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Clim. Past, 10, 1277–1290, 2014
www.clim-past.net/10/1277/2014/
doi:10.5194/cp-10-1277-2014
© Author(s) 2014. CC Attribution 3.0 License.
Treeline dynamics with climate change at the central Nepal
Himalaya
N. P. Gaire1,2, M. Koirala2, D. R. Bhuju1,2, and H. P. Borgaonkar3
1Faculty of Science, Nepal Academy of Science and Technology, Khumaltar, Lalitpur, GPO Box 3323, Kathmandu, Nepal
2Central Department of Environmental Science, Tribhuvan University, Kathmandu, Nepal
3Indian Institute of Tropical Meteorology, Pune, India
Correspondence to: N. P. Gaire (npgaire2007@gmail.com)
Received: 7 August 2013 – Published in Clim. Past Discuss.: 28 October 2013
Revised: 22 April 2014 – Accepted: 15 May 2014 – Published: 4 July 2014
Abstract. Treeline shifting in tandem with climate change
has widely been reported from various parts of the world. In
Nepal, several impacts of climate change on the physical en-
vironment have been observed, but study on the biological
impacts is lacking. This dendrochronological study was car-
ried out at the treeline in the high mountain slope of Kalchu-
man Lake (3750–4003ma.s.l.) area of Manaslu Conserva-
tion Area in the central Nepal Himalaya to explore the impact
of climate change on the treeline dynamic. Two belt transect
plots (size: 20 m wide, >250 m long) were laid which in-
cluded treeline as well as tree species limit. Ecological map-
ping of all individuals of dominant trees Abies spectabilis and
Betula utilis was done and their tree cores were collected.
Stand character and age distribution revealed an occurrence
of more matured B. utilis (max. age 198years) compared to
A. spectabilis (max. age 160years). A. spectabilis contained
an overwhelmingly high population (89%) of younger plants
(<50years) indicating its high recruitment rate. Population
age structure along the elevation gradient revealed an upward
shifting of A. spectabilis at the rate of 2.61myear−1since
AD1850. The upper distribution limit of B. utilis was found
to be stagnant in the past few decades. An increment in plant
density as well as upward shifting in the studied treeline eco-
tones was observed. The temporal growth of A. spectabilis
was correlated negatively with the monthly mean and min-
imum temperature of June to September of the current and
previous year. The regeneration of A. spectabilis, on the other
hand, was positively correlated with August precipitation and
monthly maximum temperature of the month of the current
year. The growth and regeneration of A. spectabilis was more
sensitive to maximum and minimum temperature rather than
average temperature. The growth of the B. utilis was mainly
limited by moisture stress during the pre-monsoon season.
As these two species presented species-specific responses to
climate change with differential pattern in regeneration con-
dition, much wider differences are anticipated in their pop-
ulation status as climate continues to change throughout the
century.
1 Introduction
During the past 100years the global average surface temper-
ature has increased by 0.74◦C±0.2◦C, and it is projected to
rise by 1.4–5.8◦C by AD2100 (IPCC, 2007), with the most
pronounced and rapid changes at high altitudes and latitudes.
However, recent studies have shown spatial and temporal he-
terogeneity in the past long-term temperature trend (Marcott
et al., 2013; PAGES 2k Consortium, 2013). Rapid climate
change has many biophysical impacts (IPCC, 2007) and al-
ready left several biological fingerprints including change in
species composition of ecological communities, range and
distribution shift of species as well as changes in phenology
of the organisms (Parmesan and Yohe, 2003; Root et al.,
2003; Parolo and Rossi, 2008; Chen et al., 2011; Gottfried
et al., 2012; Kirdyanov et al., 2012; Pauli et al., 2012; Webb
et al., 2012).
The high-altitude limit of forests, commonly known as
treeline, timberline or forest line, represents one of the
most conspicuous vegetation boundaries (Körner, 1998;
Holtmeier, 2009). The position of a treeline is mainly due
to strong growth limitation by low-temperature conditions
Published by Copernicus Publications on behalf of the European Geosciences Union.
1278 N. P. Gaire et al.: Treeline dynamics with climate change at the Himalaya
(Körner and Paulsen, 2004; Holtmeier, 2009). Worldwide,
high-altitude climatic treelines are associated with a seasonal
mean ground temperature of 6.7◦C±0.8 SD during the grow-
ing period (Körner and Paulsen, 2004). So, natural treeline
ecotones are sensitive biomonitors of past and recent climate
change and variability (Kullman, 1998), and are well suited
for monitoring climate change impact (Becker et al., 2007).
The high-elevation treeline is assumed to represent an ideal
early-warning feature that responds to climate change po-
sitionally, structurally and compositionally (Kullman, 1998,
2001, 2007; Kirdyanov et al., 2012). Many dendroecologi-
cal studies have documented that trees at the treeline often
respond to climatic warming with an increase in recruitment
or tree density as well as upward advances in the treeline
position (Bradley and Jones, 1993; Camarero and Gutiérrez,
2004; Danby and Hik, 2007; Kullman, 2002, 2007; Batllori
and Gutiérrez, 2008; Kullman and Öberg, 2009; Leonelli
et al., 2011; Kirdyanov et al., 2012). A meta-analysis of a
global data set, including 166 sites for which treeline dynam-
ics had been recorded since AD1900, showed that the tree-
line either advanced (52% of sites) or remained unchanged,
while only few treelines (1%) declined under heavy anthro-
pogenic disturbance (Harsch et al., 2009). Treelines that ex-
perienced strong winter warming and treelines with a diffuse
form are more likely to advance (Harsch et al., 2009).
Himalayan ecosystems are facing the impacts of climate
change. However, uncertainties about our knowledge on the
relationships of Hindu Kush–Karakoram–Himalaya (HKH)
treelines to other ecological conditions and processes such
as carbon balance, freezing and frost, drought, soil temper-
ature, wind, snow cover, soils, regeneration, etc. are yet to
be explored (Schickhoff, 2005). Treelines from Tibet and ad-
jacent mountainous regions have shifted very little with cli-
mate change (Liang et al., 2011; Gou et al., 2012; Lv and
Zhang, 2012). However, a previous study reported an upward
movement of the tree species limit due to climate change in
the Himalayas (Dubey et al., 2003). High-altitude regions in
the interior of the Nepal Himalaya are little affected by an-
thropogenic activities and may therefore provide valuable in-
formation to evaluate the isolated consequences of climate
change (Cook et al., 2003).
The atmospheric temperature of Nepal has been increas-
ing consistently after the mid-1970s with higher rate than the
global average (Shrestha et al., 1999; IPCC, 2007), and the
warming has been found to be even more pronounced in the
high altitudes of the Nepal Himalaya (Shrestha et al., 1999;
Shrestha, 2008). However, no specific trend in precipitation
has been observed (Shrestha, 2008). The effect of warming
temperature in the Nepal Himalaya is reflected by shrink-
ing permafrost areas (Fukui et al., 2007) and rapidly retreat-
ing glaciers (Fujita et al., 1998; Bajaracharya et al., 2007;
Bolch et al., 2012; Yao et al., 2012), among other phenom-
ena. Impacts on biological processes including range shifting
of species are also expected but scientific studies on these as-
pects are scarce (Schickhoff, 2005). Past works on tree rings
in Nepal have identified several promising species for den-
drochronological study, including Abies spectabilis and Be-
tula utilis (Bhattacharyya et al., 1992; Cook et al., 2003; Sano
et al., 2005; Dawadi et al., 2013), which can grow up to the
treeline ecotone (Schickhoff, 2005; Ghimire et al., 2008). Re-
cently, researchers from Nepal have initiated dendroecolog-
ical studies covering various treeline sites of the Nepal Hi-
malaya (Bhuju et al., 2010; Suwal, 2010; Gaire et al., 2011).
However, concrete results on the treeline shifting due to cli-
mate change are yet to be explored.
The present study was carried out to (i) ascertain the
present position of upper forest, treeline and species limits,
(ii) characterize the stand structure and dynamics at the forest
line and treeline, and (iii) analyse the response of tree growth
and regeneration with climate change using both dendroeco-
logical and dendroclimatological techniques. For this study,
the treeline is defined as the ecotone up to where 2 m tall trees
can be found, and the species limit is defined as the highest
position to which seedlings or saplings of the tree species are
present. Treeline dynamics describe changes in the regenera-
tion and population dynamics as well as positional change of
the tree species in the treeline ecotone.
2 Materials and methods
2.1 Site and species selection
The study was carried out at Manaslu Conservation Area
(MCA, area: 1663km2), a high mountain protected site in
the central Nepal Himalaya, established in AD1998. MCA
has a diverse natural resource base with sparse human pop-
ulation and is relatively inaccessible. The area includes nine
bioclimatic zones ranging from the lower subtropics to the
nival zone with only marginal infrastructure, such as roads.
It is the least explored protected area of the country. Local
people depend on agriculture, animal husbandry and utiliza-
tion of natural resources for their sustenance. Buddhism has
positively contributed in protecting the forest and biodiver-
sity (Chhetri, 2009).
The study site is a mountain slope adjacent to Kalchuman
Lake situated at 3690 m above mean sea level (a.m.s.l.). With
human settlements located not more than 2500ma.m.s.l.,
the study site is little disturbed anthropogenically. There is
a dense forest in between the settlement and study sites.
Soil is rich in humus, dark in colour, and its depth varies
locally with the steepness of the slope. The tree canopy
of the treeline ecotone is formed by A. spectabilis and
B. utilis with a Rhododendron campanulatum understory and
some scattered Sorbus microphylla. Above the treeline oc-
cur scrubs of Rhododendron anthopogon and some herba-
ceous species. The Himalayan silver fir, A. spectabilis is a
tall evergreen tree endemic to the Himalaya and found be-
tween the lower temperate and lower alpine zone (2400–
4400m) from Afghanistan to Bhutan (Ghimire et al., 2008).
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N. P. Gaire et al.: Treeline dynamics with climate change at the Himalaya 1279
Similarly, B. utilis is a medium-sized deciduous tree which
forms monospecific as well as mixed forests at the upper
limit of the treeline (Ghimire et al., 2008).
The climate of the study area is monsoon dominated. The
mean annual rainfall over the past 30years (1980–2009)
at nearby meteorological station at Chame was 967mm
(SD = 280). The monthly average temperature was found
to be highest in July and lowest in November (Fig. 1a).
The highest recorded temperatures were 23.4◦C during June
1998, the lowest −4.5◦C during January of 1999 and 2000.
Over the past 30years, the station experienced a decreas-
ing trend in rainfall by 3.9mmyear−1(n=30, R2=0.014,
p < 0.52) (Fig. 1b) and an increasing trend in mean an-
nual temperature by 0.017 ◦Cyear−1(Fig. 1c). In this station
monthly mean minimum temperature was decreasing while
monthly mean maximum temperature was increasing signifi-
cantly (Supplementary Fig. S1). Similarly, mean annual rain-
fall at Larke, Gorkha, was 1252mm (SD = 535). In Larke,
during the past 30years (1980–2009) there was a significant
(n=30, R2=0.26, p < 0.003) decreasing trend of rainfall
by 28mmyear−1(Fig. 1b). This decreasing trend is more
pronounced and significant (p < 0.0003, R2=0.46, n=23)
after 1987 with a decrease in annual rainfall by 55 mm year−1
between 1987 and 2009.
2.2 Field visit and data collection
Field work was carried out in three expeditions: two in
2010 (May–June and September–October) and one in 2012
(October). After careful observation in transect walk at the
treeline ecotone, the upper species limits of A. spectabilis
and B. utilis were ascertained. Two altitudinal transect plots
(20 m wide and >250 m long), named Transect 1 (T1) and
Transect 2 (T2) were marked at two sites of the treeline eco-
tone. The plots were oriented with their longer side parallel
to the maximum slope and covered the current species limit
and treeline ecotone (Fig. 2). T1 was above the continuous
forest, while T2 was situated above the middle part of the
lake. It was also assumed that the lake might hinder the dis-
persal of seedlings in the area. Individual plants were catego-
rized and enumerated into three height classes: trees (>2m),
saplings (0.5–2m) and seedlings (<0.5m), following Wang
et al. (2006) and Kullman (2007).
Census counts were carried out inside each plot for
A. spectabilis and B. utilis. For every A. spectabilis individ-
ual, their geographic location in the plot (latitude, longitude,
and altitude); size (diameter at breast height (DBH), height);
growth form and internodes interval of all individuals less
than 2m were recorded. The age of trees was calculated by
tree core analysis, while that of seedlings and saplings were
estimated by counting the branch whorls and scars left along
the main stem (Camarero and Gutiérrez, 2004; Wang et al.,
2006; Liang et al., 2011). This age estimation was also val-
idated by comparing it with the age obtained by the number
of tree rings in the basal sections collected from the root col-
1980 1985 1990 1995 2000 2005 2010
Annual precipitation (mm)
0
500
1000
1500
2000
2500 Chame
Larke
Jan Feb Mar Apr May Jun Jul Aug Sep Oct Nov Dec
Precipitation (mm)
0
50
100
150
200
250
Temperature (°C)
2
4
6
8
10
12
14
16
Precipitation
Temperature
1980 1985 1990 1995 2000 2005 2010
Average annual temperature (°C)
9.5
10.0
10.5
11.0
11.5
y = 0.017 * x + 10.55
R
2
= 0.114, p = 0.067
(a)
(b)
(c)
y = -3.855 * x + 8657
R
2
= 0.014, p<0.52
y = -28.22 * x + 57365
R
2
= 0.262, p<0.003
Figure 1. Climatic trend in the local stations: mean monthly (1980–
2009) precipitation and temperature at Chame, Manang station (a);
annual trend of precipitation at Larke, Gorkha and Chame (b), and
trend of mean annual temperature at Chame (c).
lar of saplings and seedlings (n=34). The age estimates by
whorl count and ring count correlated positively (R2=0.91,
P < 0.0001), but the internodes or whorl count ages are
systematically lower by 1.57±0.33years (max. 4years) in
saplings and 1.27years in seedlings. These suggest that the
whorl count method can give a fairly accurate indication
of the age of saplings and seedlings of conifer species like
A. spectabilis.
2.3 Tree core and cut-stump collection and analysis
Tree cores were collected using the increment borer (Haglöf,
Sweden) following the standard technique suggested by
Fritts (1976) and Speer (2010). The cores were collected
from the base of each and breast (1.3 m) height of some indi-
viduals of A. spectabilis and B. utilis in the plots. Cores were
also collected from the larger A. spectabilis and B. utilis trees
outside of the plots. A total of 249 cores and cut-stump sam-
ples (172 A. spectabilis and 77 B. utilis) were collected (Sup-
plementary Table S1). Collected core and cut-stump samples
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1280 N. P. Gaire et al.: Treeline dynamics with climate change at the Himalaya
Figure 2. Location map of the study area showing the position of
the study plots and local meteorological stations (a), and a photo of
the study site showing treeline, species limit and some portion of
Kalchuman Lake (b).
were taken to the Dendro-lab of the Nepal Academy of Sci-
ence and Technology for laboratory analysis.
Collected cores were air dried, mounted, sanded and pol-
ished using successively finer grades of sand paper (100–
1000 grits size) until optimal surface resolution allowed an-
nual rings to be visible under the microscope. Each ring
was counted under the stereo zoom microscope and assigned
a calendar year. The width of each ring was measured to
the nearest 0.01mm precision with the LINTABTM mea-
suring system attached to a PC with the TSAP Win soft-
ware package (Rinn, 1996). All the tree cores were cross-
dated by matching patterns of relatively wide and narrow
rings to account for the possibility of ring-growth anomalies
such as missing or false rings or measurement error. Cross-
dating was done using the alignment plotting technique and
also looking the math graphs. The quality of cross-dating
of each sample was checked using the computer program
COFECHA (Holmes, 1983).
The corrected ring-width data were standardized using
the computer program ARSTAN (Cook, 1985). The ring-
width series were standardized using conventional detrend-
ing methods with appropriate options of a negative expo-
nential, linear or cubic spline curve to each series. Each
ring-width-index series was then pre-whitened using autore-
gressive modelling to remove any autocorrelation effects
(Cook, 1987). Finally, three chronologies – namely standard,
residual and ARSTAN – were prepared using the corrected
sample. Various chronology statistics like mean sensitivity,
standard deviation, autocorrelation, mean series correlation,
signal-to-noise ratio, expressed population signal (EPS) and
variance explained were calculated to assess the quality of
the site chronologies. Temporal changes in the mean ra-
dial growth were assessed by doing regime shift (significant
changes in mean radial growth) analysis (Rodionov, 2004).
Regime shift was detected based on a statistical test whereby
data are processed in time sequence and the hypothesis of
a regime shift or discontinuity is tested for each new obser-
vation (Rodionov, 2004, 2006). We set the cutoff parameter
at 10years in order to detect changes in mean radial growth
driven by high-frequency events and used a 95 % level of sig-
nificance.
2.4 Population demography, regeneration and treeline
dynamics
The age obtained from cross-dated samples was used for de-
mographic analysis after the necessary correction for years to
core height and years to centre of missed pith. Such correc-
tion was made using age–height regression and age–diameter
regression combined with the fitting of a circle template to
the ring curvature so as to estimate the distance of the core
to the centre (Camarero and Gutiérrez, 2004; Speer, 2010).
For B. utilis cores from representative sample trees cover-
ing different diameter at breast height (DBH) classes were
collected. Based on the ages of 39 Betula trees, a regres-
sion analysis model between DBH and age was established
(Fig. 3), and the relationship was used to estimate the age of
all B. utilis trees from which tree cores were not taken.
The regeneration rate was determined by age histogram
using the number of seedlings, saplings and tree individuals.
The treeline dynamics was analysed by density distribution
of tree, sapling and seedling as well as the elevation-wise age
distribution of the studied species. The upper species limit
expansion was studied by observing the age of each indi-
vidual in the entire plot following Camarero and Gutiérrez
(2004) and Liang et al. (2011). In order to calculate the rate
of species limit shift, the maximum elevation of live indi-
viduals and the position of the oldest individual within each
transect was determined. Then, the species limit shift rate
(myear−1) was calculated by dividing the change in species
limit elevation (position) by the time elapsed.
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N. P. Gaire et al.: Treeline dynamics with climate change at the Himalaya 1281
DBH (cm)
010 20 30 40 50 60
Age (years)
0
50
100
150
200
250
300
Age = 3.765 * DBH + 23.34
R
2
= 0.847, p < 0.0005
Figure 3. Diameter at breast height (DBH) versus age of Betula
utilis from the Kalchuman Lake area, Manaslu Conservation Area.
2.5 Climatic influence on radial growth and
regeneration
Before proceeding to the response analysis of tree growth and
climate, the seasonality of tree growth was defined. Field ob-
servation and tree ring data have shown that radial growth of
A. spectabilis at treeline sites ceases in September–October
(Sano et al., 2005). Because climate in the preceding grow-
ing season often influences tree growth in the following year
(Fritts, 1976), we analysed the influence of temperature and
precipitation since June of the previous growth year until Oc-
tober of the current growth year. Simple Pearson correlation
coefficients were used to quantify relationships between tree-
ring chronologies and climate variables, i.e. monthly average
(Tmean), maximum (Tmax) and minimum (Tmin) temperature
and total monthly precipitation. Similarly, influence of sea-
sonal climate on radial growth was also assessed for four
seasons – namely winter (December–February, DJF), pre-
monsoon (March–May, MAM), monsoon (June–September,
JJAS) and post-monsoon (October–November, ON). In ad-
dition to this, influence of annual average temperature and
total precipitation was also calculated. One of the major dif-
ficulties in undertaking dendroclimatic research in Nepal re-
lates to the paucity of long meteorological records for statis-
tically calibrating the tree rings because most of the weather
stations in Nepal were only established after 1960 for pre-
cipitation and 1970 for temperature (Bhattacharyya et al.,
1992; Cook et al., 2003). Available climatic data (1980–
2009) of the nearest stations at Chame (28◦330N, 84◦140E
and 2680ma.s.l.) of Manang and Larke Samdo (28◦400N,
84◦370E and 3650ma.s.l.) of Gorkha were used (Fig. 2).
Missing values were replaced by average value of the same
month’s data.
To investigate the relationship between regeneration and
climate change, recruitment or age data were summed across
5-year intervals as the finest resolution to take into account
uncertainties in age estimates and compared with monthly
climate records compiled into 5-year averages over the same
time period (Camarero and Gutiérrez, 1999; Wang et al.,
2006). To describe the climate–recruitment relationships,
monthly climatic data (mean, maximum and minimum tem-
peratures, and total precipitation) from Chame, Jagat and
Larke stations were used. Climatic factors limiting regenera-
tion were identified from linear correlation analysis between
regeneration and climatic data.
3 Results
3.1 Treeline position and structural parameters
The species limit of A. spectabilis was recorded at 3984m
(GPS e-Trex) in Transect 1 (T1) and 3955 m in Transect 2
(T2) and its treeline at 3907 m in T1 and 3830m in T2. In the
case of B. utilis the treeline and species limit were recorded
at the same elevation in both transects, i.e. at 3996m in T1
and 4003m in T2.
Structural parameters (age, DBH, and basal area) revealed
that both of the species were more matured in T1 than
in T2 (Supplementary Table S2). The maximum age of
A. spectabilis and B. utilis was higher in T1 than in T2.
A. spectabilis tree density ranged from 50 to 280treesha−1.
The total basal area of B. utilis in both plots was higher than
A. spectabilis. The DBH distribution of A. spectabilis showed
bimodal distribution in T1 with peaks at 0–10cm and 35–
40cm DBH class (Supplementary Fig. S2). In T2, the DBH
distribution of the same species had an inverse-J-shaped dis-
tribution indicating continuous regeneration in the area. DBH
distribution of B. utilis in T1 presents a unimodal bell shape,
indicating poor regeneration in recent years. However, DBH
distribution of the same species in T2 indicates better recruit-
ment of individuals. Similar trends were observed in height
distribution. Age class distribution of the species was het-
erogeneous with an inverse-J-shaped and unimodal to multi-
modal bell-shaped distribution (Supplementary Fig. S3).
3.2 Age structure, regeneration and treeline dynamics
The demographic distribution of A. spectabilis and B. utilis
revealed the recruitment and mortality pattern over time
(Supplementary Fig. S3 and Fig. 4). The age distribution of
A. spectabilis indicated that the species was established in the
early 1850s in T1 (Fig. 4a) and in the 1950s in T2 (Fig. 4b).
The population of A. spectabilis was dominated by young in-
dividuals comprising 89% of the population below 50years
age. The recruitment of A. spectabilis was slow in the 1850s,
accelerating after 1950, and again after 1980. This could be
related to increased temperature in the area. The recruitment
of B. utilis started from the 1820s in T1, then reached T2 in
the 1840s (Fig. 4a, b). The proportion of young population
of B. utilis was low (13% of the population <50years old)
as compared to middle-aged trees (42% being 50–100years
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1282 N. P. Gaire et al.: Treeline dynamics with climate change at the Himalaya
old) and older ones (45% over 100years old). Recruitment
of B. utilis was lower at the beginning in both transects, but
increased slowly to peak in the 1880s in T1 and the 1930s
in T2. Regeneration oscillated between the 1880s and 1940s
and declined steadily since then.
The comparative age of the two tree species shows that
the regeneration of the B. utilis was higher before the recruit-
ment of A. spectabilis in the treeline community. Spatial–
temporal assessment of the upper species limit distribution
of A. spectabilis and B. utilis revealed that the position of
B. utilis was higher than the position of A. spectabilis in both
transects (Fig. 4a, b). Similarly, B. utilis colonized the area
earlier than A. spectabilis. Seedlings of the A. spectabilis
species were found about 80 m higher than trees indicat-
ing upward migration (Fig. 4c). Matured trees and young
seedlings were mostly dominant at lower elevation indicat-
ing stand densification as well. T2 had a lower number of
seedlings than T1. Seedlings of B. utilis were not recorded in
both transects, but some were observed just outside the plots.
On the basis of the temporal and spatial distribution of the
ages of B. utilis at an elevation gradient, we calculated that
the seedlings of the species were established at 3860m be-
tween the 1810s and 1820s and at 3990m during the 1890s
(Fig. 4a). The A. spectabilis on the other hand made a tree-
line community around 1850 at 3765m and reached 3907m
(present A. spectabilis treeline) during the 1950s. Seedlings
of this species are now established at 3984m, which is
close to the upper limit of B. utilis. The average upward
movement of the upper distribution limit of A. spectabilis
at the study sites was calculated to be 2.61myear−1(1.56–
3.66myear−1). The upward shifting of A. spectabilis was
more pronounced in T2 with migration rate of 3.66 myear−1,
while it was 1.56myear−1in T1.
The densities of saplings and seedlings of A. spectabilis
(Supplementary Table S3) indicated that its regeneration was
higher in T1 than in T2 and also higher than that of B. utilis
in both transects. As there was a presence of a large number
of saplings and seedlings of A. spectabilis in the site but no
seedlings of B. utilis, it is anticipated that the structure and
composition of this treeline community will change in the
future.
3.3 Tree-ring chronology
A 229-year-long (AD 1782–2010) standard tree ring chronol-
ogy of A. spectabilis was prepared using 46 cores from
29 trees (Fig. 5). The chronology revealed that there was no
constant increment in the growth of trees but it fluctuated
through time. The years 1818, 1819, 1974 and 1999 were
characterized by particularly poor growth, whereas the years
1789, 1814 and 2009 resulted in particularly wide rings. Re-
sults of the regime shift analysis suggest that there have been
constant changes in mean radial growth which are domi-
nated by short periods of above-average radial growth. Two
major periods with low radial growth were 36years start-
Figure 4. (a, b) Spatial and temporal variation in the recruitment
of tree species in T1 and T2. (c) Temporal upward shifting of Abies
spectabilis along an elevation gradient in the study site.
ing in 1854 and 63years starting in 1940. The period cen-
tred on 1905 is one of the shortest with below-average radial
growth. In the recent period (after 2000) the radial growth
is increased. Several statistics that were calculated for the
time span of AD1782–2010 and for the period of overlap
(AD1920–2005) of all tree-ring series indicated a high den-
drochronological potential (Supplementary Table S4). The
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N. P. Gaire et al.: Treeline dynamics with climate change at the Himalaya 1283
0
5
10
15
20
25
30
35
40
45
50
0
0.2
0.4
0.6
0.8
1
1.2
1.4
1.6
1827 1847 1867 1887 1907 1927 1947 1967 1987 2007
Number of radii
Ring width index
Year
Std chronology
Regime shift
Number of radii
EPS<85
%
Figure 5. Tree-ring standard chronology of Abies spectabilis from
the Kalchuman Lake area of Manaslu, central Nepal.
value of mean sensitivity and standard deviation is 0.136
and 0.18, respectively. The mean series correlation within
tree was high (0.467) as compared to the mean correlation
between tree (0.192) and among all radii (0.196). The EPS,
an indication of how well the site chronology estimates the
population chronology, was above (0.918) the threshold limit
of 0.85 (Wigley et al., 1984). The signal-to-noise ratio was
11.23 and the percentage of variance explained by the first
eigenvector was 24.4%.
3.4 Response of tree growth and regeneration to climate
change
The radial growth of the Abies spectabilis at the studied tree-
line was limited by the low temperature with a positive corre-
lation between the ring width chronology and monthly max-
imum temperature in most of the month and a negative re-
lationship with minimum temperature (Fig. 6a). The radial
growth in A. spectabilis correlated negatively with the mean
and minimum monthly temperature of June–September (r >
−0.45, P < 0.01) months of the current year (Fig. 6a).
However, above-average monthly mean and minimum tem-
perature in the previous year growing period (i.e. June–
September) influence negatively the current year growth.
Looking at the influence of seasonal climate on the growth,
effect of monsoon season (JJAS) temperature on the growth
was stronger than individual counterpart months with a sig-
nificant negative correlation with seasonal mean (r= −0.58,
P < 0.01) and minimum (r= −0.66, P < 0.01) tempera-
ture.
Though weak, radial growth was negatively correlated
with monthly precipitation of most months of the current
year (Fig. 6a). The relationship between ring width and pre-
cipitation of February of the current year was slightly nega-
tive (r= −0.38, P < 0.05). The precipitation during the pre-
vious year June correlated positively (r=0.45, P < 0.05)
with current year growth. The correlation between the radial
growth and seasonal sum of precipitation was weak and not
significant statistically.
-0.8
-0.7
-0.6
-0.5
-0.4
-0.3
-0.2
-0.1
0.0
0.1
0.2
0.3
0.4
0.5
0.6
pJ pJ pA pS pO pN pD J F M A M J J A S O Avg MAM JJAS ON DJF
Correlation coefficient
Month and season
Tmean Tmax Tmin Precipitation
(a)
-1.0
-0.8
-0.6
-0.4
-0.2
0.0
0.2
0.4
0.6
0.8
1.0
1.2
J F M A M J J A S O N D Avg
Correlation coefficient
Month
Tmean Tmax Tmin Precipitation
(b)
Figure 6. Relationship between the Abies spectabilis’ radial growth
with climate data (monthly mean, maximum, minimum, seasonal
average temperature, and monthly and seasonal sum of precipita-
tion) (a), and relationship between regeneration A. spectabilis with
monthly climate (b); the black horizontal line indicates a signifi-
cant correlation at the 95% confidence limit for a two-tailed test
and dashed horizontal line indicates a significant correlation at the
99% confidence limit for a two-tailed test. (a) shows the response
of radial growth to the monthly climate of June in the previous
year to October of the current year as well as seasonal averages.
pJ–pD signify June–December of the previous year; MAM, JJAS,
ON and DJF are the mean temperature and the sum of precipitation
of March–May, June–September, October–November and previous
year December to the current year February, respectively. For cli-
matic influence on the regeneration of the A. spectabilis a climatic
window of 12months of current years as well as annual average of
temperature or annual sum of precipitation are used.
Regeneration of the A. spectabilis was favoured posi-
tively by the above-average monthly maximum tempera-
ture during most of the months and above-average precipi-
tation during dry warm summer months (Fig. 6b). The rela-
tionship between regeneration and monthly maximum tem-
perature of the most of the months of the current year
was positive and statistically significant (r > 0.8, P < 0.05)
while the relationship was negative but significant with the
monthly minimum temperature of June–September of the
current year (r > −0.7, P < 0.05) (Fig. 6b). The relation
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1284 N. P. Gaire et al.: Treeline dynamics with climate change at the Himalaya
between regeneration and precipitation in August (r=0.96,
P < 0.01) was positive and statistically significant (Fig. 6b).
It is also evident from Fig. 6a and b that the growth and re-
generation of A. spectabilis is more sensitive to minimum or
maximum temperature rather than average temperature be-
cause correlation with these is more positive or negative than
average.
4 Discussion
4.1 Position, structure and dynamics of treeline
The position and dynamics of the treeline are the result of
the interaction of several factors including topoclimate (ra-
diation, temperature, precipitation, length of growing sea-
son, and snow cover), topography (slope inclination, relief
forms), ecology of tree species (regeneration, seed disper-
sal, successional stage), site history (climate oscillations,
fire, human impact, insect attacks), current biotic (browsing,
trampling, diseases and insect pests) and anthropogenic in-
fluences (burning, logging, grazing, recreation and tourism)
(Holtmeier, 2009). The position of the treeline, species line
and species composition varies across the globe as well as at
sites within the region (Körner, 1998; Miehe et al., 2007).
In Nepal, the position of the treeline varies between the
eastern, central and western region (Schickhoff, 2005). Re-
gardless of the plots, the A. spectabilis treeline in the present
study was found at 3907ma.s.l.and the B. utilis treeline
at 4003ma.s.l., which is comparable to the other studies
(Körner and Paulsen, 2004; Bhuju et al., 2010; Liang et
al., 2014). However, along the western slope of Mt An-
napurna, the upper timberline (B. utilis,R. campanulatum)
at north-facing slopes ascends to 4000–4100ma.s.l.and to
even 4400ma.s.l.on the Nilgiri northern slope (Schickhoff,
2005). Bhuju et al. (2010) found the treeline at 4050ma.s.l.
in Pangboche of Sagarmatha (Everest) region in eastern
Nepal, while at the Lauribina of Langtang in central Nepal
it was observed at about 3900ma.s.l.(Gaire et al., 2011).
Generally, the upper treeline elevations in the Hindu Kush–
Karakoram–Himalaya (HKH) region increases along two
gradients: a NW–SE gradient (corresponding to higher tem-
perature sums at the same elevations along the mountain
arc) and a peripheral–central gradient from the Himalayan
southern slope to the Great Himalayan range and the Tibetan
highlands (related to the combined effects of continentality
and mass elevation both leading to higher temperature sums)
(Schickhoff, 2005).
The size class distribution of the tree species reflects its
regeneration status (Lv and Zhang, 2012). In our study, the
structural parameters of the studied species not only var-
ied between the plots but also between the species. Com-
pared to T1, the smaller maximum DBH and younger age
of both the A. spectabilis and B. utilis in T2 indicated some
influence of the Kalchuman Lake in the seed dispersal in
the T2 from downside seed source. The DBH class dis-
tribution of A. spectabilis shows a bimodal distribution in
T1 with peaks in lower and intermediate DBH classes. The
peak in low DBH class indicates that regeneration in re-
cent years is good. The DBH distribution of B. utilis indi-
cated poor regeneration during recent years. The age dis-
tribution was an inverse-J-shaped to multimodal bell-shaped
with intra and inter-species differences. Such kind of differ-
ences in the age and DBH class distribution of A. spectabilis
and B. utilis have been observed in the other treeline eco-
tones of Nepal (Shrestha et al., 2007; Bhuju et al., 2010;
Suwal, 2010; Gaire et al., 2011), indicating site- and species-
specific regeneration condition in the Nepal Himalaya. Sim-
ilarly at the A. spectabilis treeline near the Everest region
on the Tibetan side, Lv and Zhang (2012) observed a mul-
timodal age distribution with peaks during 1840–1860 and
in recent years. The differential spatio-temporal regeneration
pattern reflected in the multimodal size (age and/or DBH)
class distributions were also observed in various treeline
sites of different mountains: for example Picea schrenkiana
in central Tianshan mountains, NW China (Wang et al.,
2006), Juniperus przewalskii in the Qilian Mountains, NE
Tibetan Plateau (Gou et al., 2012), Pinus uncinata in the
Pyrenees Mountains, NE Spain (Camarero and Gutiérrez,
2007; Batllori and Gutiérrez, 2008), Picea glauca in SW
Yukon, Canada (Danby and Hik, 2007), Larix gmelinii in
the Putorana Mountains, northern Siberia (Kirdyanov et al.,
2012), Picea glauca,Picea mariana and Larix laricina,
Abies lasiocarpa near Churchill, Manitoba, Canada (Mamet
and Kershaw, 2012).
Recruitment, critical determinant of the rate of forest or
treeline shift (Camarero and Gutiérrez, 2007), has been found
to be more sensitive to climate than adult mortality in harsh
environments where competition is low, because recruitment
has lower climatic thresholds than adult mortality (Lloyd,
1997). The recruitment of A. spectabilis was high during the
1940–1950s and after the 1980s which might have been facil-
itated by the warm winter temperature in the area and in the
country (Cook et al., 2003; Sano et al., 2005). In the present
study, the establishment of A. spectabilis was high in recent
decades as compared to the previous decades, which is con-
sistent to the findings of other studies in the treeline in the Hi-
malaya and other mountains (e.g. Gaire et al., 2011; Batllori
and Gutiérrez, 2008; Liang et al., 2011; Lv and Zhang, 2012).
Lv and Zhang (2012) found a significant tree recruitment in
the recent three decades and sporadic recruitment in earlier
periods AD1760–1960 in the treeline of the Tibetan side
of the Everest region. Liang et al. (2011) also found an in-
creased recruitment of Smith fir (Abies georgei) after the
1950s with an abrupt increase in the 1970s in the Tibetan
mountains. Batllori and Gutiérrez (2008) also observed past
and recent synchronous recruitment trends of Pinus uncinata
with climate change at the treelines in the Iberian eastern
range of the Pyrenees.
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N. P. Gaire et al.: Treeline dynamics with climate change at the Himalaya 1285
The regeneration at the treeline can be sporadic or episodic
(Cuevas, 2002; Lv and Zhang, 2012). Patchy spatial distri-
bution of A. spectabilis, with gaps in some diameter and
age classes observed in the present study, indicates episodic
regeneration. The dominance of multimodal age distribu-
tions also observed in other alpine or forest–tundra ecotone
studies suggested that recruitment in treeline forest ecosys-
tems is episodic or sporadic rather than gradual (Szeicz
and MacDonald, 1995; Cuevas, 2002; Batllori and Gutiér-
rez, 2008; Lv and Zhang, 2012). The spatial distribution of
B. utilis was more regular compared to A. spectabilis. In this
study, establishment of B. utilis in recent decades has been
very poor as compared to previous decades. A similar trend
has been reported from other treeline sites (e.g. Bhuju et al.,
2010). Recruitment of the species was slow in the beginning
in both transects and increased gradually to reach at peak
in the 1880s in T1 and the 1930s in T2 with a slight oscil-
lation between the 1880s and 1940s. Similarly, the regen-
eration of B. utilis before the arrival of A. spectabilis was
high. The maximum age of B. utilis was higher than Rhodo-
dendron campanulatum (Prabina Rana, personal communi-
cation, 2013) and the maximum age of R. campanulatum was
higher than that of A. spectabilis. Hence, this area might have
been colonized by shade intolerant B. utilis trees followed
by shade tolerant understory tree R. campanulatum and was
later invaded by A. spectabilis trees.
Seedling establishment is an important factor dictating the
altitudinal limits of treeline species (Hughes et al., 2009).
Evidently, treeline rise depends on seeds produced at the lo-
cal treeline rather than propagulae from more distant sources
at lower elevations (Kullman, 2007). At and above the tree-
line in the study site in the Kalchuman Lake area, we ob-
served neither long-living krummholz nor sub-fossil wood
of A. spectabilis and B. utilis. Matured and young seedlings
of A. spectabilis and matured Betula were mostly domi-
nant in the lower elevation. However, some seedlings of
A. spectabilis, probably due to global warming, have been
thriving at much higher elevation than tree individuals. This
study indicated both stand densification and upward migra-
tion as recorded in many other areas (Camarero and Gutiér-
rez, 2004; Danby and Hik, 2007; Gehrig-Fasel et al., 2007;
Kullman, 2007; Vittoz et al., 2008; Batllori and Gutiérrez,
2008; Kullman and Öberg, 2009; Kirdyanov et al., 2012).
Consistent with the observed trend in the other treelines
(Kullman, 2001, 2002; Wang et al., 2006; Kullman and
Öberg, 2009; Gou et al., 2012), spatio-temporal age distri-
bution showed that there was regeneration as well as upward
migration of the B. utilis until the end of the 1960s though
the exact rate is not calculated. The peak in the tree estab-
lishment in the past corresponds to the warm episode in both
winter and summer reconstructed temperature in the country
(Cook et al., 2003). However, we observed a stagnant upper
distribution limit or treeline of B. utilis in the recent decades
along with poor regeneration in spite of temperature warm-
ing in the area.
Average upward shifting of the upper distribution limit of
A. spectabilis at the treeline ecotone was about 2.61 myear−1
with some local variation in the area. This upward migra-
tion trend of A. spectabilis is consistent with the upward
migration (34m per decade) of A. spectabilis in the tree-
line of the Samagaun region of the MCA (Suwal, 2010),
and of Pinus wallichiana (19 and 14m per decade on south-
and north-facing slope) in the western Himalayas (Dubey et
al., 2003). Several other studies have reported treeline shift-
ing in different regions of the world (e.g. Camarero and
Gutiérrez, 2004; Danby and Hik, 2007; Gehrig-Fasel et al.,
2007; Harsch et al., 2009; Kullman 2001, 2002; Kullman and
Öberg, 2009; Chauchard et al., 2010; Leonelli et al., 2011;
Mamet and Kershaw, 2012; Kirdyanov et al., 2012). Kullman
and Öberg (2009) presented a regional-scale treeline rise of
Betula pubescens ssp. czerepanovii,Picea abies and Pinus
sylvestris in the southern Swedish Scandes of 70–90 m on av-
erage with maximum up-shifts of about 200m since around
AD 1915. Danby and Hik (2007) found an increased tree den-
sity as well as an advancement of Spruce (Picea glauca) tree-
line elevation by 65–85m on south-facing slopes in south-
west Yukon, Canada during the early to mid-20th century.
Similarly, Kirdyanov et al. (2012) observed an upslope shift
of the Larix gmelinii treeline position by approximately 30–
50m in altitude in the Putorana Mountains, northern Siberia
during the last century. However, Liang et al. (2011) found
no significant upward movement in fir treelines in the Ti-
betan Plateau despite the warming in the region in the past
200years.
4.2 Climatic factors affecting tree growth and
regeneration dynamics
Growth of a tree is associated with several abiotic fac-
tors including climate (Fritts, 1976). The radial growth of
A. spectabilis fluctuated over time with changing climate and
we did not observe constant increment or decrement in the
growth. However, a few studies have reported enhanced ra-
dial growth of the western Himalayan conifer during recent
years (e.g. Borgaonkar et al., 2011).
The radial growth of A. spectabilis in the treeline is more
responsive to temperature change. Tree growth was posi-
tively correlated with temperature (Tmean,Tmax and Tmin)
from October of the previous year to April of the current year
which indicates that temperature before the growing season
has a main influence on the radial growth during the subse-
quent growing period. High winter temperature may induce
early melting of snow with the easy availability of melt water
for growth in the growing season. Studies also have reported
that monthly and seasonal winter temperatures are more
limiting than growing-season temperatures to annual radial
growth in many upper treeline sites with a positive relation-
ship with winter temperature (Bräuning, 2004; Pederson et
al., 2004; Borgaonkar et al., 2011). Bräuning (2004) found a
strong positive relationship between the A. spectabilis ring
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1286 N. P. Gaire et al.: Treeline dynamics with climate change at the Himalaya
width chronology and temperature from November of the
previous year to January of the current year in western Nepal.
Borgaonkar et al. (2011) found a strong positive relation-
ship between the mean annual and winter (DJF) temperatures
of the concurrent year and growth of western Himalayan
conifers. The negative relationship, observed in the present
study, with the pre-monsoon (MAM) and monsoon season
(JJAS) temperature points towards some threshold temper-
ature or moisture stress because increase in temperature in
the pre-monsoon and monsoon season without adequate rain-
fall increases the evapotranspiration which leads to a soil-
moisture deficit and limits tree growth (Fritts, 1976; Cook et
al., 2003; Yadav et al., 2004). Other studies from the Nepal
Himalaya revealed that tree-ring width of A. spectabilis is
controlled by pre-monsoon (March–May) climate with neg-
ative correlation with temperature and positive correlation
with precipitation indicating that moisture availability in this
season limits tree growth (Cook et al., 2003; Sano et al.,
2005; Chhetri and Thapa, 2010; Gaire et al., 2011). Most
of these studies were carried out in the areas much lower
than treeline ecotone. Similarly, Yadav et al. (2004) ob-
tained a negative as well as weakened relationship between
the mean temperature of the summer months and growth of
A. spectabilis from treeline sites of the western Himalaya. In
the present study, responses of radial growth of A. spectabilis
to minimum and maximum temperatures in the current year
were in the opposite direction. This indicated to the preva-
lence of threshold temperature above or below which the
responses become less sensitive to temperature or nonlin-
ear to inverted U-shaped relationship (Paulsen et al., 2000;
D’Arrigo et al., 2004; Yadav et al., 2004; Kullman, 2007).
In this study, the radial growth of A. spectabilis was less
strongly correlated with precipitation, having significant neg-
ative correlation only with February precipitation. As pre-
cipitation in these months falls in the form of snow, high
snow accumulation delays the growth initiation, shorten-
ing the growth period and ultimately resulting in the for-
mation of a narrow ring. A deep snow pack in late win-
ter has been shown to effectively reduce radial growth rates
by maintaining low soil temperatures and delaying initiation
of cambial expansion (Fritts, 1976; Pederson et al., 2004).
Above-average moisture during the June of the previous year
positively affects the current year’s growth because above-
average moisture during summer and early autumn may pro-
mote storage of carbohydrates and bud formation, thus en-
hancing growth during the following year (D’Arrigo et al.,
2001; Fritts, 1976). The weak correlation of A. spectabilis ra-
dial growth with the precipitation also might be due to its sen-
sitivity to temperature compared to precipitation. Conversely
the weak correlation could be due to variation in the precip-
itation between the sampling sites and the local stations be-
cause in the Himalaya precipitation fluctuates greatly, even
within a small geographic area (Barros et al., 2004).
The relationship between regeneration of A. spectabilis
and the monthly maximum temperature of January–
December of the current year was positive while the rela-
tion was negative with the monthly minimum temperature of
the current year, indicating that low temperature adversely
affects seedling survival. Severe soil frosts during cold win-
ters were considered to be critical factors in the control of
seedling survival by causing needle and shoot desiccation
or fine-root mortality (Körner, 2003; Pederson et al., 2004;
Kullman, 2007). Since extremely low cool-season temper-
atures facilitate the formation of abrasive ice crystals that
physically damage trees and as a result often limit estab-
lishment at the treeline, this research also supports other re-
cent studies documenting the critical impact of warmer win-
ter temperatures on increased tree establishment in the high-
elevation ecotones (Kullman, 2007; Kullman and Öberg,
2009; Harsch et al., 2009). The regeneration of A. spectabilis
was positively correlated with the precipitation of May–
August months. The positive relationship with the precipi-
tation of these summer months implies that as the temper-
ature had already attained the minimum threshold required
for growth, high rainfall aids the survival and growth of
seedlings and saplings. During summer months the temper-
ature in the study area would often be high. So, low rainfall
may create a desiccation situation and will have a stronger
effect on recruitment because germinants are more sensitive
to drought stress (Hughes et al., 2009; Fajardo and McIn-
tire, 2012). Comparing with the past long-term reconstructed
climatic data (Cook et al., 2003), the regeneration of Abies
seems to be good during the episode of warm winter and cool
summer. Hence, temperature plays a crucial role in growth
and regeneration of A. spectabilis at the natural climatic tree-
line of the Himalaya.
From a similar study at the timberline on the Tibetan
side of the Everest region, Lv and Zhang (2012) found that
A. spectabilis recruitments in 5-year classes were positively
correlated with their corresponding monthly mean air tem-
peratures in June and September and with Palmer Drought
Severity Index in June. The relationship was inverse for re-
generation and ring width. From a study in a treeline on the
southeastern Tibetan Plateau, Liang et al. (2011) found a
significant positive correlation between the Smith fir (Abies
georgei var. smithii) recruitment and both summer and winter
temperatures. Wang et al. (2006) reported that several con-
secutive years of high minimum summer temperature and
spring precipitation was the main factors favouring the estab-
lishment of Picea schrenkiana following germination within
the treeline ecotone in the central Tianshan mountains.
Due to the lack of young seedlings and saplings as well
as long climatic data, we could not calculate the climatic
variables limiting growth and regeneration of B. utilis. How-
ever, in a recent study in birch timberlines from the Nepal
Himalaya including our study site, Liang et al. (2014) using
CRU grid-based data found a significant positive relationship
between tree-ring width chronologies of B. utilis and pre-
cipitation in May and the pre-monsoon (MAM) season and
a less strong negative relationship with temperature. They
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N. P. Gaire et al.: Treeline dynamics with climate change at the Himalaya 1287
concluded that Himalayan birch growth at the upper timber-
lines is persistently limited by moisture availability during
the pre-monsoon season. The poor regeneration and lack of
recent shifting of the B. utilis in the area might result from
the increasing moisture stress (Liang et al., 2014) as avail-
able precipitation data from the nearby station have shown a
decreasing trend in the precipitation. In addition to the influ-
ence of climate change, the lack of recent regeneration of the
B. utilis seedlings could be due to the influence of increased
canopy cover by its own and associated tree species as well
as dense shrub scrub because B. utilis seedlings could not es-
tablish under their own closed canopy even if they produce
viable seeds (Shrestha et al., 2007) because the birch seedling
growth is facilitated by direct sunlight (Shrestha et al., 2007;
Hughes et al., 2009).
4.3 Treeline dynamics with climate change
The relationships between regeneration, treeline shifts and
climate change may be more complex because climate may
affect tree recruitment and treeline advance rate in differ-
ent ways (Camarero and Gutiérrez, 2004; Wang et al., 2006;
Kirdyanov et al., 2012). A treeline ascent implies several
consecutive processes: production of viable seeds, disper-
sal, availability of adequate regeneration sites, germination,
seedling survival and persistence until the individual reaches
adulthood (Wang et al., 2006; Kullman, 2007). Climate vari-
ability affects all these sequential stages, but the same cli-
matic variable can enhance one of these processes while in-
hibiting another one (Camarero and Gutiérrez, 2004; Wang
et al., 2006). At a global scale, treelines are considered
to be constrained primarily by growing season temperature
(Körner and Paulsen, 2004). However, at many alpine tree-
line ecotones, both winter and summer temperatures are of-
ten key constraints on tree recruitment (Harsch et al., 2009;
Liang et al., 2011) including other local site conditions,
species’ traits and feedback effects (Danby and Hik, 2007;
Batllori and Gutiérrez, 2008). In the present alpine treeline
study the establishment of the A. spectabilis is also controlled
by both winter and summer climatic events.
An increasing number of studies have demonstrated that
tree population density at treelines can respond quickly to ris-
ing temperatures (Camarero and Gutiérrez, 2004; Kullman,
2007; Liang et al., 2011) compared to the changes in tree-
line position because of the great longevity and phenotypic
plasticity of tree individuals (MacDonald et al., 1998; Lloyd,
2005). If temperature is the primary and dominant driver
for both recruitment and growth, these processes should be
positively synchronized (Fajardo and McIntire, 2012). Some
previous studies at treelines found concurrent synchronies
(MacDonald et al., 1998; Batllori and Gutiérrez, 2008) and
lagged synchronies of tree growth and regeneration in both
positive and negative directions with climate change (Fajardo
and McIntire, 2012). We found a synchronous regenera-
tion of A. spectabilis in the treeline ecotone with climate
warming. The climatic conditions that enhance radial growth
of A. spectabilis were almost similar to the climatic con-
ditions that enhance regeneration with slight variation in
some months, which is similar to the findings of other stud-
ies (Szeicz and MacDonald, 1995; Camarero and Gutiérrez,
1999). In this study both the recruitment and radial growth of
A. spectabilis was found to be associated positively with win-
ter temperature and negatively with summer (May–August)
months’ mean and minimum temperature.
In spite of the regeneration of A. spectabilis above the ex-
isting treeline, the pace of future treeline shifting with cli-
mate change may not necessarily be the same because a
seedling takes many years from establishment to reach its
tree height (in the treeline, Abies took more than 30years
to reach 2 m height) and then to develop into a forest stand
(Lloyd, 2005). On the other hand, there are no seedlings of
B. utilis in and above the treeline. It will take several decades
for newly established Betula seedlings to develop and form a
treeline even if they establish now. As A. spectabilis in the
treeline is more responsive to temperature change, an ad-
vance of this at the natural treeline of the Himalaya with
climate change may continue if a long-term warming trend
stimulates growth frequently enough even in cooler years or
if low temperature events/periods which limit growth and re-
generation are insignificant or if there would not be water
deficit in plants which could offset the expected positive ef-
fects from temperature increase in tree establishment, growth
and the upslope advance of treeline (Paulsen et al., 2000;
Daniels and Veblen, 2004; Wang et al., 2006; Kullman and
Öberg, 2009). In the case of B. utilis, Liang et al. (2014) re-
ported that the birch treeline of the Himalaya is a rare case
of a drought-induced alpine timberline and Himalayan birch
at its upper distribution boundary is increasingly at risk of
survival. Therefore, downslope range shift may occur as a re-
sponse to global-change-type droughts (Liang et al., 2014).
With the supportive evidence of differential life history, re-
generation condition and the species-specific response of
these two treeline species to climate change, it is clear that, in
addition to treeline position, the community structure in the
studied treeline in the Himalaya is going to change, if current
climate change and response pattern continues.
5 Conclusions
The present study provided a recruitment pattern and dynam-
ics history of Himalayan fir and Mountain birch at the high-
altitude treeline of the Manaslu region, central Nepal Hi-
malaya. Although regeneration patterns varied between the
species, increasing trends of stand densification as well as
upward shifting of the studied treeline is evident. The up-
ward shift of A. spectabilis at MCA was estimated to be
2.61myear−1. In spite of upward migration of B. utilis up
to the mid-20th century, its upper distribution limit has been
stagnant in recent years. The regeneration of A. spectabilis
www.clim-past.net/10/1277/2014/ Clim. Past, 10, 1277–1290, 2014
1288 N. P. Gaire et al.: Treeline dynamics with climate change at the Himalaya
was positively related with monthly maximum temperature
in most of the months of the current year and precipitation in
May–August although the growth of the B. utilis can be also
limited by pre-monsoon precipitation (Liang et al., 2014).
Spatial and temporal variations in age structure and regen-
eration pattern of these two species and their species-specific
response to climate indicated that the plant communities at
the treeline ecotone in the Nepal Himalaya were sensitive to
climate change and the studied treeline is changing. Stud-
ies incorporating multiple species and covering other proxy
evidence like pollen from lake sediments could enhance our
understanding of spatio-temporal treeline and vegetation dy-
namics in association with climate change.
The Supplement related to this article is available online
at doi:10.5194/cp-10-1277-2014-supplement.
Acknowledgements. NAST provided a PhD fellowship to the first
author. We are grateful to the National Trust for Nature Conserva-
tion for the permission to conduct this study at MCA. We thank
Arbindra Shrestha, Janardan Mainali and local field assistants for
their help during field work. Amalava Bhattarcharyya, Moinuddin
Ahmed, Kumar Mainali and Madan Krishna Suwal provided with
valuable suggestions in improving the paper and plotting the fig-
ures. Russell E. Train Education for Nature Programme, WWFUS
provided a professional grant to the first author to participate in
dendrochronology training in China. APN and PAGES supported
participation in the PAGES Goa 2013 meetings. We are grateful to
the four anonymous reviewers, Thorsten Kiefer and Anne-Laure
Daniau for their constructive comments.
Edited by: A.-L. Daniau
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