, USDA-ARS,
Madison, WI 53706, USA
Forest regeneration following disturbance is a key ecological process, influencing forest structure and function, spe-
cies assemblages, and ecosystemclimate interactions. Climate change may alter forest recovery dynamics or even
prevent recovery, triggering feedbacks to the climate system, altering regional biodiversity, and affecting the ecosys-
tem services provided by forests. Multiple lines of evidence including global-scale patterns in forest recovery
dynamics; forest responses to experimental manipulation of CO
, temperature, and precipitation; forest responses to
the climate change that has already occurred; ecological theory; and ecosystem and earth system models all indicate
that the dynamics of forest recovery are sensitive to climate. However, synthetic understanding of how atmospheric
and climate shape trajectories of forest recovery is lacking. Here, we review these separate lines of evidence,
which together demonstrate that the dynamics of forest recovery are being impacted by increasing atmospheric CO
and changing climate. Rates of forest recovery generally increase with CO
, temperature, and water availability.
Drought reduces growth and live biomass in forests of all ages, having a particularly strong effect on seedling recruit-
ment and survival. Responses of individual trees and whole-forest ecosystems to CO
and climate manipulations
often vary by age, implying that forests of different ages will respond differently to climate change. Furthermore, spe-
cies within a community typically exhibit differential responses to CO
and climate, and altered community dynam-
ics can have important consequences for ecosystem function. Age- and species-dependent responses provide a
mechanism by which climate change may push some forests past critical thresholds such that they fail to recover to
their previous state following disturbance. Altered dynamics of forest recovery will result in positive and negative
feedbacks to climate change. Future research on this topic and corresponding improvements to earth system models
will be a key to understanding the future of forests and their feedbacks to the climate system.
Keywords: biogeochemistry, climate feedback, FACE, irrigation, regime shift, succession, throughfall manipulation, warming
Received 19 December 2012 and accepted 29 January 2013
The dynamic process of forest regeneration following
disturbance is of key importance, with ramifications on
several scales. On a local level, forest recovery involves
wholesale rearrangement of vegetative structure,
carbon (C) and nutrient cycling, ecosystem physiology,
and community structure (Table 1). On a landscape
level, disturbancerecovery dynamics play an impor-
tant role in the maintenance of species diversity, as
different species use forests of different ages as habitat
patches. On a regional to global level, secondary forests
are consequential for their role in climate regulation.
Forests recovering from disturbance (secondary forests)
are strong C sinks and play an important role in the
global C cycle (Running, 2008; Pan et al., 2011). For
instance, in recent years (20002007), regrowth of tropi-
cal forests following agricultural abandonment took up
an estimated 1.7 Pg C yr
(Pan et al., 2011), which is
equivalent to ca. 20% of annual global fossil fuel emis-
sions. Beyond their influence on climate through their
role in the global carbon cycle, secondary forests also
influence climate through biophysical mechanisms (Liu
Correspondence: Kristina J. Anderson-Teixeira, Smithsonian
Conservation Biology Institute, National Zoological Park, 1500
Remount Rd., Front Royal, VA 22630 USA, tel. 1 540 635 6546, fax
1 540 635 6506, e-mail:
Published 2013. This article is a U.S. Government work and is in the public domain in the USA. 1
Global Change Biology (2013), doi: 10.1111/gcb.12194
et al., 2005; Maness et al., 2012; O’Halloran et al., 2012);
for example, in northern regions albedo decreases with
forest age and strongly shapes the net climate regula-
tion services of secondary forests (Randerson et al.,
2006; Jin et al., 2012; O’Halloran et al., 2012).
A large and growing proportion of forests have been
affected by major disturbances. Globally, secondary for-
ests recovering from anthropogenic disturbances such
as agriculture and wood harvesting cover an estimated
27 million km
(Hurtt et al., 2011), and an estimated
1.2 million km
are in use as forestry plantations
(Kirilenko & Sedjo, 2007). In addition, natural distur-
bances affect a significant proportion of Earth’s ecosys-
tems; disturbances such as fires, storms, droughts, and
insect outbreaks affect over 100 000 km
annually in
North America alone (Amiro et al.,2010).Climatechange
is generally increasing the incidence of natural distur-
bances (Dale et al.,2001),includingfires(Westerlinget al.,
Table 1 Typical trajectories of change in forest properties following stand-replacing disturbance
Forest property Typical trajectory References
accumulation rate
Rapid initial increase, peak at intermediate age followed
by slow decline to near zero in old-growth forests.
Lichstein et al., 2009; Yang et al., 2011;
Hember et al., 2012
Leaf biomass or area Rapid initial increase, relatively stable thereafter. Uhl & Jordan, 1984; Bormann & Likens,
1994; Law et al., 2003; Goulden et al.,
2011; Yang et al., 2011
Fine root biomass Rapid initial increase, relatively stable or modest
decline thereafter.
Vogt et al., 1983; Claus & George, 2005;
Yuan & Chen, 2010
Carbon cycle
Gross primary
Rapid initial increase, relatively stable or modest
decline thereafter.
Amiro et al., 2010; Goulden et al., 2011
Net primary
Rapid initial increase, modest decline thereafter. Gower et al., 1996; Law et al., 2003;
Pregitzer & Euskirchen, 2004; Goulden
et al., 2011
Relatively constant. Law et al., 2003; Pregitzer & Euskirchen,
2004; Goulden et al., 2011
Net ecosystem
C balance
Initially negative (C source), increasing to maximum
(C sink) at intermediate ages, declining thereafter.
Controversy remains as to whether it declines to zero
(C neutrality).
Law et al., 2003; Pregitzer & Euskirchen,
2004; Zhou et al., 2006; Baldocchi, 2008;
Luyssaert et al., 2008; Amiro et al., 2010;
Goulden et al., 2011
Foliar [N] Relatively constant with age, although both decreases
(more common) and increases have been observed.
Davidson et al., 2007; Drake et al., 2010;
Yang et al., 2011
N mineralization Mixed responses; both increases and decreases have
been observed.
Vitousek et al., 1989; LeDuc & Rothstein,
Canopy transpiration Rapid initial increase, modest decline thereafter. Roberts et al., 2001; Delzon & Loustau,
2005; Amiro et al., 2006; Drake
et al., 2011a
Hydraulic limitation Increases with age. Drake et al., 2010, 2011a
Sensitivity to variation
in water availability
Decreases with age. McMillan et al., 2008; Drake et al., 2010;
Voelker, 2011
Community dynamics
Species turnover Rapid initial turnover, decelerating decrease in
turnover rate as the forest ages.
Anderson, 2007b
Species richness Initial increase, sometimes peaking and declining
modestly in older forests.
Shafi & Yarranton, 1973; Finegan, 1996;
Anderson, 2007b
Competition Increasing competitive advantage to late-successional
species (e.g., shade tolerant, slow growing, higher
wood density, longer lived)
Bazzaz, 1979; Bazzaz & Pickett, 1980;
Finegan, 1984
Size structure Initially, relatively even aged; competitive thinning
and seedling recruitment drive convergence
toward inverse square relationship between
abundance and diameter (diverse age structure)
Enquist et al., 2009
Published 2013. This article is a U.S. Government work and is in the public domain in the USA., Global Change Biology, doi: 10.1111/
2006) and biotic disturbances such as insect outbreaks
(Evangelista et al.,2011;Hickeet al.,2011).Theoryand
models predict that future climate change may cause
even more drastic changes (e.g., Westerling et al.,2011),
depending on the future course of greenhouse gas emis-
sions and the resultant shifts in climate (IPCC, 2007).
Because secondary forests are strong carbon sinks with
considerable value for greenhouse gas mitigation (Ander-
son-Teixeira & DeLucia, 2011) and also represent a poten-
tial bioenergy source (e.g., US DOE, 2011), secondary
forests are likely to play substantial roles in climate miti-
gation initiatives and bioenergy production (Kirilenko &
Sedjo, 2007).
Although there is strong and abundant evidence that
climate change will affect forests of all ages, we lack
synthetic understanding of how climate change will
interact with forest age to shape the dynamics of forest
recovery. Because forests undergo substantial reorgani-
zation of following major disturbance (Table 1), it is
likely that climate change will have different effects on
forests of different ages, thereby altering the trajectory
of succession relative to those observed for historical
climates. Climate change may alter one or more distinct
features of successional trajectories (Fig. 1). First, the
rate at which the forest moves along the successional
trajectory may be altered without necessarily implying
any changes to the successional pathway or the state of
mature forests; for example, increased productivity
may accelerate biomass accumulation without altering
the biomass of mature forests. Second, the state of
mature forests may be altered; for example, maximum
biomass may be altered if future climates place differ-
ent biophysical constraints on the number and size of
trees that can persist. Third, the successional pathway
may be altered; for example, tree establishment may be
delayed by altered climatic conditions such that pro-
portionally more time is spent in an early-successional
shrub phase. Distinguishing how climate change affects
forests of different ages and thereby how it alters suc-
cessional trajectories is critical in understanding how
climate change will impact both recently disturbed and
mature forests.
This review considers how altered atmospheric CO
and climate are likely to impact trajectories of forest
recovery, with a particular focus on how climate
change may alter the rate of succession, the state of
mature forests, and successional pathways (Fig. 1). We
consider five major lines of evidence relating forest
structure and function to directional variation in CO
and climate (i.e., average conditions, as opposed to
intra- or interannual variation), each of which yields
insight into how forest recovery may be altered under a
changing climate. First, we review how climatic influ-
ences the dynamics of forest recovery across broad
climatic gradients. Second, we summarize the results of
experimental studies quantifying the effects of elevated
, elevated temperature, and altered precipitation
regimes on the dynamics of forest recovery. Third, we
review observations of altered forest recovery under
contemporary multivariate environmental change.
Fourth, we consider how altered community dynamics
may impact forest recovery. Fifth, we review model
predictions. Finally, we synthesize findings from these
separate lines of evidence, identify remaining uncer-
tainties, and discuss the implications for ecological
communities, biogeochemical processes, and the
climate system.
Dynamics of forest recovery across broad climatic
Although few studies have evaluated how climate
influences forest recovery across broad climatic gradi-
ents (Prach & Rehounkov!
a, 2006; Anderson, 2007a),
there is clear evidence that climate exerts a strong influ-
ence on the rate of succession, the state of mature for-
ests, and their successional pathways (Fig. 1). The rate
of forest regrowth following disturbance is strongly
influenced by climate (Brown & Lugo, 1982; Johnson
et al., 2000; Anderson et al., 2006). Globally, the rate of
living biomass accumulation increases with tempera-
ture, being on average three to four times faster in the
tropics than in high-latitude forests (Fig. 2a; Anderson
et al., 2006). Likewise, biomass accumulation rate
increases with precipitation at a global scale (Fig 2b).
Within the tropics, rates of biomass accumulation are
Fig. 1 Schematic diagram illustrating three ways in which cli-
mate change may impact the dynamics of forest recovery. Rela-
tive to the historical trajectory of change in a forest property
(here, biomass) with age, climate change may alter (1) the rate
of change, (2) the state to which the property converges as the
forest matures (‘mature state’), or (3) the successional pathway
(i.e., the sequence of states through which any given ecosystem
property passes and the relative amount of time spent in each).
Published 2013. This article is a U.S. Government work and is in the public domain in the USA., Global Change Biology, doi: 10.1111/
dramatically higher in moist climates (precipitation
10002500 mm yr
) than in dry climates (precipitation
<1000 mm yr
); however, the positive influence of
precipitation appears to saturate, with rates in wet cli-
mates (precipitation >2500 mm yr
) less than or equal
to those in moist climates (Brown & Lugo, 1982; Mar!
Spiotta et al., 2008). Although further research is
required to fully understand the mechanisms through
which temperature, water availability, and their sea-
sonal dynamics affect rates of biomass accumulation in
secondary forests, we can say conclusively that warmer
temperatures and higher moisture availability are asso-
ciated with higher rates of biomass accumulation.
Growth in secondary forests is strongly linked to ele-
mental cycling. Biogeochemical cycles of elements
including C, nitrogen (N), and phosphorous (P) are
coupled to biomass accumulation through stoichiome-
tric constraints on the elemental composition of vegeta-
tion, such that rates at which these elements are
sequestered in vegetation are grossly proportional to
rates of biomass accumulation (Yang et al., 2011).
Indeed, mirroring the climate dependence of rates of
biomass accumulation (Fig. 2ab; Anderson et al.,
2006), it has been observed that the rate of N uptake by
a regrowing tropical forest in Costa Rica is four times
that of a regrowing temperate forest at Hubbard Brook,
USA (Russell & Raich, 2012). However, the trajectory of
forest recovery is also shaped by biogeochemistry
climate interactions. For example, climate influences
temporal patterns of N availability during secondary
succession (Vitousek et al., 1989), rates of change in soil
C and N (Li et al., 2012), and plant tissue stoichiometry
(Wright et al., 2004). Thus, climate may indirectly influ-
ence forest recovery through its influence on biogeo-
chemistry, as occurs in the case of forests developing
on Hawaiian lava flows (Anderson-Teixeira et al., 2008;
Anderson-Teixeira & Vitousek, 2012).
A few studies have compared rates or pathways of
secondary succession across broad climatic gradients.
Following clear cutting in western Oregon, climate
shapes both the rate and pathway of forest succession;
in the western Cascades region, conifer regeneration is
slower and follows a longer establishment delay com-
pared with the more mesic Coastal Range region (Yang
et al., 2005). In subalpine forests of the Colorado Rock-
ies, the rate of succession is more than twice as rapid in
mesic sites as compared with xeric sites (Donnegan &
Rebertus, 1999). Likewise, in the Medicine Bow moun-
tains of Wyoming, succession to a mature sprucefir
forest is most rapid in a mesic drainage bottom, slower
on a less mesic north-facing slope, and rarely occurs at
more arid sites prior to stand-clearing fire (Romme &
Knight, 1981). In the Czech Republic, the rate of succes-
sion in vegetative communities (including forests and
nonforests) is strongly influenced by climate; mean
annual change in dominant species cover during the
first 12 years of succession decreases dramatically with
increasing elevation (increasing precipitation and
(a) (b)
(c) (d)
Fig. 2 Influence of climate on forest recovery rates (a, b) and on aboveground C stocks of mature forests (c, d). The rate of aboveground
biomass accumulation in forests recovering from stand-clearing disturbance varies globally with respect to (a) mean annual tempera-
ture (MAT) and (b) precipitation (MAP). Data, which are from Anderson et al. (2006), represent natural regeneration in 68 unmanaged
forests worldwide. Solid and dashed lines represent an exponential fit and its 95% confidence interval, respectively. Similarly, above-
ground C stocks (biomass +coarse woody debris; CWD) in mature forests vary globally with respect to (c) MAT and (d) MAP. Data
from Anderson-Teixeira et al. (2011). Dashed lines represent hypothesized bioclimatic limits.
Published 2013. This article is a U.S. Government work and is in the public domain in the USA., Global Change Biology, doi: 10.1111/
decreasing temperature; Prach et al., 2007). These exam-
ples provide evidence that climate strongly influences
both the rate and pathway of succession.
In addition to its influence on the rate and trajectory
of forest recovery, climate also shapes the types of
steady-state conditions toward which secondary forests
can eventually converge. Globally, aboveground
biomass of forests is influenced by temperature and
precipitation (Fig. 2cd; Keith et al., 2009; Anderson-
Teixeira et al., 2011; Larjavaara & Muller-Landau,
2012), and climate strongly influences most other major
components of ecosystem C cycles, including gross and
net primary productivity (GPP and NPP, respectively;
Luyssaert et al., 2007), soil and whole-ecosystem respi-
ration, and soil organic carbon (Raich & Schlesinger,
1992; Jobb!
agy & Jackson, 2000). Similarly, species diver-
sity varies globally with respect to climate (Brown et al.,
1998). In sum, climate can determine the state to which
forests converge following disturbance both directly
through biophysical constraints and indirectly through
its influence on biogeochemistry and the surrounding
The broad-scale patterns described above demon-
strate that climate strongly influences the rate of forest
recovery, successional pathways, and the structure and
function of mature forests. However, transient dynam-
ics under a rapidly changing climate may diverge from
expectations based on these contemporary patterns,
and increasing atmospheric CO
concentrations will
alter physiological constraints on forests. To under-
stand the more immediate responses of forest recovery
to elevated CO
and climate change, we turn to experi-
mental manipulations.
Forest responses to experimental manipulation of
, temperature, and precipitation
Experiments manipulating CO
, temperature, and pre-
cipitation demonstrate that altered climatic conditions
will alter ecosystem and community dynamics in sec-
ondary forests. The responses of terrestrial ecosystems
in general to these experimental manipulations have
been previously reviewed (e.g., Pendall et al., 2004; De
Graaff et al., 2006; Norby & Zak, 2011; Wu et al., 2011;
Beier et al., 2012; Dieleman et al., 2012; Lu et al., 2012);
here, we focus specifically on the responses of forests
and any age dependency of their responses (Fig. 3).
Responses to CO
Tree-dominated ecosystems all in temperate or boreal
regions have been exposed to elevated CO
Free-Air Carbon dioxide Enrichment (FACE), Open
Top Chamber (OTC), and Whole Tree Chamber (WTC)
experiments. Given the logistical difficulties of elevat-
ing CO
in forests with tall canopies, the majority of
these experiments have been performed on young
forests or trees, with only one FACE experiment in a
mature forest to date (Table S1).
Elevated CO
consistently enhances photosynthesis,
or GPP at the ecosystem level (Ceulemans & Mousseau,
1994; Curtis & Wang, 1998; Ainsworth & Long, 2005;
onen et al., 2007). In young forests, this results in
increased NPP and biomass; at least at the onset of the
experiment (DeLucia et al., 1999; Norby et al., 2005).
However, whereas substantial NPP and biomass
increases have occurred at the onset of experiments,
this NPP stimulation has persisted in some but not all
forests (Oren et al., 2001; Seiler et al., 2009; McCarthy
et al., 2010; Norby et al., 2010). Moreover, this response
becomes less consistent as forests become older, and
NPP did not increase in the only mature forest exposed
to elevated CO
(Fig. 3; K
orner et al., 2005; Bader et al.,
2009). Similarly, leaf and fine root biomass are consis-
tently stimulated in young forests, but may decline in
old forests (Fig. 3; K
orner et al., 2005; Bader et al., 2009).
Thus, although there is strong evidence that CO
ization increases the rate of biomass accrual in young
forests, a question remains as to whether elevated CO
increases the biomass and productivity of mature for-
ests (Fig. 1; K
orner et al., 2005; Hyv
onen et al., 2007;
Norby & Zak, 2011). In large part because of this uncer-
tainty, it remains unclear whether the net carbon
balance of mature forests will increase in response to
fertilization (Fig. 3).
The ability of forests to sustain increased NPP under
elevated CO
as they age and, ultimately, the potential
for mature forests to increase C storage under elevated
depends in large part upon biogeochemistry. One
potential explanation of observed declines in NPP stimu-
lation under elevated CO
as forests age is that increased
productivity immobilizes nutrients in woody tissue or soil
organic matter such that soil N and other nutrients
needed to sustain growth become depleted and may
eventually limit growth (Luo et al.,2004).ProgressiveN
limitation can be alleviated through a variety of mecha-
nisms: trees can increase their N use efficiency, invest
more C in belowground nutrient acquisition, or access
deep N pools (McKinley et al.,2009;Iversen,2010;Drake
et al.,2011b;Norby&Zak,2011).Nlimitationcanalsobe
mitigated if greater N mineralization occurs under high
fixation is stimulated by elevated CO
(Zanetti et al.,1996;Hungateet al.,2004;Luoet al.,2004;
Hoosbeek et al.,2011;Norby&Zak,2011).Earlywork
suggested that N mineralization declines under elevated
(Hungate et al.,1999),butmorerecentstudiessug-
gest that there can be a priming effect through time from
increased atmospheric CO
that stimulates soil micro-
Published 2013. This article is a U.S. Government work and is in the public domain in the USA., Global Change Biology, doi: 10.1111/
bial activity, which in turn degrades slowly cycling
organic matter pools and release mineral N (Carney
et al.,2007;Langleyet al.,2009;Drakeet al.,2011b).
Elevated CO
may also increase N availability by
increasing labile C to drive the energetics of N
fixation (Hungate et al.,1999).However,ina
scrub-oak system in Florida, N
fixation was negatively
impacted by 7 years exposure to elevated CO
gate et al., 2004), and N
-fixation rates have continued
to decline perhaps due to canopy closure and light
limitation (Duval, 2010). Thus, N deficiency may be
avoided and NPP stimulation sustained over time
through a variety of mechanisms; however, it remains
uncertain whether this can continue indefinitely or
whether NPP stimulation in all forests would eventu-
ally decline given sufficient time (Hyv
onen et al., 2007;
Norby & Zak, 2011).
The responses of tree growth to elevated CO
variable among species (Bazzaz, 1990; Saxe et al.,
1998; Pe~
nuelas et al., 2001; K
orner et al., 2005; Seiler
et al., 2009; Dawes et al., 2011), and differential spe-
cies responses have commonly been observed in
-enrichment experiments (Table S1). For exam-
ple, of the three codominant canopy tree species
(Fagus sylvatica,Quercus petraea,Carpinus betulus) in
the mature deciduous forest exposed to elevated
Fig 3 Schematic diagram illustrating typical forest successional trajectories under ambient climate (solid lines; as reviewed in Table 1)
and how these are affected by experimental CO
fertilization, warming, and drought (increases in blue, decreases in red; color satura-
tion scales with certainty). Responses to CO
and climate change are based on a comprehensive review of experimental studies (Tables
S1S3). Responses are considered to have high certainty when observed in multiple sites and low certainty when observed in only one
study. *Indicates a response that is time dependent; it may change from negative to positive as increased N mineralization stimulates
biomass growth (Melillo et al., 2011).
Published 2013. This article is a U.S. Government work and is in the public domain in the USA., Global Change Biology, doi: 10.1111/
, shade-tolerant Fagus exhibited increased annual
basal area increments in response to CO
in two of
four treatment years, whereas growth of the other
species remained the same or declined (K
orner et al.,
2005). Similarly, proportional species’ contributions
to whole-ecosystem productivity shifted in a Florida
scruboak ecosystems exposed to elevated CO
dominant Quercus myrtifolia exhibited strong biomass
growth, Q. chapmanii exhibited less of an effect, and
subdominant Q. geminata showed no growth stimu-
lation (Dijkstra et al., 2002). Thus, differential species
growth responses consistently alter proportional spe-
cies’ contributions to whole-ecosystem productivity
and will likely change the composition of future
Understory vegetation can influence ecosystem func-
tioning and future community composition (Nilsson &
Wardle, 2005) and, therefore, impacts of global change
on juvenile trees and influential nontree species serve
as a window into the forests of the future. Moreover, as
the majority of forest biodiversity is in the understory
stratum, impacts on understory species as well as sym-
biotic mycorrhizal fungi bear consequences for tree
recruitment, carbon cycling, forest health and biodiver-
sity (Gilliam, 2007). Understory community responses
to CO
enrichment have been commonly observed
(Table S1). At ORNL-FACE, the woody understory
increased in importance relative to the total stand and
to herbaceous plants, indicating a potential acceleration
of succession under elevated CO
(Souza et al., 2010).
Consistent with earlier work using pots and growth
chambers (Bazzaz & Miao, 1993; Kubiske & Pregitzer,
1996; Kerstiens, 1998, 2001; H
attenschwiler & K
2000), CO
enrichment at DukeFACE tended to favor
slow-growing, shade-tolerant species with low rates of
productivity in understory conditions, again suggesting
that succession may be accelerated in temperate forests
under future conditions, with implications for bio-
sphereatmosphere carbon feedbacks (Mohan et al.,
2007). In addition, CO
enrichment may favor woody
vines (lianas; e.g., Sasek & Strain, 1990). This has been
observed in two FACE studies (Table S1); for example,
at DukeFACE, the woody vine poison ivy (Toxicoden-
dron radicans) growth was disproportionately enhanced
under elevated CO
(Mohan et al., 2006). Lianas have
been expanding in abundance in many regions of the
world often to the detriment of recruiting and mature
trees (Dillenburg et al., 1995; Ingwell et al., 2010; Schnit-
zer & Carson, 2010; Schnitzer & Bongers, 2011) and
the positive feedback of elevated CO
for vines may
hinder the establishment of secondary forests globally.
Thus, increasing atmospheric CO
may substantially
alter the rate and pathway of succession as well as the
composition of mature forest communities (Fig. 1).
Responses to warming
Over the last three decades, several tree-dominated eco-
systems of various ages almost all in temperate and
boreal regions have been exposed to experimental
warming (Table S2). These experiments have warmed
either aboveground vegetation or the soil (through use
of buried cables); there are few soil-and-air warming
experiments done at the scale of canopy trees (Slaney
et al., 2007; Bronson & Gower, 2010).
Soil warming in northern forests results in faster
decomposition and microbial processing of soil C and
N, which directly releases more CO
to the atmosphere
because of enhanced soil respiration (Table S2; Rustad
et al., 2001; Melillo et al., 2002, 2011). By increasing N
mineralization rates, soil warming can have an indirect
N fertilization effect, which generally increases above-
ground production and lowers C allocation to fine root
biomass (Fig. 3; Zhou et al., 2011). The net ecosystem C
balance in response to warming depends largely on the
counteracting effects of C release through increased soil
respiration and C sequestration through increased bio-
mass growth (Fig. 3); in a 60- to 70-year-old even-aged
oakmaple forest in central Massachusetts subjected to
7 years of soil warming (Harvard Forest), soil C losses
were increasingly offset by stimulated growth of can-
opy trees (after a lag of several years; Melillo et al.,
2011; Butler et al., 2012). Additional changes may be
driven by aboveground warming; over the first few
years of warming in a 12-year-old black spruce (Picea
mariana) plantation in Manitoba, soil respiration
increased under soil warming but decreased under
soil-and-air warming (Bronson et al., 2008). In this
study, elevated soil and air temperatures increased
spruce tree shoot growth (Bronson et al., 2009) but did
not change rates of photosynthesis or autotrophic respi-
ration (Bronson & Gower, 2010). Much remains to be
learned about how warming affects whole forested eco-
systems, particularly in subtropical and tropical forests,
where only one warming experiment has been con-
ducted to date (Cheesman & Winter, 2012). Moreover,
although we may posit that forest age modulates
warming responses based on the magnitude of struc-
tural and functional changes associated with forest
recovery (Table 1), there is of yet no clear evidence that
the direction of forest responses to warming varies by
age (Fig. 3).
Growth responses to warming vary among tree spe-
cies (Table S2), and this is likely to affect successional
dynamics and forest composition. For example,
although the large oaks at Harvard forest accounted for
the majority of C uptake and storage in woody tissue,
smaller maples exhibited a greater stimulation of
growth in response to soil warming stimulation (Mel-
Published 2013. This article is a U.S. Government work and is in the public domain in the USA., Global Change Biology, doi: 10.1111/
illo et al., 2011; Butler et al., 2012; Mohan et al., unpub-
lished results). Similarly, a warming experiment in a
recently timbered oakhickory forest in Pennsylvania
found altered phenology (with differential responses
among species) and community composition (Rollin-
son, 2010; Rollinson & Kaye, 2011). Thus, warming is
likely to alter species’ growth and phenology and,
thereby, the rate and pathway of succession and ulti-
mately the community composition of mature forests
(Fig. 1).
Responses to altered precipitation
There have been a number of precipitation manipula-
tion experiments in forests of a range of ages spanning
from boreal to tropical regions (Table S3; Beier et al.,
2012). Across this range of climates and forest ages,
tree growth and survival were generally increased by
water addition and reduced by water removal (Fig. 3;
e.g., Hanson et al., 2001; Nepstad et al., 2002; Plaut
et al., 2012; Vasconcelos et al. (2012), as was GPP or
NPP at the ecosystem level (Nepstad et al., 2002; Al-
berti et al., 2007). Soil respiration rates also tended to
increase under irrigation and decrease under drought
(Table S3; e.g., Conant et al., 2000; Sotta et al., 2007).
However, water addition only accelerated forest C
cycling up to a point; some more mesic forests did not
respond to precipitation manipulation (De Visser et al.,
1994; Bergh et al., 1999) or had accelerated C cycling
under reduced precipitation (Cleveland et al., 2010).
Sensitivity to precipitation manipulation often varied
by size class, but results were mixed as to whether
small or large trees were more sensitive (Hanson et al.,
2001; Nepstad et al., 2007). Whereas altered precipita-
tion had a strong effect on seedling emergence and
survival (Richter et al., 2012; Volder et al., 2012) and at
times had a stronger effect on small than on large trees
(Hanson et al., 2001), there were also instances where
exposed canopy trees suffered greater drought-related
stress (Nepstad et al., 2007; Schuldt et al., 2011). Thus,
within-stand relationships between tree age and
drought sensitivity do not necessarily mirror across-
stand relationships, where growth sensitivity to varia-
tion in water availability declines with stand age
(Table 1). As with experimental manipulation of CO
and temperature, differential species responses were
commonly observed under precipitation manipulation
experiments (Table S3; e.g., Yavitt & Wright, 2008),
portending future changes to community composition
under altered precipitation regimes. Thus, in sum-
mary, precipitation manipulation experiments have
demonstrated that water availability affects rates of
forest recovery, mature forest states, and probably suc-
cessional pathways (Figs 1 and 3).
Responses to multivariate environmental manipulation
Joint effects of altered CO
, temperature, and precipita-
tion are rarely purely additive (Dieleman et al., 2012),
and understanding the interactive effects produced by
combined manipulations remains an important chal-
lenge. Experimental manipulation of more than one of
these elements in a factorial design has occurred in sev-
eral intact forests or experimental mesocosms (Tables
S1S3). These studies demonstrate that tree growth and
carbon cycling in young forests are generally acceler-
ated under combined higher CO
, warmer, and wetter
conditions (Tables S1S3; e.g., Wan et al., 2004;
Comstedt et al., 2006; Slaney et al., 2007; Tingey et al.,
2007; Bauweraerts et al., 2013). In addition, different
species have responded differently to different ele-
ments of climate change; for example, at the alpine tree
line in Switzerland, growth of Larix decidual responded
positively to CO
but was unresponsive to soil warm-
ing, whereas Pinus cembra had a slight positive response
to warming but responded minimally to CO
et al., 2011). The limited number of studies and the
complexity of multifactor experiments make it prema-
ture to generalize about how forests of different ages
will respond to interactive elements of global change.
However, observed responses of forests to the environ-
mental change that has already occurred to which we
turn next reveal how secondary forests are respond-
ing to multivariate climate change to date.
Altered forest recovery under contemporary
multivariate environmental change
Historical reconstructions indicate that forests of all
ages have responded to the changes in CO
and climate
that have already occurred. By nature, these historical
records do not directly separate the effects of CO
, tem-
perature, and precipitation from one another and from
other potentially confounding environmental changes
(e.g., atmospheric deposition, ozone, management,
altered disturbance regimes). Rather, they provide a
picture of how the dynamics of forest recovery are
responding to contemporary multivariate environmen-
tal change.
Tree-ring and observational records extending back
decades to centuries have demonstrated the climate
dependence of forest productivity. Tree-ring records
have revealed increasing growth rates in numerous
forests including high-elevation forests in western
Washington (Graumlich et al., 1989), conifers in the
white mountains of California (Lamarche et al., 1984;
Salzer et al., 2009), Pinus ponderosa forests in the US
Pacific northwest (Soul!
e & Knapp, 2006), aspen (Popu-
lus tremuloides) secondary forests in Wisconsin (Cole
Published 2013. This article is a U.S. Government work and is in the public domain in the USA., Global Change Biology, doi: 10.1111/
et al., 2010), Abies and Quercus forests in France (Bec-
ker, 1989; Becker et al., 1994), and numerous other for-
ests throughout Europe (Spiecker, 1999; Babst et al.,
2013). These increased growth rates are generally
attributable to increased atmospheric CO
, tempera-
ture, or moisture (e.g., Graumlich et al., 1989; Soul!
Knapp, 2006; Salzer et al., 2009; Cole et al., 2010). In
contrast, tree growth rates have decreased in response
to warming or drought stress in many other forests
around the world (Allen et al., 2010), including white
spruce (Picea glauca) in interior Alaska (Barber et al.,
2000), conifers in the southwest United States
(Williams et al., 2013), and tropical forests in Panama,
Malaysia, and Costa Rica (Feeley et al., 2007b; Clark
et al., 2010). Similarly, a review documenting evidence
of altered forest productivity over the last half century
indicates that the productivity of many forests is
increasing whereas that of others is declining (Boisve-
nue & Running, 2006). Although powerful for under-
standing the historical influence of climate on forest
productivity, these records are limited in that they do
not characterize responses across a range of forest
By comparing biomassage relationships (deter-
mined through a chronosequence approach) with cur-
rent biomass accumulation rates in forests of various
ages (determined through repeated sampling), a couple
of studies have identified accelerated biomass accumu-
lation in forests following stand-clearing disturbances.
Specifically, accelerated biomass accumulation has been
observed in temperate deciduous forests in the eastern
United States (McMahon et al., 2010a; see also Foster
et al., 2010; McMahon et al., 2010b) and in temperate-
maritime forests in the Pacific northwest (Fig. 4; Hem-
ber et al., 2012). Likely explanations of these increases
in secondary forest biomass accumulation rates include
increased atmospheric CO
, increased temperature,
increased moisture, and increased growing season
length (McMahon et al., 2010a; Hember et al., 2012).
Thus, climate change appears to be increasing the rate
of forest regrowth in some temperate forests; however,
parallel studies have yet to be conducted in other
The long time frame of forest recovery precludes
comparison of forests that have matured under differ-
ent climates, which would be necessary to determine
whether climate change is altering recovery trajectories
such that forests converge to an altered state as they
mature (Fig. 1). However, long-term monitoring of
mature forests can provide evidence as to whether
climate change is affecting the state of forests that
matured under past climates. Long-term monitoring of
old-growth forests provides mixed evidence as to
whether their total carbon storage capacity is changing;
many old-growth forests throughout the world appear
to be net C sinks (Baker et al., 2004; Luyssaert et al.,
2007; Chave et al., 2008; Lewis et al., 2009); however,
this effect is diminished at larger spatiotemporal scales
of measurement (Clark, 2002; Feeley et al., 2007a; Chave
et al., 2008). There is strong evidence of directional
change in community composition of forests through-
out the world; for example, long-term records from the
50 ha forest dynamics plot on Barro Colorado Island,
Panama, indicate increased dominance of drought-
tolerant species (Feeley et al., 2011). In addition, there
have been general increases in forest die-back globally
a phenomenon attributed to climate changetype
drought (Breshears et al., 2005; Allen et al., 2010;
Williams et al., 2013).
Thus, there is evidence of historical change in both
rates of forest regrowth and the state of forests that
matured under past climates. There is also some evi-
dence of changing successional trajectories driven by
altered community dynamics (reviewed below).
Although concurrent changes in multiple environmen-
tal factors including atmospheric CO
, climate, atmo-
spheric deposition, herbivore communities, disturbance
regimes, and management make it difficult to isolate
the cause of these changes, their global distribution and
directional correlation with trends in CO
and climate
provide strong evidence that they are at least partially
attributable to increasing atmospheric CO
and climate
Fig. 4 Evidence of increasing rates of biomass accumulation in
coastal Douglas fir (Pseudotsuga menziesii) and western hemlock
(Tsuga heterophylla) forests in southwest British Columbia, Can-
ada. Plotted is the residual average biomass increment (B) from
1267 permanent inventory plots after correction for factors
including stand age, site quality, nitrogen availability, and
biomass (the five lines represent different correction methods,
as detailed in Hember et al., 2012). Linear regression represents
a significant positive trend. Reprinted from Hember et al. (2012).
Published 2013. This article is a U.S. Government work and is in the public domain in the USA., Global Change Biology, doi: 10.1111/
Community dynamics and the potential for state
Successional pathways may be altered when elevated
and climate change affect community dynamics,
either directly through differential effects on the perfor-
mance of various species and size classes or indirectly
through altered disturbance regimes and consequent
competitive outcomes.
Climate change will alter community dynamics by
altering the physical environment in which species of
varying physiological strategies are competing. Experi-
mental climate change manipulations (reviewed above;
Tables S1S3) and decades of forestry research on the
climate sensitivity of forest regeneration (Fowells &
Stark, 1965; Ferrell & Woodard, 1966; Thomas & Wein,
1985) have demonstrated that increased CO
altered climate will differentially affect growth rates of
trees by size and species, thereby altering population
dynamics, competitive interactions, and species compo-
sition of both young and mature forests. In addition,
climate change can differentially favor or inhibit com-
mon forest pathogens, providing another mechanism of
impact on community structure (reviewed by Sturrock
et al., 2011). Such community changes can affect ecosys-
tem function, altering production, C stocks, and biogeo-
chemistry. For example, model predictions of climate
change effects on forest productivity can be very differ-
ent if the community is allowed to develop dynami-
cally, compared with using parameters based on
average forest characteristics, which is a common prac-
tice in biogeochemical models examining the effects of
climate change (Bolker et al., 1995).
Beyond its direct effects on the dynamics of forest
recovery through physiological mechanisms, climate
change may also impact successional pathways indi-
rectly by altering the frequency, timing, severity, and
spatial extent of disturbances including fires, droughts,
storms, floods, and herbivore or pathogen outbreaks
(e.g., Dale et al., 2001; Westerling et al., 2006; Allen
et al., 2010; Sturrock et al., 2011). Frequency and inten-
sity of disturbance have been theoretically shown to
have very different effects on community diversity
(Miller et al., 2011; in determining microcosm diversity
(Hall et al.,2012).Thus,changesindisturbancefrequency
and intensity have the potential to shift community
composition, even when species are restricted to (and
are still viable in) their historic ranges. Moreover, large
changes to disturbance regimes are not required to
facilitate changes in community composition; in the
annual plant model of Miller et al. (2011), changing
disturbance mortality by just a few percentage points
can send a species to extinction. Changes in community
composition driven by altered disturbance regimes
may have dramatic consequences for ecosystem func-
tioning. For example, in Alaskan boreal forests, increas-
ing fire frequency and severity have shifted
competitive dominance from conifers to deciduous spe-
cies, affecting biomass and soil C accumulation, albedo,
and energy partitioning (Beck et al., 2011).
Disturbance can also provide niche opportunities for
invaders (Shea & Chesson, 2002), and the successful
invasion can dramatically alter successional trajectories
and also feedback to further modify disturbance
regimes (Mack & D’Antonio, 1998). Although distur-
bances are commonly believed to increase invader suc-
cess, recent work suggests that it is changes to
disturbance regimes, rather than disturbance events per
se that most strongly influence a communities’ suscepti-
bility to invasions (Moles et al., 2012). In this light,
climate change is likely to change the composition of
some communities by altering disturbance regimes to a
point where invader species can become dominant.
In some cases, climate change may push forests past
critical thresholds such that, upon perturbation, they
undergo drastic changes in community composition
and ecosystem properties (‘catastrophic shift’) and fail
to return to their previous state (Fig. 5). In many sys-
tems, the observed state of the community is not the
only possible stable state; a variety of empirical results
demonstrate the existence of alternative stable states in
nature (D’Antonio & Vitousek, 1992; Savage & Mast,
2005; Schr
oder et al., 2005; Odion et al., 2010; Scheffer
et al., 2012). Large changes in the environment can
bring about large changes in ecosystems, but smooth,
gradual changes in abiotic conditions also can cause
abrupt shifts in ecosystem properties and functioning
(Scheffer et al., 2001). Systems that are structured by
disturbance and are susceptible to abiotic forcing (such
as regenerating forests) may be more likely to display
alternative stable states (Didham et al., 2005). When dis-
turbance keeps systems in perpetual flux, as is the case
for many forests, no true stable equilibrium (in the clas-
sical, dynamical systems sense) is reached. Instead, for-
ests undergo periodic cycles of disturbance and
regeneration, and it is these cycles that constitute the
‘state’ of the system.
Many forests are resilient (sensu Grimm & Wissel,
1997) to commonly experienced disturbances, but
effects of climate change, such as changes to the distur-
bance and precipitation regimes, can change the
composition and productivity of forest communities
(Thompson et al., 2009), forcing the system into
different cyclical behaviors. Although different initial
trajectories can lead to different mature forest states,
there is also the possibility that different initial trajecto-
ries can lead to the same mature state, or that similar
initial trajectories can lead to distinct mature states
Published 2013. This article is a U.S. Government work and is in the public domain in the USA., Global Change Biology, doi: 10.1111/
(Fig. 1, ‘alternative transient states’ sensu Fukami &
Nakajima, 2011).
In some forested regions, the existence of alternative
stable states implies that forests may not return to their
previous state following disturbance. A general mecha-
nism underlying such alternative stable states is that
seedlings and young forests are often more vulnerable
to disturbances such as drought, herbivory, and fire
than their mature counterparts (Table 1; Stromayer &
Warren, 1997; Thompson & Spies, 2010). As a result,
conditions that support the persistence of mature
forests may not be amenable to forest regeneration. For
example, following fire, conifer regeneration may be
delayed or prevented by drought or competitive inhibi-
tion by grasses or shrubs (Savage & Mast, 2005;
Roccaforte et al., 2012). There are also systems in which
postfire establishment of pyrogenic vegetation or
vulnerability of young stands to crown fire reduces the
probability of forest regeneration (D’Antonio & Vitousek,
1992; Savage & Mast, 2005; Thompson & Spies, 2010;
Staver et al., 2011). For example, in the Klamath region
of Oregon and California, high-intensity fire shifts the
community from a high-biomass mixed conifer forest
to a pyrogenic low-biomass shrubchaparralhard-
wood community, in which state it may be maintained
by subsequent fires of any intensity (Odion et al., 2010;
Thompson & Spies, 2010). Alternative stable states may
also be driven by hydrologic, microclimatic, or biogeo-
chemical mechanisms; for example, postfire forest
resilience may be impacted by changes to soil biogeo-
chemistry and hydrological functioning (Ffolliott et al.,
2011; Smithwick, 2011).
Climate change may gradually alter the landscape of
alternative states, having minimal impact on mature
forests, but shifting conditions such that forests will be
unlikely to reestablish following disturbance (Fig. 5).
The probability of forest regeneration may be reduced
by mechanisms such as reduced probabilities of seed-
ling establishment under more arid conditions, reduced
competitive advantage of seedlings relative to grasses
or shrubs, or increases in disturbance frequency or
severity. Although ecological theory points toward the
risk that some forests may unlikely to return to their
previous state following stand-clearing disturbance as a
result of global change (Fig. 5), empirical evidence
remains scant. In the southwestern United States, pon-
derosa pine forests meet the criteria for forests that may
be vulnerable to climate change-induced catastrophic
shifts and are often failing to reestablish following fire
(Dore et al., 2008; Roccaforte et al., 2012); however, a cli-
mate change mechanism has not been demonstrated.
Understanding the potential for climate change to dra-
matically alter or prevent postdisturbance recovery
remains an important challenge.
Thus, based on empirical findings and theoretical
concepts, community composition and ecosystem func-
tion of regenerating forests under climate change are
likely to change, both quantitatively, and in terms of
stability. Given that climate change, disturbance
regimes, and community dynamics interact in complex
ways to shape ecosystems, correctly predicting the
behavior of forests over the next century will require
greater understanding of the potential for altered com-
munity dynamics to dramatically impact carbon
cycling, biogeochemistry, and ecosystematmosphere
Ecosystem and earth system model projections
Ecosystem and earth system models (ESMs) provide a
means to project dynamically how ecosystems will be
impacted by multiple interacting environmental
changes over spatiotemporal scales that exceed the
limits of observation and experimentation. ESMs vary
in complexity from fully coupled global circulation
Fig. 5 Schematic diagram illustrating the potential for
disturbance to force ecosystems from one stable state to another
as the climate changes. Colored shapes represent the landscape
of stable ecosystem states under different climate regimes, and
balls represent states in which ecosystems can stably exist in
this case, the state to which the system converges at maturity
(which will be associated with a stable disturbancerecovery
regime). The plot below illustrates hysteresis, wherein alternate
stable states exist. As the climate changes, basins of attraction
shift such that the stable state at maturity eventually switches
from one state to the other. During the transition, however, dis-
turbance (indicated by black arrow) may hasten the shift from
one stable state to another. Modified from Scheffer et al. (2001).
Published 2013. This article is a U.S. Government work and is in the public domain in the USA., Global Change Biology, doi: 10.1111/
models (GCMs), which include two-way feedbacks
between the land, atmosphere, and oceans to make pre-
dictions about climate, to simpler models with less
interaction between the earth system components (e.g.,
one-way feedbacks to the atmosphere such as land-
cover changes to net terrestrial CO
uptake). ESMs
include land components embedded with physiological
and biogeochemical mechanistic representations of the
interactions between vegetation, the atmosphere, and
either prognostic disturbances (i.e., fire) or prescribed
disturbances (i.e., harvest). Vegetation is represented in
terms of broadly defined plant functional types (PFTs;
e.g., temperate conifers). When coupled with a specific
class of ecosystem models (dynamic global vegetation
models; DGVMs), processes are included that allow
vegetation type to change based on climate conditions
(e.g., forest to grassland or woodland). Recent advance-
ments to some ESMs (CESM/CLM4.0, ORCHIDEE,
TEM) now include dynamic response variables for the
long-term physiological changes related to CO
temperature (Krinner et al., 2005; Thornton et al., 2007;
Zaehle & Friend, 2010). The complexity of these models,
and the variety of factors upon which model predic-
tions and associated uncertainty depend, preclude the
possibility of any one model incorporating all of the
known complexity of forest regeneration. However, for
models to make predictions about forest recovery
following disturbance, they need to be able to capture
the interactive effects of changing environmental condi-
tions and disturbance on forest recovery dynamics.
No model pays detailed attention to the roles of for-
est age and successional changes in species composition
in shaping the dynamics of forest recovery. Rather,
regenerating forests are generally parameterized as
mature forests, although sometimes there are two age
classes (e.g., fire BGC; Smithwick et al., 2009), and car-
bon allocation to wood may vary dynamically with age
(e.g., CLM4; Hudiburg et al., 2013). We are aware of
only one model where C allocation to nonwoody
components or physiology changes dynamically with
age (and this improves performance in describing age
trajectories of woody productivity; Davi et al., 2009).
Changes in community composition (i.e., PFTs), physi-
ological differences between early- and late- succes-
sional species, and age structure within a forest
(Table 1) generally are not incorporated (exception is
ED2; Medvigy et al., 2009). As a result, models have
difficulty accurately reproducing trajectories of change
in biomass or other components of the C cycle associ-
ated with forest age (Table 1). Nevertheless, to the
extent that forest responses are consistent across age
classes (Fig. 3), models can predict productivity
responses of young forests to elevated CO
and climate
change. Simulated climate change effects on forest
growth vary by model, region, and climate change sce-
nario; the direction of change in forest growth is
expected to vary regionally and to depend on the
course of atmospheric CO
and climate change
(Kirilenko & Sedjo, 2007). For example, in lodgepole
pine (Pinus contorta) forests regenerating from fire in
the Yellowstone region, woody production, live
biomass, N mineralization, and total ecosystem C are
projected to increase under two different future climate
scenarios, with percent increase depending on the cli-
mate scenario (Smithwick et al., 2009). Thus, models
demonstrate likely changes in forest productivity under
future climates; however, without giving specific atten-
tion to changes in physiology and C allocation with for-
est age, they say little about the responses of
regenerating forests specifically.
Because disturbance type, severity, size, and fre-
quency affect postdisturbance C dynamics and biogeo-
chemical cycling (Amiro et al., 2010; Smithwick, 2011),
future trajectories of forest recovery are likely to be dri-
ven by climate changedisturbance type interactions. In
most models, disturbance events are generally imple-
mented by altering forest biomass pools through remo-
vals (harvest), combustion (fire), or transfer of live to
dead material (insect outbreaks), with the amount
transferred scaled to disturbance severity. For fire and
insect outbreaks, the timing of transfer of biomass to
litter and forest floor components varies because tree
death can occur slowly, and snag fall rates are depen-
dent on a variety of factors including forest type
(Campbell et al., 2007; Edburg et al., 2011). At this time,
we are unaware of any model capable of representing
the specific dynamics (e.g., recruitment, altered hydrol-
ogy, or biogeochemistry) associated with distinct dis-
turbance types, severities, and sizes. Therefore, models
currently say little about how changing disturbance
severity and size are likely to impact forests; however,
they do reveal how altered disturbance frequency is
likely to impact forests. For example, in the Yellow-
stone region, fire burn area and frequency are projected
to increase under a range of future climate scenarios,
quite possibly to the extent that current forest commu-
nities will have insufficient time to recover before the
next fire event, making the current suite of conifer spe-
cies unlikely to persist (Westerling et al., 2011). Thus,
models demonstrate that climate change is likely to
have significant impacts on forested landscapes
through its influence on disturbance regimes.
Despite their uncertainties, ESMs have demonstrated
that forest recovery will be substantially altered under
future climates. Rates of recovery will change, with
direction and magnitude varying regionally and
depending on future courses of atmospheric CO
climate change. Altered disturbance regimes will inter-
Published 2013. This article is a U.S. Government work and is in the public domain in the USA., Global Change Biology, doi: 10.1111/
act with altered recovery trajectories, at times driving
biome shifts (Westerling et al., 2011). In combination,
the direct and indirect effects of climate change are pre-
dicted to have substantial impacts on regional C bal-
ances and forestry operations; for example, harvestable
forest biomass in Canada is projected to be reduced
2662% for the 21st century, depending on the model
assumptions of predicted growth rate, soil carbon
decay rate, and area burned by fire (Metsaranta et al.,
2011). However, specific representation of physiological
and community changes associated with forest age
(Table 1) will be required to understand how forest
recovery trajectories will be altered by climate change
(Fig. 1).
As reviewed above, there is strong evidence that
increasing atmospheric CO
, warming, and altered
precipitation regimes will alter trajectories of forest
recovery. This conclusion is supported by global patterns
in both forest regrowth rates and biomass of mature
forests (Fig. 2); responses of forests of various ages to
, temperature, and precipitation manipulation
(Fig. 3; Tables S1S3); observations of altered forest
recovery under contemporary multivariate environ-
mental change (Fig. 4); our understanding of succes-
sional community dynamics and alternative stable
states (Fig. 5); and models. Because forests undergo
major structural, physiological, biogeochemical, and
compositional changes as they age (Table 1), it is logical
that responses to climate change vary as a function of
forest age (Fig. 3). Depending on differential responses
of forests of different ages, climate change can impact
rates of forest recovery, states of mature forests, and/or
recovery pathways (Fig. 1, Table 2), and understanding
the impact of climate change on forests therefore
requires attention to the role of forest age (Fig. 1).
Through its influence on young forests, climate
change will impact rates of forest recovery (Fig. 1,
Table 2). Multiple lines of evidence point to accelerated
regrowth in mesic northern forests under future climates
(Figs 24; Tables S1S2); however, responses of tropical
forest regeneration rates to elevated CO
and increasing
temperature remain uncertain. For forests globally, there
is strong evidence that biomass accumulation rates will
decrease under more arid conditions (Figs 2b and 3;
Table S3) sometimes to the point where forests may
never recover (Fig. 5). Changes to rates of nutrient
accumulation in biomass, biogeochemical cycling, and
community change are likely to parallel responses of
biomass accumulation rate (Table 2).
Climate change will also impact the state toward
which forests converge as they age (Fig. 1; Table 2). A
challenge of central importance is in understanding
how climate change responses of young forests on
which the majority of manipulative experiments have
been performed (Tables S1S3) relate to the ultimate
state of these ecosystems once they reach ‘maturity’
(Fig. 1). For instance, we do not know whether
increased biomass accumulation in young forests will
translate to increased biomass of old forests or whether
these forests will simply attain maximum biomass fas-
ter. The effect of elevated CO
on mature forest biomass
and total ecosystem C remains uncertain, although
decreases in either are unlikely; meanwhile, elevated
is very likely to result in increased nutrient limita-
tion (Fig. 3; Table S1). Likewise, it remains unclear how
warming will affect mature forest biomass and ecosys-
tem C stocks (Fig. 3); it is likely that aboveground C
stocks will increase in northern climates (Fig. 2c) while
soil C stocks decrease and N mineralization increases
(Table S2). In contrast, changes in water availability
have predictable effects; reduced water availability will
reduce productivity, live biomass, and total ecosystem
C stocks (Figs 2d and 3; Table S3). In all cases, altered
community composition is very likely (Table S3).
Responses of mature forest states to combined changes
in CO
, temperature, and precipitation will vary region-
ally, and understanding how the states toward which
future forests will converge as they recovery from dis-
turbance (Fig. 1) remains an important challenge.
Climate change is also likely to impact pathways of
forest recovery (Fig. 1; Table 2), which may occur
through a variety of mechanisms including altered bio-
geochemistry (e.g., decreased N limitation during early
stages due to increased N mineralization), changing
biophysical constraints (e.g., reduced frequency of
years with enough precipitation to support seedling
establishment), or altered community dynamics. As
reviewed above, different species within the same com-
munity commonly have substantially different
responses to altered CO
or climate (K
orner et al., 2005;
Mohan et al., 2006, 2007; Seiler et al., 2009; Dawes et al.,
2011), and consequent changes to community structure
may impact ecosystem functioning in ways that cannot
be predicted based solely on characteristic physiologi-
cal responses of dominant taxa (Bolker et al., 1995). For
example, increased liana biomass under future climates
could meaningfully reduce forest biomass (Phillips
et al., 2002; Mohan et al., 2006; Ingwell et al., 2010). Dif-
ferential responses are likely to be most influential early
in succession, when species turnover rate is highest and
trees are most sensitive to environmental variation
(Table 1), and may have an enduring influence on com-
munity composition and ecosystem function (D’Anto-
nio & Vitousek, 1992; Bunker et al., 2005; Beck et al.,
2011; Hooper et al., 2012).
Published 2013. This article is a U.S. Government work and is in the public domain in the USA., Global Change Biology, doi: 10.1111/
Table 2 Probable climate change impacts on trajectories of several forest properties following disturbance (sensu Fig. 1)
Forest property
Expected response to climate change
Elevated CO
Elevated temperature* Altered water availability Multivariate change
Biomass Rate of
Very likely increase. Very likely increase in temperate
and boreal forests; Uncertain
response in tropics.
Very likely increase with water
availability/decrease with
drought stress.
Likely increase in temperate and
boreal forests (absent moisture
stress); Likely decrease under
drought stress; Uncertain response
in tropics.
Uncertain (likely
increase or no change)
Possible changes in some
regions (e.g., increase in cold
Very likely increase with water
availability/decrease with
drought stress.
Likely changes (region and time
frame specific).
Pathway Likely changes driven
by shifts in community
biogeochemistry, or
Likely changes driven by shifts
in community dynamics,
biogeochemistry, or biophysics.
Likely changes driven by shifts
in community dynamics,
biogeochemistry, or biophysics;
High potential for catastrophic
shift to low-biomass (nonforest)
state under severe drought
Likely changes driven by shifts in
community dynamics,
biogeochemistry, or biophysics.
Total C stock Rate of
Very likely increases. Likely increase in temperate and
boreal forests; Uncertain
response in tropics.
Very likely increase with water
availability/decrease with
drought stress.
Likely increase in temperate and
boreal forests (absent moisture
stress); Likely decrease under
drought stress; Uncertain response
in tropics.
Uncertain (likely increase
or no change).
Very likely decrease in soil
organic matter, possible
increase in biomass carbon
(higher latitudes). Net balance
uncertain and likely region
Very likely increase with water
availability/decrease with
drought stress.
Likely changes (region and time
frame specific).
Pathway Likely changes driven by
shifts in community
biogeochemistry, or
Likely concurrent reductions in
soil organic C and increases in
biomass C.
Likely changes driven by shifts
in community dynamics,
biogeochemistry, or biophysics;
High potential for catastrophic
shift to low C (nonforest) state
under severe drought
Likely changes driven by shifts in
community dynamics,
biogeochemistry, or biophysics
(region- and time frame specific).
Biogeochemistry Rate of
Likely acceleration of
nutrient accumulation in
vegetation; Likely
acceleration of C & N
Likely acceleration of nutrient
accumulation in vegetation in
northern forests; Likely
acceleration of C & N cycling in
temperate and boreal forests;
Likely acceleration of nutrient
accumulation in vegetation
with increased water
availability/rate decrease with
drought stress.
Likely acceleration of nutrient
accumulation in vegetation in
mesic northern forests; Likely rate
decrease under drought stress;
Uncertain response in tropics.
Published 2013. This article is a U.S. Government work and is in the public domain in the USA., Global Change Biology, doi: 10.1111/
Table 2 (continued)
Forest property
Expected response to climate change
Elevated CO
Elevated temperature* Altered water availability Multivariate change
cycling in temperate and
boreal forests.
Uncertain response in the
Likely acceleration of C &
N cycling in temperate
and boreal forests; Likely
decrease in soil N pool;
Uncertainty changes
total nutrient storage in
Likely acceleration of C & N
cycling in temperate and boreal
forests; Likely decrease in soil N
pool; Uncertain response in the
Likely decrease in nutrient
limitation under drought stress.
Likely acceleration of C & N cycling
in temperate and boreal forests;
Likely decrease in soil N pool;
Possible increase in total nutrient
storage in vegetation.
Pathway Likely increase in nutrient
Likely concurrent reductions in
soil N and increases in biomass N.
Likely changes driven by shifts
in community dynamics.
Likely increase in nutrient
limitation in mesic forests.
Rate of
Likely acceleration of
community change.
Likely acceleration of
community change; Uncertain
response in the tropics.
Uncertain (drought may reduce
the rate of community change
through retarded plant growth
or enhance it by increasing
Likely acceleration in temperate
and boreal forests (absent moisture
stress); Uncertain response in
Very likely alteration of
mature community
composition driven by
differential species
responses; Likely
increase in liana
Very likely alteration of mature
community composition driven
by differential species
Very likely alteration of mature
community composition driven
by differential species
Very likely alteration of mature
community composition driven by
differential species responses;
Likely increase in non-native
Pathway Very likely alteration
driven by differential
size- and species
responses, sometimes
causing shift to
alternative state.
Very likely alteration driven by
differential size- and species
responses, sometimes causing
shift to alternative state.
Very likely alteration driven by
differential size- and species
responses, sometimes causing
shift to alternative state.
Very likely alteration driven by
differential size- and species
responses, sometimes causing shift
to alternative state.
*Elevated temperature responses assume no change in moisture stress; responses to changes in water availability are listed in ‘Altered water availability’ column.
‘Rate of change’ refers to the rate at which the forest approaches its mature state. ‘Mature state’ refers to the state to which forests converge as they age. ‘Pathway’ refers to the
sequence of states through which any given ecosystem property passes and the relative amount of time spent in each.
Published 2013. This article is a U.S. Government work and is in the public domain in the USA., Global Change Biology, doi: 10.1111/
In the most dramatic cases, altered successional
pathways may result in catastrophic shifts to an
alternate stable state (e.g., a forest to grassland tran-
sition; Fig. 5). There are documented instances
where, following disturbance, young forests fail to
establish or persist under conditions that are amena-
ble to persistence of mature forests (Thompson &
Spies, 2010; Roccaforte et al., 2012). When these con-
ditions are linked to climate, as they often are (e.g.,
sufficient moisture for seedling establishment, fire
regime), climate change is likely to force a transition
to an alternate stable state (Fig. 5). As a result,
directional changes to forest ecosystems that would
happen gradually in the absence of disturbance may
be greatly accelerated by disturbance (Fig. 5).
There remain several important unanswered ques-
tions regarding the impact of climate change on the
dynamics of forest recovery:
(1) How does forest age modulate responses to climate
change? Forests of different ages have responded differ-
ently to climate manipulations (Fig. 3; Tables S1S3);
however, at this point climate manipulation experi-
ments provide only circumstantial evidence of age dif-
ferences in climate change response. Systematic
comparison of responses of forests of different ages to
experimental CO
or climate manipulation and to natu-
ral climate variability will be crucial to understanding
and modeling climate change impacts on forests of all
(2) How will successional trajectories differ under future
climates? Beyond understanding how age modulates
forest responses to climate change, we face the chal-
lenge of understanding how climate change will impact
entire trajectories of forest recovery (Fig. 1). It is impor-
tant to note that, because the climate history under
which a stand has developed affects its current state
and future trajectory, changes to entire trajectories can-
not be understood simply by integrating across
responses of forests different ages. Rather, it will be
important to understand how altered biogeochemical
dynamics and community composition shape succes-
sional pathways and the states toward which forests
converge as they mature.
(3) Where and when will state changes occur? Climate
change-driven regime shifts (Fig. 5) will have dramatic
consequences, yet they remain difficult to document
and predict. There is a need for experimental, observa-
tional, and modeling studies to identify the conditions
under which such shifts are likely and the mechanisms
through which they may occur.
(4) How will tropical forest regeneration respond to cli-
mate change? Although tropical forests are well repre-
sented in global-scale comparisons (Fig. 2),
precipitation manipulation experiments (Table S3), and
long-term monitoring of mature forests (e.g., Chave
et al., 2008), we are aware of only one study manipulat-
ing CO
or temperature at the whole-tree level in a field
setting in the tropics (Cheesman & Winter, 2012). This
constrains our ability to predict climate change
responses of tropical forests. Understanding how
climate change will affect tropical forest regeneration is
particularly important given the widespread use of
slash-and-burn agriculture in the tropics and the signif-
icant role of tropical forest regrowth in the global C
cycle (Pan et al., 2011).
An additional challenge lies in improved representa-
tion of forest recovery dynamics in ESMs, which are
currently simplistic in their treatment of forest recovery
dynamics. Although detailed representation of forest
recovery dynamics in global models is infeasible, we
believe that two advances will be important to improv-
ing the treatment of forest regeneration. First, the most
important stand age-dependent physiology and alloca-
tion strategies (driven by aging of dominant species
and changes in species composition) should be identi-
fied and incorporated. This will allow improved repre-
sentation of the dynamics of forest recovery in current
and future climates. Importantly, this will help to iden-
tify situations where young forests fail to establish
despite the persistence of their mature counterparts,
suggesting climate change-driven regime shifts (Fig. 5).
Second, although modeling individual species in ESMs
is infeasible, it will be necessary to represent the conse-
quences of demonstrated variability in species
responses to climate change and inevitable resultant
shifts in community composition and ecosystem pro-
cesses. With changing community composition, the net
ecosystem response may differ significantly from that
which would be predicted based on mean characteris-
tics of the original community (Bolker et al., 1995). In
the most dramatic cases, altered competitive interac-
tions may result in a regime shift from forest to a grass-
or shrub-dominated state (Fig. 5). Predicting regime
shifts in ESMs will be particularly important, as these
imply feedbacks to the climate system through altered
C storage, albedo, and hydrology.
Changes in the dynamics of forest recovery following
disturbance will result in potentially significant climate
feedbacks. Altered disturbancerecovery dynamics may
impact the C cycle enough to reverse the sign of a regio-
nal C cycle feedback (Kurz et al., 2008; Running, 2008;
Metsaranta et al., 2011). Moreover, albedo and evapo-
transpiration are important components of the climate
regulation services of ecosystems (Anderson-Teixeira
et al., 2012), change systematically over the course of for-
est recovery (Randerson et al., 2006; Kirschbaum et al.,
2011; Jin et al., 2012; O’Halloran et al., 2012), and may
shift substantially in response to climate change partic-
Published 2013. This article is a U.S. Government work and is in the public domain in the USA., Global Change Biology, doi: 10.1111/
ularly if the new community differs dramatically from
the old (Beck et al., 2011). Altered forest recovery
dynamics will result in particularly strong feedbacks to
climate change when a critical threshold is passed such
that forests fail to recover (Fig. 5), resulting in dramatic
reductions in C storage and altered biophysical proper-
ties. For example, in semiarid regions such as the US
southwest, current forest communities may not be sup-
ported under future more arid conditions and may not
re-establish following disturbance, resulting in a posi-
tive C cycle feedback (Breshears et al., 2005; Williams
et al., 2010, 2013; Anderson-Teixeira et al., 2011; Rocca-
forte et al., 2012). Thus, recently disturbed forests may
play a key role in shaping terrestrial feedbacks to climate
This review has demonstrated that the dynamics of
forest recovery are likely to be significantly impacted
by rising atmospheric CO
and climate change. This
will have repercussions for biodiversity, climate, and
even economics, as the forestry industry and emerging
woody bioenergy industry stand to be affected by
altered forest regeneration rates (Kirilenko & Sedjo,
2007; Metsaranta et al., 2011; Hanewinkel et al., 2013).
Because the course of forest recovery shapes forest
structure and function for decades or centuries, climate
change impacts on secondary forests will have a lasting
legacy. Although the proportion of recently disturbed
forests is relatively small at any given time, disturbance
eventually affects all forests, and the proportion of for-
ests that have regenerated under altered climate condi-
tions will steadily grow. In these ways, climate change
will broadly impact forested regions through its influ-
ence on forest recovery dynamics.
Thanks to Jonathan Thompson and Sean McMahon for helpful
discussion and to three anonymous reviewers for helpful com-
ments. This research was supported by DOE grant
#DE-SC0008085 to KJAT, EHD, and BDD.
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Supporting Information
Additional Supporting Information may be found in the
online version of this article:
Table S1. Summary of experimental manipulations of CO
in tree-dominated ecosystems through Free-Air Carbon
dioxide Enrichment (FACE), Open Top Chamber (OTC), or
Whole Tree Chamber (WTC; in situ only) methodology.
Table S2. Summary of experimental warming in tree-domi-
nated ecosystems (listed in order of forest age).
Table S3. Summary of experimental manipulations of pre-
cipitation (PPT) in tree-dominated ecosystems (listed in
order of forest age).
Published 2013. This article is a U.S. Government work and is in the public domain in the USA., Global Change Biology, doi: 10.1111/
... Recovery is a key ecological process that influences forest stability, structure, and function in the face of extreme events (Anderson-Teixeira et al. 2013;Hodgson et al. 2015;Donohue et al. 2016). Tree recovery is defined as the capacity of a system to return to a stable state and function after a disturbance (Ingrisch & Bahn 2018;Gessler et al. 2020). ...
... low NDVI values). Such repetitive droughts can alter the recovery time and push the forest beyond the "tipping point"-associated with tree mortality -and lead to a deterioration of the forest ecosystem (Anderson-Teixeira et al. 2013;Schwalm et al. 2017). We showed that the NDVI in 2015 is slightly lower that in 2020, most likely due to the 2015 drought period and heatwave from Central Europe that had a lower intensity than in 2018. ...
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Forests worldwide are increasingly exposed to extreme weather events. Drought deteriorates the health, structure, and functioning of forests, which can lead to reduced diversity, decreased productivity, and increased tree mortality. Therefore, it is an urgent need to assess the impact of drought on tree species. Due to differences in tree physiology, saplings and mature trees are likely to respond specifically to drought conditions. In contrast to mature trees, little is known about the response of saplings to drought. Here, we combine in-situ field measurements for saplings of deciduous tree species with remote sensing for forest canopy to assess drought damage, recovery, and sapling mortality patterns during a centennial drought (2018, 2019) and beyond (2020). We measured 2051 saplings out of 214 plots in Central Germany. Forest canopy health was assessed using 10 × 10 m resolution satellite observations for the same locations. We (1) demonstrate that forest canopy exhibits long-lasting drought-induced effects, (2) show that saplings have a remarkable capacity to recover from drought and survive a subsequent drought, (3) demonstrate that reduced sapling recovery leads to their mortality, (4) reveal that drought damage on saplings increases from pioneer to non-pioneer species, and mortality is ranking from Sorbus aucuparia > Sambucus nigra > Fraxinus excelsior, Acer campestre, Frangula alnus > Ulmus glabra > Carpinus betulus > Betula pendula, Fagus sylvatica > Acer pseudoplatanus > Quercus petraea > Corylus avellana, Crataegus spp., > Prunus avium, Quercus robur; and (5) link drought response to site conditions, indicating that species diversity and winter precipitation as relevant indicators of tree health. If periods of drought become more frequent, as expected, this could negatively impact mid-term forest recovery, alter long-term tree species assemblages and reduce biodiversity and functional resilience of forest ecosystems. We suggest that models of forest response to drought should differentiate between the forest canopy and understory and also consider species-specific responses as we found a broad spectrum of responses within the same plant functional type of deciduous tree species in terms of drought damage and recovery.
... Resource (e.g., water and nitrogen (N)) availability is considered as the major limiting factor in regulating secondary succession of plant communities Powers & Marín-Spiotta, 2017;Seabloom et al., 2020). Alterations in water and N availabilities associated with global change (e.g., changing precipitation regime and atmospheric N deposition) may accelerate or slow down the successional processes in plant communities directly through impacting recovery of plant growth and productivity after disturbance and indirectly by affecting species competition (Anderson-Teixeira et al., 2013;Clark et al., 2019;Poorter et al., 2016;Seabloom et al., 2020). ...
... Increased precipitation can stimulate plant growth and the accumulation rate of aboveground biomass by enhancing soil water availability (Anderson- Teixeira et al., 2013;Poorter et al., 2016;Wang et al., 2021). The enhanced plant productivity could provide more C substrate supply for root growth and soil microbial activity , and consequently increase soil respiration during secondary succession. ...
Ecological succession after disturbance plays a vital role in influencing ecosystem structure and functioning. However, how global change factors regulate ecosystem carbon (C) cycling in successional plant communities remains largely elusive. As part of an eight‐year (2012‐2019) manipulative experiment, this study was designed to examine the responses of soil respiration and its heterotrophic component to simulated increases in precipitation and atmospheric nitrogen (N) deposition in an old‐field grassland undergoing secondary succession. Over the eight‐year experimental period, increased precipitation stimulated soil respiration by 11.6%, but did not affect soil heterotrophic respiration. Nitrogen addition increased both soil respiration (5.1%) and heterotrophic respiration (6.2%). Soil respiration and heterotrophic respiration linearly increased with time in the control plots, resulting from changes in soil moisture and shifts of plant community composition from grass‐forb codominance to grass dominance in this old‐field grassland. Compared to the control, increased precipitation significantly strengthened the temporal increase of soil respiration through stimulating belowground net primary productivity. By contrast, N addition accelerated temporal increases of both soil respiration and its heterotrophic component by driving plant community shifts and thus stimulating soil organic C. Our findings indicate that increases in water and N availabilities may accelerate soil C release during old‐field grassland succession and reduce their potential positive impacts on soil C accumulation under future climate change scenarios.
... Temperate forests exhibit characteristic trajectories following disturbances that occur with predictable frequency (Runkle, 1985). However, changing nutrient, temperature, and precipitation regimes will likely alter community composition following disturbances, filtering recovering communities, and may initiate novel successional pathways (Anderson-Teixeira et al., 2013). In annual plants, temperate forests, and desert fish, community life history composition is filtered by the long-term environmental regime to maximize population growth based on demographic constraints. ...
Environmental regimes, which encompass decadal‐scale or longer variation in climate and disturbance, shape communities by selecting for adaptive life histories, behaviors, and morphologies. In turn, at ecological timescales, extreme events may cause short‐term changes in composition and structure via mortality and recolonization of the species pool. Here, we illustrate how short‐term variation in desert stream fish communities following floods and droughts depends on the context of the long‐term flow regime through ecological filtering of life history strategies. Using quarterly measures of fish populations in streams spanning a 10‐fold gradient in flow variation in Arizona, USA, we quantified temporal change in community composition and life history strategies. In streams with highly variable flow regimes, fish communities were less diverse, fluctuation in species richness was the principle mechanism of temporal change in diversity, and communities were dominated by opportunistic life history strategies. Conversely, relatively stable flow regimes resulted in more diverse communities with greater species replacement and dominance of periodic and equilibrium strategies. Importantly, the effects of anomalous high‐ and low‐flow events depended on flow regime. Under more stable flow regimes, fish diversity was lower following large floods than after seasons without floods, whereas diversity was independent of high‐flow events in streams with flashier flow regimes. Likewise, community life history composition was more dependent on antecedent anomalous events in stable compared to more temporally variable regimes. These findings indicate that extreme events may be a second‐level filter on community composition, with effects contingent on the long‐term properties of the disturbance regime (e.g., overall degree of variation) in which extremes take place. Ongoing changes to global environmental regimes will likely drive new patterns of community response to extreme events.
... For communities dominated by long-lived and slowgrowing organisms, the study of recruitment dynamics is particularly timely as mortality events linked to climate driven disturbances are becoming increasingly severe (Anderson-Teixeira et al. 2013;Hughes et al. 2019;Schweiger et al. 2020). Such systems typically possess an ensemble of strategies such as seed banks, clonality, and asexual propagation that allow rapid recovery following perturbation. ...
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For many organisms, early life stages experience significantly higher rates of mortality relative to adults. However, tracking early life stage individuals through time in natural settings is difficult, limiting our understanding of the duration of these ‘mortality bottlenecks’, and the time required for survivorship to match that of adults. Here, we track a cohort of juvenile corals (1–5 cm maximum diameter) from 12 taxa at a remote atoll in the Central Pacific from 2013 to 2017 and describe patterns of annual survivorship. Of the 537 juveniles initially detected, 219 (41%) were alive 4 years later, 163 (30%) died via complete loss of live tissue from the skeleton, and the remaining 155 (29%) died via dislodgement. The differing mortality patterns suggest that habitat characteristics, as well as species-specific features, may influence early life stage survival. Across most taxa, survival fit a logistic model, reaching > 90% annual survival within 4 years. These data suggest that mortality bottlenecks characteristic of ‘recruitment’ extend up to 5 years after individuals can be visually detected. Ultimately, replenishment of adult coral populations via sexual reproduction is needed to maintain both coral cover and genetic diversity. This study provides key insights into the dynamics and time scales that characterize these critical early life stages.
... Competition for water, which may intensify under drought, is particularly high in regrowing forests as a result of fast growth rates of understory vegetation, high stem densities, and dominance of water-demanding species (Uriarte et al., 2016b). Despite these potential vulnerabilities, research on the factors that determine how drought may influence post-hurricane ecosystem recovery is lacking (Anderson-Teixeira et al., 2013;Uriarte et al., 2016b;Bretfeld et al., 2018;McDowell et al., 2020). ...
Rapid changes in climate and disturbance regimes, including droughts and hurricanes, are likely to influence tropical forests, but our understanding of the compound effects of disturbances on forest ecosystems is extremely limited. Filling this knowledge gap is necessary to elucidate the future of these ecosystems under a changing climate. We examined the relationship between hurricane response (damage, mortality, and resilience) and four hydraulic traits of 13 dominant woody species in a wet tropical forest subject to periodic hurricanes. Species with high resistance to embolisms (low P50 values) and higher safety margins (SMP50) were more resistant to immediate hurricane mortality and breakage, whereas species with higher hurricane resilience (rapid post‐hurricane growth) had high capacitance and P50 values and low SMP50. During 26 yr of post‐hurricane recovery, we found a decrease in community‐weighted mean values for traits associated with greater drought resistance (leaf turgor loss point, P50, SMP50) and an increase in capacitance, which has been linked with lower drought resistance. Hurricane damage favors slow‐growing, drought‐tolerant species, whereas post‐hurricane high resource conditions favor acquisitive, fast‐growing but drought‐vulnerable species, increasing forest productivity at the expense of drought tolerance and leading to higher overall forest vulnerability to drought.
... Climate change directly affects forest ecosystems through physiological and demographic processes such as photosynthesis, respiration, growth, mortality, and species competition, which catalyze the dynamics of forest succession (Anderson-Teixeira et al., 2013;Boulanger et al., 2017;Rüger et al., 2020). Climate change also indirectly affects forest ecosystems by altering the frequency and intensity of fire disturbances (McDowell et al., 2020;Seidl et al., 2017). ...
Climate change could alter species composition, with feedback on fire disturbances by modifying fuel types and loads. However, the existing fire predictions were mainly based on climate-fire linkages that might overestimate the probability and size of fire disturbances due to simplifying or omitting vegetation feedback. We applied a model-coupling framework that combines forest succession, climate-fire linkages, and vegetation feedback to predict burned area, aboveground biomass, and species composition of boreal forests in Northeast China under climate change conditions. Results showed that climate change and fire would favor the recruitment of deciduous species, but these species need a long-time to replace the existing coniferous species. Burned area would increase with climate change. Climate change, historical and future fire disturbances affect aboveground biomass by altering tree mortality and regeneration. Further studies should address strategies for altering species composition through forest management practices to adaptation climate change and reduce carbon losses from fire.
... With a more direct impact on forest regeneration, global warming is also expected to increase the frequency and intensity of fires in tropical forests, resulting in negative consequences to species persistence and to the growth of early forests (Herawati & Santoso, 2011). Added to the effects of fire, prolonged droughts caused by climate change may hinder seedling recruitment and survival, which may further compromise natural regeneration (Anderson-Teixeira et al., 2013). ...
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Aim Evaluate how large‐scale forest regeneration based on a low‐cost restoration method may mitigate the effects of habitat loss and fragmentation associated to future climate changes on the distribution of birds and arboreal mammals in a tropical biodiversity hotspot; find areas with different current and future potential species richness and assess how passive restoration can reduce the risk of species extinction. Location Brazilian Atlantic Forest (BAF). Methods We built a forest regeneration scenario via a model of seed dispersal based on the potential movement of frugivorous fauna and projected the potential distribution of 356 bird species and 21 arboreal mammals based on Species Distribution Models (SDM) which employed 79,462 occurrence records and four algorithms for different climate and landscape scenarios. SDM were based on climate and landscape predictors separately and the results were combined into maps of species richness. Finally, we assessed the species’ risk of extinction based on the species–area relationship. Results Without considering the effects of climate change, the potential distribution area for each species increases on average by 72.5% (SD = 8%) in the scenario of potential regeneration. Climate change decreases the area of potential occurrence of 252 species, which may suffer a mean reduction of 74.4% (SD = 9.3%) in their current potential distribution areas. BAF regions with the largest amounts of forest had the greatest potential richness of species. In future climate scenario, 3.4% of species may become extinct, but we show that large‐scale regeneration may prevent these extinctions. Main conclusions Despite the possible negative impacts of climate change on the distribution of 67% of the studied species, which would increase the risk of species extinction, our analysis indicated that promoting large‐scale BAF restoration based on natural regeneration may prevent biodiversity loss.
... The method works as an important process of structuring forest communities over time (Brooker et al., 2007;Morrison and Lindell, 2011;Navarro-Cano et al., 2016). This allows for greater provision of ecosystem services, including carbon accumulation in living biomass and nutrient cycling (Anderson-Teixeira et al., 2013;Poorter et al., 2016). ...
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Mining contributes to the global economy on different scales and plays a fundamental role in the development of the goods and services sectors. However, the negative impacts caused by the activity are unavoidable, as they intensely degrade soil structures and modify landscapes. The use of native tree species has been effective in restoring the structure and functions of post-mining ecosystems in the Amazon. Thus, the objective of this work was to evaluate chemical indicators of soil quality, survival and initial growth of six tree species planted in degraded ecosystem by kaolin mining under the effects of liming in three pit volumes in the Eastern Amazon. The initial conditions indicated highly degraded soil, with acidic pH, low content of OM, P and K. Liming significantly reduced the level of toxic Al in the soil and provided Ca and Mg for the plants. Through Principal Component Analysis (PCA), we found that the two first components explained 69.30% of the variance of 13 functional indicators of soil quality. PC1 was positively correlated with Ca, Mg, K, SB, CEC and V% and negatively correlated with H+Al and Al saturation. Survival was higher than 80% for Clitoria fairchildiana in all treatments. Liming and pit volume were determining in growth, with high growth rates in height of Inga edulis, Inga cayennensis, Clitoria fairchildiana and Tachigali vulgaris. The species used in this study presented good initial development in restoring ecosystems after kaolin mining.
... Lastly, the impacts of climate change could lead some forest systems to reach critical ecological thresholds, where conversion to different forest types (e.g., the conversion of black spruce to deciduous forest in interior Alaska following severe wildfire [Hansen et al. 2020]), or even to non-forested systems, is possible (Koch et al. 2009, Adams 2013, Anderson-Teixeira et al. 2013, Allen et al. 2015, Teskey et al. 2015, Walker et al. 2018, Busby et al. 2020, Harris and Taylor 2020. Research by Parks et al. (2019a), for instance, suggests that a significant percentage of The impacts of climate change could lead some forest systems to reach critical ecological thresholds, where conversion to different forest types, or even to nonforested systems, is possible. ...
Technical Report
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A rapidly changing climate, including rising temperatures, changing precipitation patterns, and more extreme storms, is having profound consequences for America’s national forests. Climate-related impacts on forest systems include larger and more severe disturbances (e.g., wildfires, drought, and insect outbreaks), shifts in tree species ranges and forest composition, and changes in forest dynamics and regeneration capacity. Many of our national forests have been significantly modified by past management and land use, and forest managers are contending with ongoing threats from invasive species, disease outbreaks, and other challenges. With the added impacts on forest systems from climate change, an enormous mismatch exists between the level of restoration work currently underway and the scale of the challenge. As a result, there is a need to substantially increase the pace, scale, and quality of restoration on our national forests, and to ensure that this restoration is carried out in an ecologically appropriate and climate-smart manner. Continuing and accelerating climatic changes, and their associated impacts, have significant implications for the effectiveness of traditional forest restoration efforts, including reliance on historical conditions as benchmarks for restoration outcomes. Drawing on a growing body of evidence, research, and experimentation, this science review and synthesis looks at how climate change is inspiring an important evolution in approaches for national forest restoration and management. Over the past decade, the U.S. Forest Service has made considerable progress in understanding the effects of a changing climate on forest ecosystems and working to incorporate climate considerations into its planning and management. Nonetheless, varying perspectives on what climate change means for ecological restoration in practice and how to navigate potential trade-offs continue to pose challenges to integrating climate adaptation and mitigation in national forest planning and management. Addressing this challenge would benefit from a shared understanding among agency staff and stakeholders of what constitutes a forward-looking and climate-smart approach to national forest restoration. To this end, this report reviews and summarizes recent advances and ongoing evolution in how the concepts and principles of climate adaptation and mitigation can help promote the development and application of climate-smart forest restoration.
Technical Report
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In this report, we summarize the current state of knowledge and best estimates of how climate change is expected to impact Norwegian forest ecosystems from now to the year 2100
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While ecological dogma holds that rates of community change decrease over the course of succession, this idea has yet to be tested systematically across a wide variety of successional sequences. Here, I review and define several measures of community change rates for species presence‐absence data and test for temporal patterns therein using data acquired from 16 studies comprising 62 successional sequences. Community types include plant secondary and primary succession as well as succession of arthropods on defaunated mangrove islands and carcasses. Rates of species gain generally decline through time, whereas rates of species loss display no systematic temporal trends. As a result, percent community turnover generally declines while species richness increases—both in a decelerating manner. Although communities with relatively minor abiotic and dispersal limitations (e.g., plant secondary successional communities) exhibit rapidly declining rates of change, limitations arising from harsh abiotic conditions or spatial isolation of the community appear to substantially alter temporal patterns in rates of successional change.
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In a throughfall displacement experiment on Sulawesi, Indonesia, three 0.16 ha stands of a premontane perhumid rainforest were exposed to a two-year soil desiccation period that reduced the soil moisture in the upper soil layers beyond the conventional wilting point. About 25 variables, including leaf morphological and chemical traits, stem diameter growth and hydraulic properties of the xylem in the trunk and terminal twigs, were investigated in trees of the tall-growing tree species Castanopsis acuminatissima (Fagaceae) by comparing desiccated roof plots with nearby control plots. We tested the hypotheses that this tall and productive species is particularly sensitive to drought, and the exposed upper sun canopy is more affected than the shade canopy. Hydraulic conductivity in the xylem of terminal twigs normalised to vessel lumen area was reduced by 25%, leaf area-specific conductivity by 10–33% during the desiccation treatment. Surprisingly, the leaves present at the end of the drought treatment were significantly larger, but not smaller in the roof plots, though reduced in number (about 30% less leaves per unit of twig sapwood area), which points to a drought effect on the leaf bud formation while the remaining leaves may have profited from a surplus of water. Mean vessel diameter and axial conductivity in the outermost xylem of the trunk were significantly reduced and wood density increased, while annual stem diameter increment decreased by 26%. In contradiction to our hypotheses, (i) we found no signs of major damage to the C. acuminatissima trees nor to any other drought sensitivity of tall trees, and (ii) the exposed upper canopy was not more drought susceptible than the shade canopy.
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Increased carbon storage with afforestation leads to a decrease in atmospheric carbon dioxide concentration and thus decreases radiative forcing and cools the Earth. However, afforestation also changes the reflective properties of the surface vegetation from more reflective pasture to relatively less reflective forest cover. This increase in radiation absorption by the forest constitutes an increase in radiative forcing, with a warming effect. The net effect of decreased albedo and carbon storage on radiative forcing depends on the relative magnitude of these two opposing processes. We used data from an intensively studied site in New Zealand's Central North Island that has long-term, ground-based measurements of albedo over the full short-wave spectrum from a developing Pinus radiata forest. Data from this site were supplemented with satellite-derived albedo estimates from New Zealand pastures. The albedo of a well-established forest was measured as 13 % and pasture albedo as 20 %. We used these data to calculate the direct radiative forcing effect of changing albedo as the forest grew. We calculated the radiative forcing resulting from the removal of carbon from the atmosphere as a decrease in radiative forcing of −104 GJ tC<sup>−1</sup> yr<sup>−1</sup>. We also showed that the observed change in albedo constituted a direct radiative forcing of 2759 GJ ha<sup>−1</sup> yr<sup>−1</sup>. Thus, following afforestation, 26.5 tC ha<sup>−1</sup> needs to be stored in a growing forest to balance the increase in radiative forcing resulting from the observed albedo change. Measurements of tree biomass and albedo were used to estimate the net change in radiative forcing as the newly planted forest grew. Albedo and carbon-storage effects were of similar magnitude for the first four to five years after tree planting, but as the stand grew older, the carbon storage effect increasingly dominated. Averaged over the whole length of the rotation, the changes in albedo negated the benefits from increased carbon storage by 17–24 %.
This 2-year field study examined stomatal conductance, photosynthesis, and biomass allocation of Liquidambar styraciflua saplings in response to below- and aboveground competition with the vines Lonicera japonica and Parthenocissus quinquefolia. Vine competition did not affect stomatal conductance of the host trees. The leaf photosynthetic capacity and photosynthetic nitrogen-use efficiency were significantly reduced by root competition with vines, either singly or in combination with aboveground competition, early in the second growing season. However, such differences disappeared by the end of the second growing season. Trees competing below ground with vines also had lower allocation to leaves compared with steins. Aboveground competition with vines resulted in reduced photosynthetic capacity per unit leaf area, but not per unit leaf weight, in trees. No correlation was found between single leaf photosynthetic capacity and tree growth. In contrast, a high positive correlation existed between allocation to leaves and diameter growth. Results from this study suggest that allocation patterns are more affected than leaf photosynthesis in trees competing with vines.
Ecological orthodoxy suggests that old-growth forests should be close to dynamic equilibrium, but this view has been challenged by recent findings that neotropical forests are accumulating carbon and biomass, possibly in response to the increasing atmospheric concentrations of carbon dioxide. However, it is unclear whether the recent increase in tree biomass has been accompanied by a shift in community composition. Such changes could reduce or enhance the carbon storage potential of old-growth forests in the long term. Here we show that non-fragmented Amazon forests are experiencing a concerted increase in the density, basal area and mean size of woody climbing plants (lianas). Over the last two decades of the twentieth century the dominance of large lianas relative to trees has increased by 1.7–4.6% a year. Lianas enhance tree mortality and suppress tree growth, so their rapid increase implies that the tropical terrestrial carbon sink may shut down sooner than current models suggest. Predictions of future tropical carbon fluxes will need to account for the changing composition and dynamics of supposedly undisturbed forests.