Carbon storage in a high-altitude Polylepis

woodland in the Peruvian Andes

Ekatherina Vsquez, Brenton Ladd &

Nils Borchard

Alpine Botany
ISSN 1664-2201
Alp Botany
DOI 10.1007/s00035-014-0126-y

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Author's personal copy

Alp Botany
DOI 10.1007/s00035-014-0126-y

SHORT COMMUNICATION

Carbon storage in a high-altitude Polylepis woodland

in the Peruvian Andes
Ekatherina Vasquez Brenton Ladd
Nils Borchard

Received: 29 November 2013 / Accepted: 11 February 2014

Swiss Botanical Society 2014

Abstract Polylepis woodland occurs in Perus tropical

highlands at elevations between 3,500 and 5,000 m above
sea level and Polylepis is the most common tree at timberline in South America. The objective of this study was
to assess the total ecosystem carbon stock in a Polylepis
incana woodland, i.e., aboveground biomass (canopy trees
and understory), root biomass and soil carbon stocks were
all quantified. As part of this study, an allometric equation
for the quantification of the aboveground biomass of
individual P. incana trees was developed for the first time.
The most important carbon pool was the soil
(39.7 6.9 kg m-2) followed by the aboveground biomass of Polylepis trees (3.8 0.7 kg m-2). The total
ecosystem carbon stock was estimated to be
43.9 7.6 kg m-2; thus, 90.6 % of the ecosystem carbon
stock is soil carbon.
Keywords Polylepis  Carbon stock 
Ecosystem services  Allometric equation
E. Vasquez  B. Ladd (&)
Facultad de Ciencias Ambientales, Universidad Cientfica del
Sur, Panamericana Sur km19, Lima 33, Peru
e-mail: brenton.ladd@gmail.com
B. Ladd
Evolution and Ecology Research Centre, School of Biological,
Earth and Environmental Sciences, University of New South
Wales, Sydney, NSW 2052, Australia
N. Borchard
Agrosphere (IBG-3), Julich Research Centre, 52425 Julich,
Germany
N. Borchard
Departamento de Edafologa, Instituto de Geologa, Universidad
Nacional Autonoma de Mexico, Ciudad Universitaria,
04510 Mexico, DF, Mexico

Introduction
One promising option to reduce greenhouse gas emissions
to the atmosphere is to protect forests and woodlands,
which could preserve carbon stored in plant biomass and
soil (Guo and Gifford 2002; IPCC 2000; Lal 2005). Any
land-use change or deforestation risks loss of the stored
carbon due to biomass removal and losses of soil carbon
through enhanced decomposition or erosion (Gloor et al.
2012; Guo and Gifford 2002; IPCC 2000). In Peru, carbon inventory studies have been done mostly in the
Amazon (Gloor et al. 2012; Houghton 2012; Midgley
et al. 2010). In contrast, highland tropical ecosystems,
such as Polylepis woodland, that are common in the
Andes have received much less attention. The Polylepis
ecosystem is distributed along the length of the highlands
in the tropical and subtropical Andes at an elevation of
3,5005,000 m above sea level (Zutta et al. 2012).
Polylepis is the most prominent South American tree at
timberline (Beck et al. 2008).
Polylepis woodlands also contain a number of endemic species, and notable among these is a group of birds
specialized in the use of these habitats (Aucca et al.
2008; Servat et al. 2002). They are also important providers of water-related ecosystem services: they are a
critical component of the Andean headwaters (Aucca and
Ramsay 2005; Buytaert et al. 2006). Polylepis woodlands
intercept mist and fog which augments the supply of
potable water in Peru (Fjeldsa 2002). Many researchers
agree that the Polylepis woodlands act as water regulator
through their ability to store water in soil and slowly
release it, thus, reducing peak flows and contributing to
the availability of water in dry seasons (Fjeldsa 2002;
Fjeldsa and Kessler 1996; Kessler 2002, 2006). Therefore, we emphasize the importance of protecting the

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Alp Botany
Table 1 Summary statistics for the allometric equation which correlated tree aboveground biomass with tree height and trunk diameter
Source

df

SS

Pr [ F

g2

Height

252

16.3

0.005

62.5

2.7

0.14

10.6

Diameter

43

Error

108

Model: aboveground biomass = -16.51 ? 40.26 9 tree

(log10) ? 9.30 9 trunk diameter (log10)

height

SS sum of squares, df degrees of freedom, Pr probability

g2, percentage of variation of the R2 explained for each independent
variable

stored carbon in the following pools: aboveground biomass

(canopy trees), understory biomass, root biomass and soil.
Aboveground biomass (canopy trees)

Fig. 1 Location of the field-based carbon inventory: Nor Yauyos

Cochas National Park

Polylepis woodland ecosystem for water-related ecosystem services.

In order to estimate the benefits of conserving Polylepis
woodland in terms of carbon emissions avoided, we made a
comprehensive inventory of stored carbon in soil and biomass. We also developed an allometric equation for
Polylepis incana. Many allometric equations have been
generated for tropical and temperate tree species (Araujo
et al. 1999; Brown 1997; Nelson et al. 1999; Zianis et al.
2005), but none for this important highland species.

Materials and methods

Study area
We made a carbon inventory of a P. incana woodland in the
Nor Yauyos Cochas National Park, located in Peruvian
Andes, 320 km southeast of Lima, at 4,200 m above sea
level (Fig. 1). The height of mature trees in this woodland
ranged between 2 and 3 m. We measured the quantity of

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To calculate the aboveground biomass of the Polylepis

woodland we first developed an allometric equation for
P. incana trees. For ten randomly selected trees we measured
the height by climbing the trees (when required) with a tape
measure in hand. Trunk diameter at 50 cm aboveground
level (included bark) was also measured with a tape measure. Following measurements of tree height and trunk
diameter we harvested all ten trees. Trunk material and
foliage were cut into small pieces for further processing. Ten
plots of 10 9 10 m were then randomly selected, in which
we measured trunk diameter and height of all P. incana trees
therein and recorded the geographic coordinates of each
plot.
Carbon stored in biomass
For aboveground carbon, we started by establishing an
allometric equation for P. incana, based on data from the
ten harvested trees. We used XL-stat (Add-in Soft, Paris,
France) to perform a multiple regression analysis in which
aboveground biomass (AGB) was the dependent variable
and tree height (H) and trunk diameter (D) were the
independent variables (Table 1). Tree height and trunk
diameter were Log10 transformed prior to the regression
analysis because tree growth is known to decline with tree
age (Ladd et al. 2009). This multiple regression analysis
produced the following equation: AGB = -16.51 ?
40.26 9 H (log10) ? 9.30 9 D (log10) (Table 1), that in
turn was used to calculate the AGB of all P. incana trees
found in the ten 10 9 10 m plots (in total 132 trees).
The understory carbon stock was calculated from vegetation (grasses, herbs and shrubs) collected from sub-plots
(5 9 5 m) within each 10 9 10 m plot.
The estimates of root biomass were made for five plots
(plot 15; 10 9 10 m). We excavated root samples using

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stainless steel core sampler (15 cm long, 11 cm diameter)
at four sites in each plot, two of which were in open spaces
away from Polylepis trees and two directly adjacent to a
Polylepis tree. Soil was sieved and roots were washed and
dried using a temperature of 60 C as mentioned by
Cornelissen et al. (2003) for at least 72 h. Using the
stainless steel core sampler and a sledge hammer we were
able to extract both coarse and fine roots. In the studied
ecosystem we observed very little root biomass below
15 cm and, therefore, we limited root sampling to 15 cm
depth. Nevertheless, Polylepis trees often posses large tap
roots (lignotubers) and we did not sample these or the
coarse roots adjacent to the tap root which resulted in an
underestimation of tree biomass. In an effort to constrain
this error we excavated and harvested tap roots from 4 of
the 10 harvested trees. All biomass samples were dried to
constant weight at 60 C and converted to carbon by
assuming a 50 % carbon content in aboveground biomass
(Giese et al. 2003; Schlesinger and Bernard 2013) and a
45 % carbon content in roots (Gayoso and Schlegel
2001). All data were then expressed in kg C/m2 ground
area.
Soil carbon sampling

Measured Biomass Kg Dry Weight

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30

1:1
20

10

0
0

10

20

30

Predicted Biomass Kg Dry Weight

Fig. 2 The relationship between modeled vs. measured values of
biomass of the P. incana trees (y = y = x - 4E-14, R2 = 0.73). See
Table 1. The diagonal line indicates the 1:1 relationship

modeled values. The g2 values (Table 1) are a measure of the

percentage variation of the R2 of the regression model that is
explained by each variable. The g2 values confirmed that the
two independent variables contributed to the predictive
power to the regression model, but that tree height was the
most influential (g2 = 62.5 %): see Table 1 for further
detail.
Carbon stock in soil and biomass

Soil pits were excavated to bedrock (42 3.4 cm) in 5 of

the ten 10 9 10 m plots. Due to decreasing carbon content
in deeper soil layers (Jobbagy and Jackson 2000), samples
from two soil depths (topsoil: 010 cm; subsoil:
3040 cm) were collected from the face of the soil pits
using a metal tube of known volume. Thereafter, soil
samples were dried at 60 C and stored in a dry, dark place
until analysis of carbon by ignition at 350 C for 24 h.
Carbon content was calculated assuming that 58 % of the
mass lost by ignition was carbon (Scheffer and Schachtschabel 1998). Knowing soil bulk density (BD) and depth
of soil layers (020 cm and 20 to *40 cm) (Z), we applied
the following equation to calculate the soil carbon stock:

Applying the allometric equation to the 132 trees encountered in the ten 100 m2 plots, we obtained an average carbon
stock of 3.8 0.7 kg m-2 in the AGB of Polylepis trees.
The understory carbon stock was 0.1 0.0 kg m-2 and the
carbon stock of root biomass was 0.3 0.0 kg m-2. The
soil carbon pool was the largest pool with an average value
of 39.7 6.9 kg m-2. In total the investigated Polylepis
ecosystem plots stored in biomass and soil was
43.9 7.6 kg m-2. Yet, the soil was the most significant
sub-pool accounting for 90.6 % of the total ecosystem carbon stock.

soil carbon stock kg m2 carbon kg t1


 BD t m3  Z m

Discussion

Results
Allometric equation
A multiple regression with two variables (tree height and
trunk diameter at 50 cm) produced an allometric equation
that predicts the aboveground biomass with reasonable
accuracy (R2 = 0.73; Table 1). Inspection of Fig. 2 reveals
that most of the data adhere very closely to the 1:1 line
indicating close correspondence between the observed and

The allometric equation developed with the multiple

regression was sufficient to explain 73 % of the variance in
aboveground biomass of the individual trees, meaning that
27 % of the variance remained unexplained. Nevertheless,
this equation allows reasonable estimates of aboveground
biomass in P. incana trees and the raw data (Table 2) can be
the foundation for a larger dataset that may enable the
elaboration of a more accurate allometric equation. Further
if the data reported here is unified with data from other
species of Polylepis, it may be possible to develop a generalized allometric equation that could apply across the

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Table 2 Height, trunk diameter and weight of the Polylepis trees
harvested and then used to develop the allometric equation
No. of tree

Height (m)

Trunk
diameter
(cm)

Tap root
biomass
(kg)

Aboveground
biomass (kg)

2.94

54

18.2

2.00

34

9.8

3.00

12.3

4.18

40

23.7

3.40

42

1.95

19.3

2.70

14

4.22

11.2

1.96

16

7.0

2.18

31

10.5

3.47

34

3.91

18.6

10

1.98

28

1.86

8.8

Mean SE

2.78 0.76

30 14

3.99 0.62

13.9 5.5

entire Polylepis genus, as has been achieved for the Eucalyptus genus (Williams et al. 2005). We excavated and
weighed four tap roots of the ten harvested trees and we
found that tap root biomass was significant accounting for
15.89 2.67 % of the total biomass of the P. incana trees.
In future research, it may also be important to harvest the
tuberous tap roots that Polylepis trees often possess, as these
tap roots may allow a better estimate of carbon stock.
Another variable that could be measured in future studies to
improve the allometric equation is wood density of Polylepis
trees, which varies with altitude (Hoch and Korner 2005;
Moser et al. 2008).
Soil is the largest reservoir of organic carbon in the terrestrial biosphere (Schlesinger and Bernhardt 2013). In this
P. incana ecosystem the soil stock accounted for 90.6 % of
total carbon stock, which is comparable to the situation in
Perus alpine grasslands. In Perus high elevation Puna
grasslands in Manu National Park soil carbon accounts for
89.6 % of the total ecosystem carbon (Gibbon et al. 2010).
However, further north (closer to the equator) in an Ecuadorian Polylepis forest soil carbon accounted for only
1030 % of the ecosystem carbon stock (Fehse et al. 2002).
The fact that soil carbon accounts for such a large fraction of
total ecosystem carbon in the studied P. incana woodland
may result from the fact tree biomass is extremely low
(3.8 0.7 kg m-2) in comparison with other tropical
Andean woodlands (up to 175 t ha-1; see Moser et al. 2008
and Fehse et al. 2002). An alternative explanation for the
relatively abundant soil carbon in the P. incana woodland
relative to the Ecuadorian woodlands is that temperatures
are lower because the P. incana woodland is located further
south. Lower temperatures may limit decomposition leading
to accumulation of soil carbon.

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Despite the recognized ecological values of Polylepis

woodland (Aucca et al. 2008; Servat et al. 2002) little
quantitative work on ecosystem services has been conducted
for this ecosystem type. The ability to store carbon is an
important ecosystem service which will be diminished if
deforestation occurs (Guo and Gifford 2002). Protected
areas can contribute to the conservation of these substantial
carbon stores. In Brazil the expansion of national parks has
led to a reduced rate of deforestation (Nepstad et al. 2006;
see also Scharlemann et al. 2010). Analysis of the National
Forest inventory in Peru indicates that 71,755 ha of Polylepis woodland exists, of which only 12.9 % (9,263 ha) is
currently protected (MINAM 2012). Following the Brazilian example, the addition of Polylepis woodland to the
national parks in Peru is a potential strategy that could
protect the significant carbon stocks stored in Polylepis
woodlands.
Acknowledgments We thank J. Takahashi and the University
Cientfica del Sur for financial support of the study. We also thank the
reas Naturales Protegidas (SERNANP) for
Servicio Nacional de A
permission to destructively harvest Polylepis trees, and especially we
thank the SERNANP staff (Che et al.) that work in Yauyos Cochas
National Park for their help during the field campaign.

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