An Allometric Model Based Approach For Estimating

Biomass In Seven Indian Bamboo Species In Western

Himalayan Foothills, India
R. Kaushal
ICAR-Indian Institute of Soil and Water Conservation
S. Islam (  [email protected] )
ICAR-Indian Institute of Soil and Water Conservation
Salil Tewari
G.B. Pant University of Agriculture & Technology
J. M.S. Tomar
ICAR-Indian Institute of Soil and Water Conservation
S. Thapliyal
ICAR-Indian Institute of Soil and Water Conservation
M. Madhu
ICAR-Indian Institute of Soil and Water Conservation
T. L. Trinh
International Bamboo and Rattan Organization
J. Durai
International Bamboo and Rattan Organization

Research Article

Keywords: Bamboo, Biomass, Diameter, Height, Monte Carlo cross-validation, power function, Synthetic bamboo height

Posted Date: December 8th, 2021

DOI: https://doi.org/10.21203/rs.3.rs-1106649/v1

License:   This work is licensed under a Creative Commons Attribution 4.0 International License. Read Full License

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Abstract
The rapid growth rate, high biomass production, and annual harvesting, makes bamboo as suitable species for
commercial production. Allometric equations for many broadleaf and conifer tree species are available. However,
knowledge on biomass production and allometric equations of bamboos are limited. This study aims at developing
species specific allometric models for predicting biomass and synthetic height values as a proxy variable for seven
bamboo species in Himalayan foothills. Two power form based allometric models were used to predict above ground
and culm biomass using Diameter at breast height (D) alone and D in combination with culm height (H) as independent
variable. This study also extended to establishing H-D allometric model that can be used to generate synthetic H values
as proxy to missing H. In the seven bamboo species studied, among three major biomass component (culm, branch and
foliage), culm is the most important component with highest share (69.56 to 78.71%).Distribution of percentage (%)
share of culm, branch and foliage to above ground fresh weight varies significantly between different bamboo species.
D. hamiltonii has highest productivity for above ground biomass components. Ratio of dry to fresh weight of seven
bamboo species was estimated for culm, branch, foliage and above ground biomass to convert fresh weight to dry
weight.

1. Introduction
Bamboos are a group of perennial plants belonging to Poaceae family and subfamily Bambusoideae. A total of 1642
bamboo species occupy a wide ecological regime across the globe, mostly in tropical and sub-tropical regions1. The
rapid growth rate, high biomass production, and annual harvesting, makes bamboo as suitable species for commercial
production. Being an annual yielding crop with a wide range of product possibilities such as construction materials,
woven products, pulp and paper, food, feed and fodder, scaffolding, bio-energy, engineered products, lifestyle products,
medicine among others, offers itself to contribute to poverty reduction, job creation and green economy development.
Moreover, bamboo has proven wide range of ecosystem services especially soil erosion control, water recharge, slope
stabilisation, climate change, wildlife habitat and biodiversity conservation. Overall, bamboo contributes in achieving a
number of United Nations 2030 Agenda Sustainable Development Goals – particularly, SDG1, SDG7, SDG 11, SDG 12,
SDG 13, SDG 15, and SDG 172.

Worldwide, bamboo grows over in 35 million ha area and covers 3.2 percent of the forest areas of their host countries,
or about 1 percent of the global forest area 3,4 . In India, bamboo grows naturally throughout the country except in
Kashmir region. India is one of the richest countries of the world in terms of bamboo genetic resources with 125
bamboo species belonging to 23 genera. In term of coverage, India has largest bamboo cover, the bamboo area in the
country is 15.68 million hectare which provides livelihood to about 2 million traditional artisans through harvesting,
processing, value addition and selling bamboo products 5 . The demand of bamboo is estimated to be 26.69 million
tonnes as against supply of 13.47 million tonnes in the country 6 . Recognizing the importance of bamboo, Government
of India (GOI) allocated an outlay of Rs.12.90 billion ($177.6 million) for promoting bamboo sector and has set a target
to bring about 0.1 million ha area under bamboo plantations in forests, degraded lands and farmlands for generating
livelihood opportunities and meeting the industrial demand 7 .

Bamboo cuts across various sector: agriculture, forestry, manufacturing, construction, and energy. Particularly,
agriculture, forestry and land use contributes to about one-fifth of the GHG emissions. India has set its target to cut its
carbon emissions to net zero by 2070 at the United Nations Climate Change Conference (COP26) climate crisis summit
2021 organized in Glasgow, UK for which monitoring and measurement of tree biomass assumes a greater importance.
Assessment of biomass is helpful in timber extraction, tracking changes in the carbon stocks and quantifying the
amount of carbon dioxide which can be sequestered from the atmosphere. Destructive methods and indirect methods

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(biomass equations) are generally used for estimating tree biomass. Use of biomass equations is cost effective and
less time consuming and is recognized by United Nation as a common framework and good practice guidance for
carbon reporting in the implementation of the emerging carbon credit market mechanism8. Biomass equation predicts
biomass based on easily correlated measured variables like diameter at breast height and tree height9–10. However, in
bamboos biomass estimation becomes more complicated due to presence of different aged and multiple culms. In
bamboos, so far, occular methods are being used for assessing biomass by the traders and farmers which require long
field experience and suffer from high degree of inaccuracy and therefore results in huge losses to the farmers, foresters,
industries and other bamboo growers. Therefore, there is an urgent need to develop site specific predictive models/
equations for estimating biomass in bamboos. Of various non linear models, allometric models have been found to be
most efficient forestimating biomass because of numerous computational advantages like, high convergence of model
parameters compared to most of the other non-linear models, best goodness of fit and the low statistical errors11–14.

In literature, allometric equations are available for broadleaf and conifer tree species. However, knowledge on biomass
production in bamboos is limited. Most of allometric equations for bamboo are available for Phyllostachys species in
China15–18. In India, despite of large area under bamboo, allometric equations are available for few species viz.,
Bambusa cacharensis, Bambusa balcooa, Bambusa tulda, Bambusa vulgaris, Dendrocalamus hamiltonii,
Dendrocalamus longispathus, Melocanna baccifera, Schizostachyum dullooa and Pseudostachyum polymorphum in
north east India19–20; D. strictus in north India21–22; Bambusa bambos in eastern India23 and South India24. In north
India, not much work has been attempted on biomass estimation in bamboos. The present study was therefore
undertaken in seven commercial important bamboo species with the objectives: (1) to build species specific allometric
models for aboveground biomass and culm biomass using diameter (D), height (H), D 2H (2) to predict synthetic height
values as a proxy variable using diameter for biomass estimation.

2. Materials And Methods

2.1 Study area
The study was carried out to develop allometric models for predicting biomass in seven commercial important bamboo
species viz., Bambusa balcooa, Bambusa bambos, Bambusa vulgaris, Bambusa nutans, Dendrocalamus hamiltonii,
Dendrocalamus stocksii and Dendrocalamus strictus in the western Himalayan foothills from 2017-2020. All the
selected bamboo species are of commercial importance and on priority list of National Bamboo Mission, Government
of India and International Bamboo and Rattan Organization (INBAR). Two sites located at distance of 275 Km were
selected for collecting the data of seven different bamboo species (Figure 1). The details of the study sites are given in
Table 1. The planting material for Site 1 was collected from nursery of Uttarakhand Bamboo and Fiber Development
Board, Dehradun, India. The planting material for Site 2 was collected from different bamboo nurseries from different
parts of the country. The formal identification of the planting material was done based on morphological features
described by Seethalakshmi and Kumar (1998).

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 Table 1
Details of the study sites
Parameters Site1 Site 2

Location Research farm, ICAR-Indian Institute of Agroforestry Research Centre, G.B. Pant
Soil and Water Conservation, Dhulkot and University of Agriculture and Technology,
Selaqui Pantnagar

District/state/country Dehradun, Uttarakhand, India U.S. Nagar, Uttarakhand, India

Coordinates 30"20'59"N latitude, 77"53'05"E longitude 29°1′ 17″ N latitude and 79°29′ 14″ E
longitude

Altitude 548 m 243.8

Soil type Inceptisols Mollisols

Climate Sub-humid and humid sub-tropical

subtropical climate

Rainfall (mm) 1660 1364

Species Bambusa balcooa, B. bambos, B. vulgaris, Bambusa balcooa, Bambusa bambos,

B. nutans, Dendrocalamus hamiltonii, D. Bambusa nutans, Dendrocalamus hamiltonii,
stocksii and D. strictus and Dendrocalamus strictus

Year of plantation 2010 and 2 012 2005

Area of plantation 1.2 ha 3 ha

2.2 Sampling
For sampling, three plots of 0.018 ha (15m×12m) were selected for each species at site 1. For site 2, three plots of
0.0225 ha (15m×15m) were selected for each species. Each plot consisted of 9 bamboo clumps comprising of
numerous bamboo culms (poles). Two-stage random sampling design was adopted for selecting the clumps (1st stage
sampling unit) and culms (2nd stage sampling unit ). In total 39 plots (21 at site 1 and 18 at site 2) comprising of 351
clumps were demarcated to collect data every year from the year 2017 to 2020. For understanding the culm size
distributions within the sample plots, three clumps were selected randomly and measured for its clump girth, crown
diameter, culm number, culm diameter at breast height. Depending on the culm sizes, different diameter classes were
recognized representing the whole diameter range and from each diameter class three mature culms (> 3 year old) were
harvested randomly for each species in each clump. The age of mature culms in the clumps was determined on the
basis of culm sheath, colour of culms, position of the culms, and growth and development of branches and
leaves25.The summary statistics for biometric parameters of different bamboo species studied is given in Table 1.The
diameter of each bamboo culm was measured at breast height with the help of caliper. Culm height was measured with
the help of tape after harvesting the culm. The harvested culms were separated into different components viz., leaf,
branch and culm. Fresh weights were taken for respective components in the field. Subsamples from upper, middle, and
lower portion of the culms of different ages were oven-dried at 65°C to constant weight for determining the ratio of dry
weight to the fresh weight26.

2.3. Development of allometric equations for biomass estimation

Two power equation forms of allometric equations were developed for biomass modelling of seven bamboo species.
The first one was two-parameter general power form allometric equation as:

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where ‘a’ denotes the constant of proportionality or scale constant, ‘b’ denotes the power or shape constant, andεi is the
2 2
error component follow normal distribution with zero mean and non-constant variance σ i i.e.,εi ∼ iid N(0, σ i ). ‘x’ be a
single predictor variable. Here, diameter at breast height (D) and culm volume (D 2H) were used as x variable. Further,
three parameter power form of allometric equationwas used as:

where ‘a’ denotes the scale constant, ‘b’ and ‘c’ denote the shape constant for the predictor variable x and z, respectively.
εi be the error component follow normal distribution with zero mean and non-constant variance σ 2i i.e.,
εi ∼ iid N(0, σ 2i). For biomass modelling D was used as x and Hused as z.
The error term of both the allometric forms already discussed, assuming as heteroscedastic (i.e., not constant variance).
Therefore, a weighted fit model were used to calculate the variance of the residual variation using the function,
2 2 2
σ i = Var(εi) = σ̂ (v i) 2δ;where σ̂ is the sum of the estimated errors of the squares; v i is the weighting variable
associated with the ith plant/clump sampled; and δis the coefficient of the variance function to be estimated27−29.
Therefore, the weighted maximum likelihood nonlinear fixed effects modelling method was used to fit the biomass
equations for the seven bamboo species using the ‘nlme’ package of R Studio statistical software (R version 4.1.0). The
details of the allometric form of equations used for testing and validation of best fit are shown in Table 2. Statistical
differences between the biomass components of the seven species were examined using analysis of variance and
further post-hoc analysis was performed through Tukey’s Honest Significant Difference (HSD) test to achieve pair-wise
comparison between species biomass components.

Table 2
Allometric models selected for comparison
Model Id. Model name Model Expression

A1 Allometric1 y = aD b + ε

A2 Allometric2 y = a(D 2H) b + ε

A3 Allometric3 y = aD bH c + ε

Note: y denotes above ground fresh weight (kg), culm fresh weight (kg)

2.3.1 Model fit statistics and plot analysis to compare and choose the best model

Three popularly used model fit statistics viz. Akaike Information Criteria30, Bayesian Information Criteria31, and adjusted
R2 (Adj.R2) were used to evaluate, compare and select the best allometric equations as per the following formulae:

(i) AIC=-2ln (L)+2p, where L is the likelihood of the model, and p is the number of the parameters of the model and n is
the sample size.
(ii) BIC=-2ln(L)+pln (n), where L is the likelihood of the model, and p is the number of the parameters of the model and n
is the sample size.

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 RSS / ( n − p )
(iii) Adj. R 2 = 1 − TSS / ( n − 1 ) , where RSS=Residual sum of square, TSS=Total sum of square, and p is the number
of the parameters of the model and n is the sample size.
Model with highest Adj.R2 and lowest AIC, lowest BIC value was chosen as the best model31. Diagnostic plots of
observed values versus fit and residual trends were also used to assess model performance.

2.4 Model validation

Monte Carlo cross-validation method was applied to select, evaluate and compare the models32–35. The method divide
the dataset randomly into two parts, with about 80% for model development and 20% for cross-validation, repeated R
times (about 100 times) to obtain stable statistics. Model fitting statistics and cross-validation criteria were calculated
for each data series randomly selected and averaged over R (approximately 100) series. To evaluate the predictive
reliability of the fitted model, three statistical validation criteria viz., bias (percentage), Root Mean Squared Error (RMSE)
and Mean Absolute Percentage Error (MAPE) were used27–29, 34–38 as per the following formulae:.

wherey i= Observed value and ŷ i=Predicated value, R=Number of times cross-validation occured. A model that has the
lowest statistical error value in terms of bias, RMSE and MAPE was preferred as the best model.
2.5 Assessment of the relationship between height and diameter of bamboo

Allometric equation of the form H=𝑎𝐷𝑏+e, was used to develop relationship between height (H) in meter with diameter
at breast height(D) in cm, where ‘a’ is the normalization (scale) constant, ‘b’ is the shape parameter and e is error term
2 2
follow normal distributionwith zero mean and non-constant variance σ Hi i.e.,ei ∼ iid N(0, σ Hi). The model
development was done using the weighted maximum likelihood nonlinear fixed effects modelling methodand validation
was done using Monte Carlo cross-validation method.

3. Results
3.1 Biometric parameters of the seven bamboo species
Analysis of variance revealed that biometric parameters of the seven bamboo species viz., culm diameter, culm height,
culm fresh weight, foliage fresh weight and fresh above ground biomass weight varied significantly (p<0.05). The post-
hoc analysis through Tukey’s honest significant difference (HSD) test, revealed that D. hamiltonii has highest average
culm diameter (5.99 cm), culm height (11.91m), culm fresh weight (16.92kg), foliage fresh weight (1.53kg) and fresh
weight (20.39 kg). Average branch weight of D. hamiltonii though observed as highest (1.93kg) but not significantly
differed from B. balcooa, B. bambos, B. nutans and B. vulgaris (Table 3). Further, Tukey’s HSD test, revealed that ratio of
dry weight to fresh weight of different above ground biomass components viz., culm, branch, foliage and total above
ground biomass (AGB) significantly differed for seven bamboo species. The highest dry to fresh weight ratio for culm
(0.63), branch (0.62) and AGB (0.61) was observed in B. nutans. Whereas highest (0.53) dry to fresh weight ratio for
foliage was found in D. strictus (Table 4). Biomass distribution for the different components of the seven species
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revealed that the culm component shares the highest proportion in percentage (69.56-78.71%), followed by branch
(12.04-17.79%) and the foliage (3.50-16.94%). The allocation of branch biomass was highest (17.79%) in B. bambos. In
D. stocksii highest allocation of culm (78.71%) and leaf biomass (16.94%) were found among the seven bamboo
species (Figure 2).

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 Table 3
Summary statistics for different above ground biometric parameters of seven bamboo species grown in India.
Species Summary Culm Culm height Culm Branch Foliage Above
Statistics diameter (m) weight (kg) weight weight (kg) ground
(sample (cm) (kg) weight (kg)
size)

B. mean± 5.18±0.14bc 8.85±0.33c 9.62±0.68b 1.85±0.12a 1.29±0.09bc 12.76±0.84b

balcooa SE

(97) Min. 2.55 3.30 0.70 0.10 0.05 1.03

Max. 8.6 17.2 26.00 4.86 2.99 32.88

B. mean± 5.37±0.30bc 8.54±0.55c 10.51±1.40b 1.85±0.15a 0.51±0.08d 12.87±1.65b

bambos SE

(54) Min. 2.17 0.90 1.15 0.04 0.05 1.46

Max. 10.51 17.7 41.5 4.56 2.23 48.03

B. nutans mean± 4.74±0.16cd 10.43±0.41b 9.84±0.88b 2.05±0.28a 1.12±0.06c 13.00±1.14b

SE
(69)
Min. 2.23 3.80 0.56 0.05 0.12 0.86

Max. 8.60 17.30 32.94 12.40 2.10 45.15

B. mean± 5.56±0.18ab 9.39±0.46bc 12.41±0.95b 1.80±0.17a 1.51±0.11ab 15.71±1.16b

vulgaris SE

(62) Min. 2.77 2.7 0.9 0.08 0.04 1.15

Max. 8.60 18.80 31.58 5.895 3.545 38.34

D. mean± 5.99±0.19a 11.91±0.43a 16.92±1.17a 1.93±0.07a 1.53±0.07a 20.39±1.29a

hamiltonii SE

(121) Min. 1.91 2.80 0.49 0.29 0.10 1.05

Max. 9.87 22.78 45.20 3.90 3.69 51.27

D. stocksii mean± 4.24±0.10de 8.61±0.28c 4.97±0.24c 0.95±0.06b 1.20±0.06c 7.12±0.34c

SE
(83)
Min. 1.91 1.60 0.72 0.10 0.05 1.20

Max. 5.73 13.30 10.15 2.25 3.20 14.60

D. strictus mean± 3.78±0.11e 6.24±0.21d 3.96±0.33c 0.84±0.08b 0.59±0.03d 5.38±0.43c

SE
(176)
Min. 1.30 1.50 0.06 0.03 0.03 0.17

Max. 8.60 13.5 28.27 6.73 1.95 34.17

Note: SE: Standard Error of mean, Min.: Minimum value and Max.: Maximum value; mean± SE value within
biometric parameter, different superscript letters indicate significant differences (p<0.05) between the species.

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 Table 4
Description about ratio of dry to fresh weight of different above ground biomass components for seven bamboo
species.
Species Ratio=Wsample dry weight/ Wsample wet weight

Culm Branch Foliage AGB

B. balcooa 0.59b 0.60b 0.48d 0.58b

B. bambos 0.52e 0.49e 0.51c 0.51f

B. nutans 0.63a 0.62a 0.46e 0.61a

B. vulgaris 0.51f 0.57c 0.52b 0.52e

D. hamiltonii 0.47g 0.49e 0.46e 0.47g

D. stocksii 0.56c 0.45f 0.45f 0.53d

D. strictus 0.53d 0.53d 0.53a 0.53c

Note: Dry weight (kg) = Ratio × Fresh weight (kg). Within biometric parameter, different superscript letters indicate
significant differences (p<0.05) between the species.

3.2 Development and validation of allometric models

3.2.1Above ground biomass prediction

Three allometric equations [allometric form (1) in A1 and A2; allometric form (2) in A3] were developed for fresh above
ground biomass (AGB) and fresh weight for the seven bamboo species using the data collected from harvested culms
from >3 years old bamboo plantation. Further, allometric model A1 were again developed for bamboo height prediction
(H) using diameter at breast height (H). Results of allometric models development revealed ranged value of adjusted-R2
(Adj.R2) (0.82-0.97), AIC (190.24-562.79) and BIC (205.57-580.74), suggested differential fitting accuracy, whereas high
Adj.R2 and low AIC and BIC value for the three model, evidenced appropriateness of using allometric models (Table 5).

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 Table 5
Parameter estimates of allometric models fitted on 80% dataset of seven bamboo species for above ground fresh
weight prediction.
Species Model Weight variable Parameter Adj.R2 AIC BIC

Id. [Estimate± std. error]

a b c

B. balcooa A1 1/D 2.41 0.10±0.02 2.87±0.11 - 0.85 375.02 391.42

A2 1/D 0.40 0.15±0.03 0.79±0.04 - 0.83 387.65 404.05

A3 1/D 2.18 0.10±0.02 2.45±0.19 0.31±0.14 0.87 372.10 390.88

B. bambos A1 1/D 1.10 0.35±0.06 2.04±0.09 - 0.94 209.02 221.35

A2 1/D 0.29 0.24±0.06 0.69±0.04 - 0.92 225.08 237.40

A3 1/D 1.16 0.40±0.06 2.34±0.24 -0.29±0.20 0.94 215.00 234.37

B. nutans A1 1/D 2.02 0.21±0.04 2.56±0.11 - 0.82 269.03 283.08

A2 1/D 0.34 0.15±0.04 0.80±0.05 - 0.85 306.83 320.89

A3 1/D 1.99 0.16±0.03 2.34±0.17 0.27±0.14 0.85 273.51 295.59

D. hamiltonii A1 1/D 1.58 0.29±0.05 2.29±0.08 - 0.87 562.79 580.74

A2 1/D 0.58 0.16±0.03 0.77±0.03 - 0.88 552.35 570.30

A3 1/D 1.54 0.20±0.03 1.88±0.13 0.43±0.12 0.89 550.55 564.76

D. stocksii A1 1/D 1.48 0.37±0.05 2.01±0.9 - 0.86 208.4 223.72

A2 1/D 0.23 0.30±0.04 0.62±0.02 - 0.91 190.24 205.57

A3 1/D 0.78 0.29±0.04 1.31±0.16 0.58±0.11 0.90 198.21 222.30

D .strictus A1 1/D 1.78 0.15±0.01 2.48±0.04 - 0.96 357.64 378.24

A2 1/D 0.48 0.10±0.01 0.82±0.01 - 0.97 347.55 368.14

A3 1/D 1.75 0.11±0.01 1.92±0.1 0.57±0.09 0.97 335.17 367.52

B. vulgaris A1 1/D 0.95 0.30±0.08 2.23±0.15 - 0.84 275.31 288.56

A2 1/D 0.32 0.27±0.07 0.70±0.04 - 0.87 264.20 277.45

A3 1/D 1.20 0.26±0.06 1.50±0.21 0.63±0.14 0.87 268.31 289.12

Monte Carlo cross-validation method unfolded prediction accuracy of the fitted allometric models with the extended
value of ABias (-14.62 to 8.90%), ARMSE (0.98 to 5.12) and AMAPE (11.28 to 30.29%), revealed differential prediction
accuracy of the fitted model, whereas low error valuesfor the three model, indicating their prediction capabilities (Table
6).

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 Table 6
Validation of allometric models of above ground fresh weight
prediction on 20% dataset of seven bamboo species.
Species Model Id. ABias (%) ARMSE AMAPE (%)

B. balcooa A1 -7.13 3.09 21.67

A2 -11.52 2.87 24.85

A3 -7.10 2.97 21.22

B. bambos A1 -2.74 4.02 25.88

A2 2.77 4.17 27.95

A3 -14.62 5.12 30.29

B. nutans A1 0.50 2.14 10.83

A2 -8.87 2.22 13.96

A3 -1.64 2.02 11.28

D. hamiltonii A1 -9.13 3.90 23.93

A2 -2.57 3.71 19.59

A3 -1.87 3.24 18.96

D. stocksii A1 -8.11 1.62 24.44

A2 -4.08 1.31 17.66

A3 -4.98 1.33 18.68

D. strictus A1 -3.98 0.98 24.22

A2 -0.65 1.12 27.28

A3 -0.47 1.03 23.45

B. vulgaris A1 5.68 4.94 30.80

A2 8.46 3.91 22.50

A3 8.90 4.03 23.05

Allometric model A1 unveiled highest adj.R2 (0.94) for B. bambos, lowest AIC (209.02-269.03) and BIC (221.35-283.08)
for B. bambos and B. nutans. Further, it unveiled lowest prediction error with lowest ABias (-2.74 to 0.50%) ARMSE (2.14
to 4.02) and AMAPE (10.83 to 25.88%) for B. bambos and B. nutans. A2 model revealed lowest prediction error for D.
stocksii and B. vulgaris with lowest AMAPE (17.66 to 22.50%) and ARMSE (1.31 to 3.91). Moreover, A2 found lowest
ABias (-4.08%) for B. vulgaris. Model A3 has unveiled highest adj.R2 (0.87 to 0.97), lowest AIC (335.17-550.55) and BIC
(367.52-564.76) for B. balcooa, D. hamiltonii and D. strictus. Further, it achieved lowest prediction error for B. balcooa, D.
hamiltonii and D. strictus with lowest ABias (-7.13 to -0.47%) and AMAPE (18.96 to 23.45%) for B. balcooa, D. hamiltonii
and lowest ARMSE (2.97 to 23.45) found for B. balcooa and D. hamiltonii. Overall results of above ground biomass
modelling evidenced that A1 allometric model was most efficient for B. bambos and B.nutans, whereas A2 was most
reliable for D. stocksii and B. vulgaris, further A3 was most suitable for B. balcooa, D. hamiltonii, and D. strictus (Table 5,
Table 6). Figure 3 and Figure 4 demonstrates the results of best fitted allometric models for above ground fresh weight
prediction. The results show that the weighted best fitted allometric models capture heteroscedasticity well in residuals

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and produced much lower residuals. Further, residuals found to be random (lack of any pattern) visually which indicate
high fitting accuracy.

3.2.2 Culm biomass prediction

Results of allometric models development for fresh culm biomass extended value of Adj.R2 (0.80-0.96), AIC (200.39-
554.83) and BIC (205.72-578.54) lower than fitted model for fresh above ground biomass, suggested higher consistency
of the allometric models (Table 7). Cross-validation results of the fitted allometric models unfolded low prediction error
with extended value of ABias (-11.68 to 7.03%), ARMSE (0.79 to 4.70) and AMAPE (11.10 to 30.10%), revealed high
prediction accuracy of the the three model (Table 8).

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 Table 7
Parameter estimates of allometric models of culm fresh weight fitted on 80% dataset of seven bamboo species.
Species Model Weight variable Parameter Adj.R2 AIC BIC

Id [Estimate± std. error]

a b c

B. balcooa A1 1/D 2.91 0.06±0.01 3.00±0.10 - 0.81 324.18 340.59

A2 1/D 0.61 0.07±0.13 0.87±0.03 - 0.89 311.20 337.61

A3 1/D 2.67 0.05±0.01 2.33±0.17 0.54±0.12 0.85 315.14 340.92

B. bambos A1 1/D 0.74 0.16±0.03 2.33±0.10 - 0.95 200.39 212.72

A2 1/D 0.22 0.10±0.03 0.79±0.04 - 0.93 216.08 228.41

A3 1/D 1.04 0.25±0.05 2.61±0.25 -0.39±0.22 0.94 206.16 225.53

B. nutans A1 1/D 2.28 0.11±0.02 2.75±0.13 - 0.80 297.12 267.17

A2 1/D 0.41 0.10±0.03 0.83±0.06 - 0.81 291.67 205.72

A3 1/D 1.68 0.09±0.02 2.11±0.165 0.49±0.15 0.80 550.33 578.54

D. hamiltonii A1 1/D 1.77 0.13±0.03 2.59±0.11 - 0.84 554.83 572.78

A2 1/D 0.61 0.07±0.01 0.87±0.03 - 0.88 545.05 563.01

A3 1/D 1.68 0.09±0.02 2.12±0.166 0.49±0.14 0.87 550.33 578.54

D. stocksii A1 1/D 1.35 0.20±0.06 2.31±0.18 - 0.81 268.19 281.43

A2 1/D 0.38 0.18±0.05 0.723±0.04 - 0.84 253.26 265.50

A3 1/D 1.39 0.175±0.05 1.37±0.24 0.80±0.165 0.84 258.48 279.28

D. strictus A1 1/D 1.39 0.087±0.01 2.63±0.05 - 0.95 287.58 308.17

A2 1/D 0.43 0.06±0.01 0.86±0.02 - 0.96 309.38 329.97

A3 1/D 1.82 0.07±0.01 2.17±0.11 0.46±0.11 0.96 283.21 315.57

B. vulgaris A1 1/D 1.35 0.20±0.06 2.31±0.18 - 0.81 268.18 281.43

A2 1/D 0.38 0.18±0.05 0.72±0.04 - 0.84 253.26 266.50

A3 1/D 1.38 0.18±0.05 1.37±0.24 0.798±0.16 0.83 258.47 279.28

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 Table 8
Validation of allometric models of culm fresh weight on 20%
dataset of seven bamboo species.
Species Model Id ABias (%) ARMSE AMAPE (%)

B. balcooa A1 -9.71 2.96 25.23

A2 -7.52 2.56 20.38

A3 -8.69 2.71 21.98

B. bambos A1 -4.99 3.16 23.14

A2 7.03 3.56 24.50

A3 -11.68 4.70 29.30

B. nutans A1 2.27 1.76 12.91

A2 -2.05 1.71 11.10

A3 -3.03 1.79 18.86

D. hamiltonii A1 -7.08 4.16 26.07

A2 -8.24 3.93 23.98

A3 -11.28 3.57 24.56

D. stocksii A1 7.01 4.70 27.66

A2 6.11 3.87 21.46

A3 6.92 3.89 22.44

D. strictus A1 -5.32 0.86 28.15

A2 -3.06 0.81 30.10

A3 -2.62 0.79 27.21

B. vulgaris A1 7.01 4.70 29.66

A2 6.11 3.77 22.46

A3 6.92 3.83 25.44

ModelA1 found to be outperformer for fresh culm biomass prediction of B. bambos as it has highest fitting accuracy
with highest adj.R2 (0.95), lowest AIC(200.39) and BIC (212.72) and lowest prediction error with ABias (-4.99%) ARMSE
(3.16) and AMAPE (23.14%). A2 model was found to be outperformer with maximum fitting and prediction accuracy
over A1 and A3 model for five bamboo species viz., B. balcooa, B. nutans, D. hamiltonii, D. stocksii and B. vulgaris. For
the five mambo species, A2 model shown maximum fitting accuracy with highest Adj.R2 (0.80-0.89), lowest AIC (253.26-
545.05) and BIC(205.72-563.01). Further, highest prediction accuracy of A2 by lowest ABias (-8.24 to 6.11%), ARMSE
(1.71 to 3.93) and AMAPE (11.10-23.98%).Whereas A3 was found to be most suitable for D. strictus, as it has highest
fitting accuracy [highestAdj.R2 (0.96), lowest AIC (283.21) and BIC (315.57)] and lowest prediction error [ABias (-2.62%)
ARMSE (0.79) and AMAPE (27.21%)].
3.3 Generate synthetic values as a proxy height using H-D model

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The results of fitted general power form allometric model for H-D modelling in the seven bamboo species revealed that
the Adj.R2 value of the fitted model for the seven bamboo species was found to be >0.7 except the species B. nutans
with Adj.R2=0.69 (Table 9). Whereas, highest Adj.R2 (0.89) was observed in B. bambos. Lowest AIC (156.04) and BIC
(497.83) was observed in B. bambos. While highest AIC (497.83) and BIC (485.78) value was found in D. hamiltonii.
Further, evaluation results about prediction accuracy of the fitted models H-D were done and found that bias of the fitted
models was extended between -1.57 (in B. vulgaris) and-3.91 (in D. stocksii). The RMSE ranged between 1.16 (in D.
strictus) and 2.70 (in D. hamiltonii). Lowest value of MAPE (9.34) was found in B. bambos, whereas highest MAPE
(20.52) was observed for D. hamiltonii (Table 10).

Table 9
Parameter estimates of H-D allometric model fitted on 80% dataset of seven bamboo
species.
Species Weight variable Parameter Adj.R2 AIC BIC

[Estimate± std. error]

a b

B. balcooa 1/D 0.91 1.28±0.19 1.17±0.09 0.70 318.85 335.26

B. bambos 1/D 0.45 1.29±0.15 1.11±0.06 0.89 156.04 168.37

B. nutans 1/D 1.07 2.58±0.37 0.89±0.09 0.69 254 260.74

D. hamiltonii 1/D 0.38 2.30±0.30 0.92±0.06 0.70 497.83 485.78

D. stocksii 1/D 0.41 1.55±0.21 1.18±0.09 0.76 232.66 247.99

D. strictus 1/D 0.13 1.73±0.10 0.96±0.03 0.84 438.77 459.36

B. vulgaris 1/D 1.057 1.09±0.22 1.24±0.11 0.71 220.15 224.39

Table 10
Validation of H-D allometric model on 20% dataset
of seven bamboo species.
Species Bias (%) RMSE MAPE (%)

B. balcooa -3.33 1.61 13.73

B. bambos -2.26 1.25 9.34

B. nutans 2.14 1.47 12.80

D. hamiltonii -3.65 2.70 20.52

D. stocksii -3.91 1.58 12.61

D. strictus -1.72 1.16 16.66

B. vulgaris -1.57 1.48 10.67

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4. Discussion
Biometric parameters of the seven bamboo species

In bamboo culm has surprising biomass productivity due to its habit of producing sprouting from horizontal rhizome
systems annually that allows annual harvesting without exposing the soil surface. Bamboo plays an important role in
global carbon cycle, by sequestering atmospheric carbon through its biomass accumulation process39–42. In seven
bamboo species studied, among three major biomass component (culm, branch and foliage), culm is the most
important with highest share (69.56 to 78.71%) to above ground biomass which is more than double to combine
biomass share of branch and foliage. Moreover, bamboo culm has high commercial value due its multi-purpose
utility43. Distribution of percentage (%) share of culm, branch and foliage to above ground fresh weight varies
significantly between different bamboo species (Figure 2).

Analysis of variance (ANOVA) analysis revealed significant (p<0.05) differential productivity of above ground biometric
components (culm diameter, culm height, culm fresh weight, foliage fresh weight and above ground fresh weight) for
the seven bamboo species. Tukey’s HSD post-hoc test, revealed that on an average D. hamiltonii has highest biometric
productivity for culm diameter D (5.99 cm), culm height H (11.91m), culm fresh weight (16.92kg), foliage fresh weight
(1.53kg) and above ground fresh weight (20.39kg). Forbranch fresh weight, though D. hamiltonii was found to be have
the highest productivity but not significantly higher than B. balcooa, B. bambos, B. nutans and B. vulgaris (Table 3).
Differential growth rate of the seven bamboo species, evidenced need of species specific relationship rather general
relationship study for establishing model for biomass prediction.

Ratio of dry to fresh weight of seven bamboo species were estimated for culm, branch, foliage and above ground
biomass to convert fresh weight to dry weight using the formula: dry weight (kg) = Ratio × fresh weight (kg). The ratio in
different biomass components within species found significantly different. The highest dry to fresh weight ratio was
found for culm (0.63), branch (0.62) and above ground biomass (0.61) in B. nutans, where as, in foliage highest ratio
(0.53) was achieved for in D. strictus. The relationship equation between dry weight and fresh weight indicated that
variability of fresh weigh directly proportional to dry weight (biomass) (Table 4).

Development and validation of allometric models

From the analysis of above ground and culm biomass, it was found that biomass production in the seven bamboo
species has a high positive correlation with diameter at breast height (D) and culm height (H) but did not have
significant correlation with age. This was similar to findings point out by Yen et al.42 for bamboo species S. dullooa, P.
polymorphum and M. baccifera. So, D, combination of D and H was used for prediction of biomass using combine data
of more than three years oldbamboo plantation. Many authors already observed D and D 2H as good predictor variable
for bamboo allometric modelling 27–29, 34–35. The results revealed that D was a valid independent variable but using
volume (D 2H) as compound independent variable or incorporation of H as separate independent variable, further
improved prediction accuracy for both above ground biomass and culm biomass except bamboo species B. bambos
and B. nutans for fresh above ground biomass and B. bambos for fresh culm weight prediction (Table 5-8). Here, use of
H as additional independent variable did not improve prediction accuracy but increase prediction error may be due to
measurement error in H data or incorporation of H may have masking effect on D during model fitting. A1 with D as
independent variable alone, performed very well and usually very close to A2 and A3 model. Further biomass equation
using D as independent variable has practical advantage of easy measurement45–46. A1 model was also reported as
reliable model for biomass prediction by Yen et al.25, Kaushal et al.21.

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General power form allometric equation A2 with volume (D 2H) as an independent variable, found to be most reliable for
above ground fresh weight prediction in bamboo species D. stocksii and B. vulgaris, whereas for fresh culm weight
prediction A2 to be outperformer forfive bamboo species viz., B. balcooa, B. nutans, D. hamiltonii, D. stocksii and B.
vulgaris. This finding was consistent with allometric models established for biomass prediction in thin-walled bamboo
species by Singnar et al.47 and general findings of Chave et al.37 in woody species for above ground biomass prediction
using bole volume (D 2H).

Moreover, three parameter allometric equation A3 with D and H as independent variables, found to be most reliable in
bamboo species B. balcooa, D. hamiltonii and D. strictus for above ground fresh weight prediction, whereas for fresh
culm weight prediction A3was most appropriate model in D. strictus. For A3 use of D and H as independent variable
provided more flexibility to establish relationship with biomass response.

The fitting and prediction accuracy of best fitted allometric models for fresh above weight prediction were visually
investigated through line plots. The residual plot of prediction error values evidenced that allometric models using
weighted maximum likelihood nonlinear fixed effects modelling method capture heteroscedastic relationship between
the biomass response and the predictor variables well and produced much lower residuals. Further, residuals found to
be random (lack of any pattern) visually which indicate high fitting accuracy (Fig. 3-4).

Generate synthetic values as a proxy height using H-D model

It was very often that best fitted allometric models were not possible to apply due to missing observation of H48. To get
H value, synthetic H values were generated as proxy H using H-D allometric model. H-D allometric equation developed
for the seven bamboo species revelaed high fitting accuracy with high Adj.R2 (>0.7) except B. nutans (0.69). Highest
fitting accuracy was observed in B. bambos, whereas lowest value observedin D. hamiltoni i(Table 9-10). Similar, result
was also pointed out by Kempes et al.9 and Sileshi10. H-D relationship was developed using >3 year old bamboo
plantation where H reached to plateau i.e., there is no growth in diameter and height49. As per our knowledge, for the
first time this study has established allometric modelling for quantifying H-D relationship between the seven bamboo
species viz., B. bambos, B. nutans, D. stocksii, B. vulgaris, B. balcooa, D. hamiltonii, and D. strictus.

5. Conclusions
In this study species-specific two-parameter allometric equations with simple as well as compound regressor variable
and three parameter allometric equations were developed for above ground biomass. Further, species specific height-
diameter models were also developed to predict bamboo height, which can be used as a proxy of height variable and
used for allometric equation without measuring height of bamboo. Since the allometric equations were developed using
rigorous criteria, we can also conclude that the developed biomass estimation equations provides better biomass
predication for culm as well as above ground biomass. The models can expected to be increase accuracy in estimation
of biomass and carbon sequestration in bamboo based ecosystems of India under managed natural stands or
plantation grown under similar condition. Moreover, considering the wide multipurpose use of the bamboo species, the
developed models may give provide useful information about culm biomass and above ground biomass to forestry
professionals, bamboo farmers, and other stakeholders.

Declarations
Competing interests

The authors declare that they have no competing interests.

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Declarations

The planting material was obtained from different Government nurseries from different part of the country and for
which all the permissions were obtained. All the collection complies with relevant institutional, national, and
international guidelines and legislation.

Funding

This research was funded by National Bamboo Mission, New Delhi, State Forest Department, Uttarakhand, India and
International Bamboo and Rattan Organisation (INBAR) through its grants from IFAD and Dutch Government and from
the CGIAR Research Program on Forests, Trees and Agroforestry (FTA).

Acknowledgements

The financial help received from both the funding agency is duly acknowledged. We also thank ICAR and Director ICAR-
IISWC for providing the permission to conduct the experiment. The technical assistance provided by Mr. UVS Chauhan
and Mr. Ravish is gratefully acknowledged.

Author contributions

R.K., S.I., S.T., J.D. conceptualized the research and design the experiments; R.K. contributed to experimental materials;
J.M.S.T., S.T., executed the experimental data collection, R.K., S.I. conducted data analysis and interpretation; S.I., R.K.,
M.M. prepared the manuscript; T.L.T. and J.D. secured funding and edited the manuscript. All authors reviewed the
manuscript.

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Figures

Figure 1

Location map of the study sites.

Figure 2

Distribution of percentage (%) share of culm, branch and foliage to above ground biomass in different bamboo species.

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Figure 3

A1-A4 plots potraited observed vs best fitted allomteric models for above ground fresh weight prediction in B. balcooa,
B. bambos, B. nutans, D. hamiltonii, respectively. Whereas B1-B4 plots potraited weighted residuals vs best fitted above
ground fresh weight values in B. balcooa, B. bambos, B. nutans, D. hamiltonii, respectively.

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Figure 4

A5-A7 plots potraited observed vs best fitted allomteric models for above ground fresh weight prediction in D. stocksii,
D. strictus, respectively. Whereas B5-B7 plots potraited weighted residuals vs best fitted above ground fresh weight
values in D. stocksii, D. strictus, respectively.

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