Trees, Forests and People 5 (2021) 100098

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Trees, Forests and People

journal homepage: www.elsevier.com/locate/tfp

Review Article

A critical review of forest biomass estimation equations in India

Biplab Brahma a, Arun Jyoti Nath a,∗, Chandraprabha Deb a, Gudeta W Sileshi b,c,
Uttam Kumar Sahoo d, Ashesh Kumar Das a
a
Department of Ecology and Environmental Science, Assam University, Silchar, Assam, 788011, India
b
School of Agricultural, Earth and Environmental Sciences, University of Kwazulu-Natal, Pietermaritzburg, South Africa
c
College of Natural and Computational Sciences, Addis Ababa University, P.O. Box 1176, Addis Ababa, Ethiopia
d
Department of Forestry, Mizoram University, Aizawl, India

a r t i c l e i n f o a b s t r a c t

Keywords: Plant biomass is an integral part of the global carbon cycle and a renewable energy source that can deaccelerate
Biomass equation the rising global temperature. India has 71 million ha (M ha) of land under forests represented by tropical to
Power-law function alpine ecosystems. Numerous direct and indirect species-specific and mixed-species equations have been used for
Root mean square error
biomass estimation in India. Biomass estimation equations that facilitate the prediction of aboveground biomass
Diameter at breast height
(AGB) stocks non-destructively across India are still lacking. Therefore, the objective of this review is to (i) assess
Species-specific equation
Mixed species equation the existing species-specific biomass estimation equations for trees, bamboos, palms, and bananas in India, (ii)
assess and identify the most appropriate multi-species biomass estimation equations for AGB estimation across
India, and (iii) define the critical research gaps in biomass estimation in India. The literature search found 85
species-specific and six multi-species AGB estimation equations reported from India. It was also found that a 50%
of these equations were based on the power-law function using diameter at breast height (D) as the predictor
variable. We carried out a multi-fold validation to compare the multi-species equation’s compatibility by compar-
ing the root mean square error (RMSE). The estimated RMSE values of the six reported multi-species equations
showed that the following two equations could be effectively used for estimation of AGB: (i) lnAGB= 0.349+1.316
lnGBH and (ii) AGB= (0.18D2.16 ) × 1.32. These are adequate for predicting biomass of any woody species across
a range of conditions in India.

1. Introduction 2014; Li et al., 2015), stages of forest development, nutritional status,

Plant biomass is an integral part of the global carbon cycle, and its
management is vital for mitigating climate change (Houghton et al.,
2009; Souza et al., 2017; Nath et al., 2021). Notwithstanding, the grow-
ing biomass can remove carbon dioxide from the atmosphere and store
it for an extended period. It can play a critical role in limiting the rise
in global temperature. Recent studies have explored global mitigation
pathways to limit the increase in global average temperature to below
2 °C above the pre-industrial levels (Popp et al., 2017; Griscom et al.,
2017). Among the various carbon pools, vegetation carbon can be man-
aged with relative ease to reduce the atmospheric carbon concentra-
tions. Multiple studies have shown that vegetation’s carbon sinks poten-
tial could be significantly enhanced by adhering to sustainable forest
management principles (Griscom et al., 2017). Interest in biomass stud-
ies has increased globally due to its importance as a source of food,
energy, and fiber (Houghton et al., 2009; Wang et al., 2020). Biomass
production in a given forest depends on several factors such as location,
vegetation types/species composition (Barbosa et al., 2014; Luo et al.,
and edaphic characteristics, besides factors related to natural distur-
bance, anthropogenic pressure, and management actions (Powell et al.,
2014). Hence, over the years, major emphasis has been made to estimate
terrestrial biomass accurately by incorporating uncertainties related to
forest management (Weiskittel et al., 2015).
Different direct and indirect methods have been employed for
biomass estimation. In the direct methods, a representative sample of
trees in a given area is harvested and measured to estimate the dry
weight of different tree components (Brahma et al., 2018). However, di-
rect methods can be expensive, especially when dealing with large sam-
ple areas and several tree species (Sileshi, 2014). In the indirect method,
biomass is estimated using biomass estimation equations (BEEs). BEEs
relate measurable variables such as tree height (H), diameter at breast
height (D), and wood density (𝜌) to total tree biomass (Chave et al.,
2005). Therefore, indirect methods are often preferred over direct meth-
ods (Vashum and Jayakumar, 2012). Multi-species pan-tropical equa-
tions have been developed to estimate aboveground biomass (AGB)

∗
Corresponding author: Arun Jyoti Nath, Department of Ecology and Environmental Science, Assam University, Silchar, Assam, 788011, India.
E-mail address: arunjyotinath@gmail.com (A.J. Nath).

https://doi.org/10.1016/j.tfp.2021.100098
Received 8 April 2021; Received in revised form 8 May 2021; Accepted 9 May 2021
Available online 13 May 2021
2666-7193/© 2021 The Author(s). Published by Elsevier B.V. This is an open access article under the CC BY-NC-ND license
(http://creativecommons.org/licenses/by-nc-nd/4.0/)
B. Brahma, A.J. Nath, C. Deb et al. Trees, Forests and People 5 (2021) 100098

Quantification of biomass at the local, regional and landscape lev-

Abbreviations els is essential to support India’s nationally determined contributions
(NDCs) to the united nations framework convention on climate change
AGB aboveground biomass (UNFCCC). India is a party to this convention and aims to limit the global
BaD basal diameter atmospheric temperature rises by 2 °C by limiting the global greenhouse
BB bole biomass gas (GHG) emission. In this direction, the Indian government has imple-
BECCS bioenergy with carbon capture and storage mented various activities to improve its carbon stock, sustainable use
BEE biomass estimation equations of biomass to reduce fossil fuel emissions, and bio-based products via
BGB belowground biomass bioenergy with carbon capture and storage (BECCS). India is situated in
BWB bole wood biomass four biodiversity hotspots’ confluence and covers nearly 71 M ha of the
C carbon land area under forests, covering around 23% of the total geographical
CBH circumference at breast height area (FSI, 2019). Two major biodiversity hotspots, such as the Eastern
CrD crown diameter Himalaya and the Western Ghats, have widened India’s biomass carbon
CD collar diameter research scope.
CF correction factor The biomass equations developed in India vary significantly in terms
COP conference of Parties of the data set used, size of the field inventory, result validation, etc.
CrH crown height The uncertainty in biomass estimation hinders accurate estimation of
CRB coarse root biomass biomass carbon accounting, urging the development of suitable and ro-
CrH crown height bust equations (Weiskittel et al., 2015). This article aims to review the
CrW crown width existing biomass equations to prepare an up-to-date database of biomass
D diameter at breast height estimation equations for India. This synthesis aims to determine the un-
FAO food and agriculture organization certainties associated with existing equations when applied to a single
FR fine root tree and a forest stand. Finally, it aims to determine the best biomass
FRB fine root biomass estimation equations that can be applied to diverse forest types in the
FSI forest survey of India Indian context. Considering the limitations of species-specific equations
GBH girth at breast height for biomass estimation in a mixed-species stand, we have tested the per-
GHG greenhouse gas emission formance of published mixed-species equations through the data vali-
H tree height dation process.
H-D height-diameter
LRB lateral root branch 2. Materials and methods
NDC nationally determined contribution
NB number of branches The methodology used in this review involved two steps: (1) compi-
NPB number of primary branch lation of all available biomass equations for India and (2) validation of
PBG primary branch girth the equations.
PsS Pseudo stem Articles published on India’s biomass estimation equations were ac-
MSE mean square error cessed from Web of Science and Google Scholar. A combination of
NB number of branches keywords including "tree biomass estimation," "biomass stock estima-
RMSE root mean square of error tion," "biomass carbon stock," "allometric biomass equations," "above-
SRB stump root ground biomass stock," "belowground biomass stock," "total biomass
TaRB tap root biomass stock," "biomass estimation model development," "root biomass estima-
TB total biomass tion model development," were used to search published articles. As-
UNFCCC united nation’s framework convention on climate terisk (∗ ) has been with the keywords to avail the plurals, e.g., "above-
change ground biomass stock." A total of 200 published articles were down-
loaded for critical screening. Research articles were analyzed for infor-
mation on the equation’s various aspects and their application and to
for major forest types (Chave et al., 2005, 2014; Vashum and Jayaku- filter the desired information as per the article’s objective. Fifty-four
mar, 2012; Nath et al., 2019). Despite significant progress in biomass (54) articles were finally selected to gather the necessary information.
estimation methods, quantifying biomass stocks for diverse forest types The following criteria were used for selecting the articles: (1) The tar-
in India is very challenging due to the lack of generalized biomass esti- geted articles should have accurate information about the study sites,
mation equations (Nath et al., 2019; Giri et al., 2019). (2) The articles should have dealt with the biomass estimation model
Estimating tree volume is another indirect and alternative tree development, (3) The articles should have the proper description about
biomass estimation method (Thangjam et al., 2019; Brown, 1997). In the method of data collection, etc. The primary emphasis was on extract-
this approach, the wood density or biomass expansion factor is multi- ing information on different biomass pools such as AGB, belowground
plied with the corresponding volume to obtain the biomass of a tree biomass (BGB), total biomass (TB), and components like a branch, bole,
species (Brown, 1997; Mugasha et al., 2016; Romero et al., 2020). In In- and roots. Different variables used were equation expressions, age of
dia, the Forest Survey of India has developed volume equations for the the stand or tree (yr), the number of trees harvested, associated species
dominant trees (FSI, 1996). The physical and mechanical constraints in name, and the source extracted from the articles were tabulated sep-
H measurements of standing trees often produce significant estimation arately for each pool, and component (AGB, BGB, TB, branch, and
ambiguities. H that plays a vital role in volume equation development bole). The locations of the reported studies considered in the present
(Gonzalez-Benecke et al., 2014), is influenced by various attributes of manuscript are given in Fig. 1.
a stand (e.g., species, tree age, the slope of the terrain, and elevation) The dataset of D, H, and 𝜌 for different species were collected
(Stereńczak et al., 2019). The measurement of H in difficult terrain situ- from 7 (seven) different articles (Deb et al., 2016; Sandeep et al.,
ations and closed forest stand often leads to bias, and therefore volume 2015; Chaturvedi and Raghubanshi, 2015; Jain and Ansari, 2013;
equation using H may have an error source. To avoid this uncertainty in Tripathi, 2015; Baishya and Barik, 2011; Das and Ramakrishnan, 1987)
the measurements, researchers prefer to use BEEs for estimation of tree to validate the mixed biomass equations. The root mean square er-
biomass. rors (RMSE) (Sileshi, 2014) for mixed species biomass equations were

2
B. Brahma, A.J. Nath, C. Deb et al.
estimated. In the absence of wood density in the respective research
articles, the values recommended for the same by FAO (1997) and
Manikandan et al. (2019) were used. Finally, the average RMSE val-
ues were computed to compare the generalized equation to recommend
the best equation for biomass estimation in India.
3. Results and discussion
3.1. Aboveground Biomass estimation equations
The AGB stock represents 70‒80% of the total vegetative biomass
stock in tropical forests (2016). It has been recognized as the most cru-
cial biomass component to represent vegetation carbon stock in terres-
trial ecosystems and has been studied widely. Moreover, AGB estimation
is relatively easy than the BGB.
The review revealed that forest biomass estimations had been carried
out by both direct harvest and several indirect methods at the regional
and landscape levels. Among these, direct harvest was used in around
85% of the total studies analyzed in this article. The complex association
of forest tree species is the predominant factor limiting the implications
of direct harvesting and biomass equation development for multi-species
3
Trees, Forests and People 5 (2021) 100098
Fig. 1. : Locations of the reported studies considered
for the review article.
stands (Nath et al., 2019). The error in biomass prediction using species-
specific equations for multi-species stand could be high. However, the
use of multi-species biomass equations can reduce the error in biomass
estimation at the regional or landscape level.
Bamboos are fast growing, and therefore their growth and wood den-
sity considerably vary with age and species. Therefore, for bamboos, the
development of age-specific and species-specific equations are strongly
recommended for accurate biomass estimation. Six studies have been
carried out in India to develop biomass equations for ten different bam-
boo species (Table 1). However, the reliability of these equations refer-
ring to future use depends much on various factors such as the accessi-
bility of the used independent variables, the form of the equations, and
the number of harvested data used in equation development. D is an
easily accessible variable for any equation. It is mostly recommended to
predict different plant biomass with more accuracy than other variables
(Lui et al., 2017). H as a covariate in biomass equations have been used
frequently in volume equations for AGB estimation in bamboos. At least
15 bamboo culm biomass data has been used to develop the age-specific
Power-law equations for bamboo culms under<1 to >3 yr of age for dif-
ferent species in India. The AGB equations for Dendrocalamus strictus,
Melocanna baccifera, Pseudostachyum polymorphum, and Schizostachyum
B. Brahma, A.J. Nath, C. Deb et al. Trees, Forests and People 5 (2021) 100098

Table 1
. Details of the recommended regression equations for aboveground biomass estimation for diverse lifeforms in India.
Harvested sample
Associated Species Equations Age (yr) size (number) Sources
Bamboos
Bambusa arundinacea AGB= 15.487 + 134.779D2 H 1 25 Chandrashekara, 1996
Bambusa arundinacea AGB= 13.371 + 145.202D2 H 2 25 Chandrashekara, 1996
Bambusa arundinacea AGB= 16.372 + 148.869D2 H 3 25 Chandrashekara, 1996
Bambusa arundinacea AGB= 14.668 + 120.826D2 H >3 25 Chandrashekara, 1996
Bambusa cacharensis ln AGB= 2.078+2.140 lnD <1 27 Nath et al., 2009
Bambusa cacharensis ln AGB= 2.134+2.268 lnD 1 27 Nath et al., 2009
Bambusa cacharensis ln AGB= 2.174+2.306 lnD 2 27 Nath et al., 2009
Bambusa cacharensis ln AGB= 2.184+2.178 lnD 3 27 Nath et al., 2009
Bambusa vulgaris ln AGB= 2.281+2.149 lnD <1 20 Nath et al., 2009
Bambusa vulgaris ln AGB=2.386+2.079 lnD 1 20 Nath et al., 2009
Bambusa vulgaris ln AGB= 2.554+1.956 lnD 2 20 Nath et al., 2009
Bambusa vulgaris ln AGB=2.548 + 1.970 lnD 3 20 Nath et al., 2009
Bambusa balcooa ln AGB=2.149 + 2.284 lnD <1 20 Nath et al., 2009
Bambusa balcooa ln AGB =2.199+2.353 lnD 1 20 Nath et al., 2009
Bambusa balcooa ln AGB=2.368 + 2.214 lnD 2 20 Nath et al., 2009
Bambusa balcooa ln AGB= 2.153+2.477 lnD 3 20 Nath et al., 2009
Bambusa bambos AGB= −3.225.8 + 1730.4D 9 Kumar et al., 2006
Dendrocalamus strictus AGB=0.1002D2.266 1 to 4 112 Kaushal et al., 2016
Melocanna baccifera lnAGB= ln(−1.83) +1.11(lnD2 ) 1 to 5 180 Singnar et al., 2017
Pseudostachyum polymorphum lnAGB= ln(−7.76) +0.82(ln(D2 H)) 1 to 5 90 Singnar et al., 2017
Schizostachyum dullooa lnAGB= ln(−4.33) +0.51(ln(D2 H)) 1 to 4 132 Singnar et al., 2017
Schizostachyum pergracile AGB=−2.99+1.83D <1 15 Thokchom and Yadava, 2017
Schizostachyum pergracile AGB= −3.12+2.12D 1 15 Thokchom and Yadava, 2017
Schizostachyum pergracile AGB = −3.09+1.95D 2 15 Thokchom and Yadava, 2017
Schizostachyum pergracile AGB = −2.92+1.63D 3 15 Thokchom and Yadava, 2017
Schizostachyum pergracile AGB = −2.34+1.51D >3 15 Thokchom and Yadava, 2017
Bananas
Musa balbisiana AGB= (exp(−4.54+0.874(lnD2 Hp)) x 1.06 140 Laskar et al., 2020
Trees
Acacia auriculiformis lnAGB= −1.974+ 0.827lnD2 H 165 Kumar et al., 1998
Ailanthus triphysa lnAGB= −2.212+1.804 ln D + 0.470 ln H 170 Kumar et al., 1998
Avicennia marina AGB= 3.2918D-0.9664 19 Trivedi et al., 2017
Azadirachta indica AGB=0.016H 4.862 9 Mohamed et al., 2018
Azadirachta indica AGB=0.498BaD1.729 9 Mohamed et al., 2018
Azadirachta indica AGB=0.213D2.109 9 Mohamed et al., 2018
Azadirachta indica AGB=19.63CrH2.599 9 Mohamed et al., 2018
Azadirachta indica AGB=9.235CrW1.326 9 Mohamed et al., 2018
Barringtonia acutangula AGB= 0.04D2.38 20 Nath et al., 2017
Casuarina equisetifolia AGB= −0.378+0.033D2 H 127 Kumar et al., 1998
Casuarina equisetifolia lnAGB= −1.785+2.495lnD 2 to 9 90 Vidyasagaran and Paramathma, 2014
Citrus sinensis AGB=(exp(0.79 + 0.20(lnD2 H))) × 1.08 8 to 30 58 Sahoo et al., 2020
Eucalyptus tereticornis AGB= 0.493D1.818 7 10 Kumar et al., 2019
Grewia optiva AGB=0.692D1.667 4 to 23 25 Verma et al., 2014
Hevea brasiliensis AGB= (exp (−3.31+ 0.95 (lnD2 H)) × 0.12 6 to 34 67 Brahma et al., 2017
Hevea brasiliensis lnAGB= −2.9 + 2.57lnD 6 15 Brahma et al., 2017
Hevea brasiliensis ln AGB= −2.01+2.25lnD 15 19 Brahma et al., 2017
Hevea brasiliensis ln AGB = −1.71+0.78lnD2 H+𝜀 27 15 Brahma et al., 2017
Hevea brasiliensis lnAGB = −3.08+0.93lnD2 H 34 18 Brahma et al., 2017
Hevea brasiliensis AGB= 2.216G2.702 2 to 5 51 Chaudhuri et al., 1995
Hevea brasiliensis AGB= 2.284G2.672 2 to 5 47 Chaudhuri et al., 1995
Hevea brasiliensis AGB= 2.278479G2.6823 2 to 5 98 Chaudhuri et al., 1995
Hevea brasiliensis AGB= 0.02020G2.492 34 Dey et al., 1996
Jatropha curcus AGB= 0.091(CD)1.725 8 16 Rao et al., 2016
Litchi chinensis AGB= (24.83/[1 + 0.03 exp(−0.41 × CD)]ˆ(1/0.01) 2 to 10 30 Naik et al., 2018
Leucaena leucocephala lnAGB= −1.543+2.248lnD 147 Kumar et al., 1998
Mallotus philippensis ln AGB= −1.24 + 1.28 lnGBH 9 Rana et al., 1989
Mangifera indica AGB= 2.886(PBG × NPB)1.039 3 to 85 74 Ganeshamurthy et al., 2016
Pinus roxburghii lnAGB =−6.398 + 2.655 lnGBH 16 to 128 26 Chaturvedi and Singh, 1987
Pinus kesiya AGB =1.3503–3.4145D+4.8678D2 –1.352D3 40 Baishya and Barik, 2011
Populus deltoides AGB= 1.72D1.21 1 to 9 27 Das et al., 2011
Psidium guajava AGB=0.069CD2.456 4 to 14 17 Rathore et al., 2018
Quercus leucotrichophora ln AGB= 0.685 + 1.254 lnGBH 23 Rawaat and Singh, 1988
Quercus floribunda ln AGB= 0.028 + 1.429 lnGBH 17 Rawaat and Singh, 1988
Rhododendron arboreum ln AGB= 1.176 + 0.855 lnGBH 8 Rawaat and Singh, 1988
Shorea robusta ln AGB= - 1.79 + 1.89 lnGBH 9 Rana et al., 1989
Tectona grandis lnAGB= 8.902 + 7.873/(1+(lnpD2 H/14.05)−6.780 100 Chaturvedi and Raghubanshi, 2015
∏
Tectona grandis AGB=0.06 p( D2 /4)H 5 to 40 70 Sandeep et al., 2015
Tectona grandis AGB=0.4989D2 –0.202D-21.971 1 to 30 33 Jha, 2015
Tectona grandis AGB= 0.26+730.55D2 H 15 to 20 12 Chandrashekara, 1996
Terminalia chebula AGB= 0.1296D2.0827 10 Panwar et al., 2014
Mixed woody species AGB=0.18D2.16 × 1.32 303 Nath et al., 2019
Mixed woody species AGB= 0.32pD2 H0.75 × 1.34 303 Nath et al., 2019
Mixed woody species ln AGB= 3.428 + 0.310 lnpD2 H 470 2012
Mixed woody species ln ABG= −3.206+1.337 lnpD2 H 460 Chaturvedi and Raghubanshi, 2013
Mixed woody species ln AGB= −4.31 + 2.21 lnGBH 18 Naik et al., 2018
Mixed woody species ln AGB= 0.349 + 1.316 ln GBH 48 Rawat and Singh, 1988

Note: AGB= Aboveground Biomass, BaD=Basal diameter, CD= Collar diameter, CrH= Crown height, CrW=Crown width, CrD= Crown diameter, D=Diameter at breast
height of the tree/culm (DBH), G= Girth at 15 cm from bud union, GBH= Girth at breast height, H= Tree/culm height, PBG= Primary branch girth and NBG=Number
of the primary branch, p= wood density, 𝜀= Model error.

4
B. Brahma, A.J. Nath, C. Deb et al.
Table 2
Details of the recommended regression equations for bole biomass estimation.
Associated Species
Bamboos
Bambusa arundinacea
Bambusa arundinacea
Bambusa arundinacea
Bambusa arundinacea
Dendrocalamus strictus
Dendrocalamus strictus
Bananas
Musa balbisiana
Trees
Acacia nilotica
Acacia nilotica
Acacia nilotica
Acacia catechu
Albizia lebbeck
Albizia procera
Alnus nepalensis
Alnus nepalensis
Alnus nepalensis
Barringtonia acutangula
Bauhinia racemosa
Casuarina equisetifolia
Cinnamomum camphora
Cinnamomum camphora
Citrus sinensis
Dalbergia sissoo
Dalbergia sissoo
Dalbergia sissoo
Dalbergia sissoo
Dalbergia sissoo
Eucalyptus tereticornis
Eucalyuptus tereticornis
Eucalyuptus tereticornis
Embilica officinalis
Gaultheria griffithiana
Gaultheria griffithiana
Grewia optiva
Gmelina arborea
Gmelina arborea
Hevea brasiliensis
Hardwickia binata
Jatropha curcus
Jatropha curcus
Jatropha curcus
Jatropha curcus
Jatropha curcus
Litchi chinensis
Lannea coromandelica
Leucaena leucocephala
Leucaena leucocephala
Lyonia ovalifolia
Lyonia ovalifolia
Mallotus philippensis
Miliusa tomentosa
Neolitsea zeylanica
Neolitsea zeylanica
Populus deltoides
Populus deltoides
Populus deltoides
Populus deltoides
Populus deltoides
Populus deltoides
Populus deltoides
Populus deltoides
Populus deltoides
Pinus kesiya
Pinus patula
Pinus patula
Pinus patula
Pinus roxburghii
Psidium guajava
Pyrularia edulis
Pyrularia edulis
Trees, Forests and People 5 (2021) 100098
Harvested sample
Suggested Equations Stand/ Tree age (yr) size (number) Sources
CB= 9.599 + 111.303D2 H 1 25 Chandrashekara, 1996
CB= 9.457 + 77.815D2 H 2 25 Chandrashekara, 1996
CB= 10.170 + 97.031D2 H 3 25 Chandrashekara, 1996
CB=8.298 + 77.672D2 H >3 25 Chandrashekara, 1996
CB=0.1051 × D2.4900 1 to 4 112 Kaushal et al., 2016
CB=0.0537D2.4260 1 to 4 112 Kaushal et al., 2016
PsS=(exp(−4.66–0.829(lnD2 Hp)) × 1.06 140 Laskar et al., 2020
BWB=109.474(1-e − 0.09D ) 2.685 4.5 to 6.5 75 Rizvi and Ahlawat, 2014
ln BWB=−6.459 + 1.879 lnD 4 20 Singh and Toky, 1995
ln BWB= −9.071 + 2.325 lnD 8 20 Singh and Toky, 1995
BB=−1.9418+1.3604D 25 Singh et al., 2011
BB= −0.2976+0.4172D 25 Singh et al., 2011
BB=0.038 D2.505 8 32 Newaj et al., 2016
lnBB =1.532 + 2.461 ln D 7 to 46 17 Sharma and Ambasht, 1991
ln BB = −8.762 + 0.209 ln Age Singh and Yadava, 1994
ln BB = −13.776 + 2.117 ln D Singh and Yadava, 1994
BB= 2.015D0.051 200 Nath et al., 2017
BB= 0.0431+0.0025CBH2 H 19 Kale et al., 2004
lnBB= 2.5584+2.6097 lnD 2 to 9 90 Vidyasagaran and Paramathma, 2014
ln BB =4.143+1.056 lnAge Singh and Yadava, 1994
ln BB = 1.387 + 1.750 ln D Singh and Yadava, 1994
BB= (exp(−0.19 + 0.80(lnH))) × 1.09 8 to 30 58 Sahoo et al., 2020
BB= −3.0374+ 2.7698D 5 12 Lodhiyal and Lodhiyal, 2003
BB=−11.0369+3.6005D 10 12 Lodhiyal and Lodhiyal, 2003
BB=22.0780+2.8541D 15 12 Lodhiyal and Lodhiyal, 2003
BB= −0.367+1.3457D 25 Singh et al., 2011
BB=0.832 D 1.593 14 42 Newaj et al., 2016
BB= 0.522D1.766 6 10 Kumar et al., 2019
ln BWB=−2.632 + 1.672 lnD 4 20 Singh and Toky, 1995
ln BWB=−24.266 + 5.724 lnD 8 20 Singh and Toky, 1995
BB=0.056 CD2.000 12 30 Newaj et al., 2016
ln BB = 2.880 + 0.714 lnAge Singh and Yadava, 1994
ln BB = −5.176 + 1.827 ln D Singh and Yadava, 1994
BB= 0.348D1.704 4 to 23 25 Verma et al., 2014
BB= −4.152+ 1.667D+ 0.026D2 1 to 6 75 Swamy et al., 2004
BB= −6.354+2.321D 1 to 6 75 Swamy et al., 2004
BB= (exp (−4.57+1.05 (lnD2 H)) × 1.03 6 to 34 67 Brahma et al., 2017
BB=0.232 D2.046 17 30 Newaj et al., 2016
BB= 0.019 CD2.036 8 16 Rao et al., 2016
BB=0.017PH1.149 8 16 Rao et al., 2016
BB= 0.016NB1.174 8 16 Rao et al., 2016
BB=1.66E−06 CrW2.618 8 16 Rao et al., 2016
BB=3.21E−10 CD4.331 8 16 Rao et al., 2016
BB=(9.78/[1 + 4.54 exp(−0.34 × CD)]ˆ(1/0.60)+E) 2 to 10 30 Naik et al., 2018
BB= −1.84+0.002CBH2 H 15 Kale et al., 2004
ln BWB=−3.063 + 1.721 lnD 4 20 Singh and Toky, 1995
ln BWB= −14.308 + 4.534 lnD 8 20 Singh and Toky, 1995
ln BB = −1.138 + 0.565 ln Age Singh and Yadava, 1994
ln BB = −4.607 + 1.204 ln D Singh and Yadava, 1994
ln BB= −2.14 + 1.40 lnGBH 9 Rana et al., 1989
BB= −0.68+0.0024CBH2 H 17 Kale et al., 2004
ln BB =−0.438 + 0.580 lnAge Singh and Yadava, 1994
ln BB =0.869 + 1.069 ln D Singh and Yadava, 1994
BB= 1.24D1.24 1 to 9 27 Das et al., 2011
BB=0.0034+4.2077D 5 27 Lodhiyal et al., 1995
BB= −1.0397+4.6333D 6 27 Lodhiyal et al., 1995
BB=−1.4539+6.0494D 7 27 Lodhiyal et al., 1995
BB=−7.3826+7.2773D 8 27 Lodhiyal et al., 1995
BB=−0.0036+0.5706D 5 27 Lodhiyal et al., 1995
BB=−1.4445+0.6393D 6 27 Lodhiyal et al., 1995
BB=−0.2034+0.8434D 7 27 Lodhiyal et al., 1995
BB= −1.0400+1.0249D 8 27 Lodhiyal et al., 1995
BB=1.8364–5.3988D +6.5046D2 –1.7410D3 40 Baishya and Barik, 2011
BB= −4.733+1.192D 15 12 Bargali and Singh, 1997
BB= 0.544+1.195D 18 12 Bargali and Singh, 1997
BB= 0.010+1.126D 21 12 Bargali and Singh, 1997
ln BB= −6.42 + 2.60 lnGBH 16 to 128 26 Chaturvedi and Singh, 1982
BB= 0.035 CD2.319 4 to 14 17 Rathore et al., 2018
ln BB = −8.172+ 1.094 lnAge Singh and Yadava, 1994
ln BB = −10.356 + 1.850 ln D Singh and Yadava, 1994
(continued on next page)
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B. Brahma, A.J. Nath, C. Deb et al. Trees, Forests and People 5 (2021) 100098

Table 2 (continued)

Associated Species Suggested Equations Stand/ Tree age (yr) Harvested sample Sources
size (number)

Quercus dealbata ln BB = 8.956 + 0.712 lnAge Singh and Yadava, 1994

Quercus dealbata
Quercus leucotrichophora
Quercus floribunda
Quercus fenestrata
Quercus fenestrata
Quercus griffithii
Quercus griffithii
Quercus griffithii
Quercus griffithii
Rhododendron arboreum
Rhododendron arboreum
Rhododendron arboreum
Shorea robusta
Styrax serrulatus
Styrax serrulatus
Tectona grandis
Tectona grandis
Tectona grandis
Tectona grandis
Tectona grandis
Tectona grandis
Ziziphus xylopyrus
Mixed woody species
Mixed woody species
Mixed woody species
ln BB = 12.465 + 0.712 ln D Singh and Yadava, 1994
ln BB= −0.523 + 1.367 lnGBH 23 Rawat and Singh, 1988
ln BB= −1.109 + 1.518 lnGBH 17 Rawat and Singh, 1988
ln BB = −5.145 + 0.915 lnAge Singh and Yadava, 1994
ln BB = −1.631 + 1.818 ln D Singh and Yadava, 1994
ln BB = 3.202 + 1.192 lnAge Singh and Yadava, 1994
ln BB = −9.103 + 1.913 ln Age Singh and Yadava, 1994
ln BB = −2.655 + 2.574 lnD Singh and Yadava, 1994
ln BB = −23.603 + 3.877 ln D Singh and Yadava, 1994
ln BB= 1.120+0.704 lnGBH 8 Singh and Yadava, 1994
ln BB = −5.689 + 1.084 ln Age Singh and Yadava, 1994
ln BB = −21.265+ 2.495 ln D Singh and Yadava, 1994
ln BB= −2.83+ 1.98lnGBH 9 Rana et al., 1989
ln BB = −4.536 + 1.038 ln Age Singh and Yadava, 1994
ln BB = −10.953 + 1.836 lnD Singh and Yadava, 1994
BB= 0.025D 2.817 20 to 47 Buvaneswaran et al., 2006
BB= 0.0581D 2.523 20 to 47 Buvaneswaran et al., 2006
lnBB=8.512+10.49/1+(lnpD2 H/15.36)−5.252 100 Chaturvedi and Raghubanshi, 2015
lnBB= −2.85+2.655 lnCBH 15 Kale et al., 2004
BB=0.3699D2 –0.1537D-17.8 1 to 30 33 Jha et al., 2015
BB= 0.942+512.69D2 H 15 to 20 12 Chandrashekara, 1996
lnBB= −3.20+2.87 ln CBH 15 Kale et al., 2004
ln BB= −3.258+1.283 lnpD2 H 460 Chaturvedi and Raghubanshi, 2013
ln BB= −0.86 + 1.43 lnGBH 48 Rawat and Singh, 1988
ln BB= −5.03 + 2.33 lnGBH 18 Rana et al., 1989

Note: CB= Culm Biomass, BB= Bole Biomass, BWB= Bole wood biomass, D= Diameter at breast height of the tree/culm (DBH), CBH= Circumference at breast height
of the tree, H= Tree/culm height, CD= Collar diameter of the tree, 𝜌=wood density, CrW=Crown width, CrD= Crown diameter and NB=Number of primary branch,
PsS= Pseudo Stem.

dullooa were developed using at least 90 culms. Only one study was
found in India describing AGB estimation of banana plants. An exponen-
tial form of regression was used using D2 H as an independent variable
from 140 harvested banana plants (Laskar et al., 2020).
Species-specific equations for the AGB estimations of 26 woody tree
species in India were reported (Table 1). Multiple biomass equations for
Azadirachta indica, Casuarina equisetifolia, Hevea brasiliensis, and Tectona
grandis were found in the literature. The minimum dataset for equation
development significantly affects model accuracy. Therefore, it is rec-
ommended to have more than forty harvested samples for AGB model
development of woody species (Sileshi, 2014). The harvested data fre-
quency graph showed about 60% of studies reported in India had used
less than 40 numbers of harvested biomass data in equation develop-
ment (Fig. 2).
A total of 13 studies used 40 to 180 harvested biomass data in AGB
equation development. Around 460 to 470 trees have been harvested
from ten different species to develop an AGB estimation equation for
saplings to the juvenile stage (Chaturvedi et al., 2012; 2013). AGB equa-
tion developed (Nath et al., 2019) for the tropical to sub-tropical forests
were associated with at least twenty-three different woody trees. The
dataset consists of 303 trees from four different types of forests.
3.1.1. Bole biomass estimation
Tree bole contributed the highest share to the tree biomass, which
ranged from 34 to 56% of the AGB (Nath et al., 2017; Naik et al., 2018;
Singh et al., 2011; Swamy et al., 2004) and 30 to 53% of the total tree
biomass (Naik et al., 2018; Newaj et al., 2016). For estimation of bole
or stem biomass, different independent variables such as D, (collar di-
ameter) CD, circumference at breast height (CBH), number of branches
(NB), crown width (CrW), plant height (H) have been widely used by
the researchers (Table 3). In India, 85 and 6 species-specific equations
have been reported for tree bole and bamboo culm biomass estimation.
At the same time, three equations for bole biomass estimation for mixed
stands were found in the literature.
Power-law equations based on D of the 112 culms were developed
for 1 to 4 yr old Dendrocalamus strictus. Incorporating H as a co-variable
with D has been also used to develop the age-specific equations for 1 to
>3 yr old Bambusa arundinacea. Power-law equations can well describe
the growth of the trees and biomass changes in them (Sileshi, 2014).
About 90% of India’s total recommended equations were based on
Power-law form. The minimum dataset used for the equation devel-
opment showed that approximately 86% of India’s total studies had
used less than forty trees. However, only two studies were found in
the Power-law form developed using more than forty harvested trees
(Nath et al., 2017; Swamy et al., 2004). Since the standing tree D is eas-
ily measurable, the equations recommended by Nath et al. (2017) and
Swamy et al. (2004) are most suitable for the bole biomass (BB) es-
timation of Barringtonia acutangula and Gmelina arborea, respectively.
Three studies used the Power-law equations to estimate biomass in
mixed woody species of India (Rana et al., 1989; Chaturvedi et al., 2012;
Chaturvedi and Raghubanshi, 2013). However, these studies either con-
sidered the juvenile or small woody diameter trees or evaluated mini-
mum harvest data for biomass equation development.
3.1.2. Branch biomass estimation
Tree branches contribut 3‒20% of the AGB stocks (Chaturvedi and
Raghubanshi, 2015; Verma et al., 2014; Jha, 2015; Swamy et al., 2004;
Lodhiyal and Lodhiyal, 2003; Lodhiyal et al., 1995). Different biomass
equations for various tree species have been developed by several re-
searchers for branch biomass estimation under 1 to 128 yr old planta-
tions in India (Table 3).
Four species-species regression equations for bamboo and 75 species-
specific equations for branch biomass estimation of woody species were
found published from India (Table 3). The age-specific Power-law equa-
tions for 1 to >3 yr old bamboos (Bambusa arundinacea) have developed
using H as a co-variable with D. Results of this study were based on lim-
ited number of harvest data. Such equations can be used in future ref-
erences after multi-fold validation processes. We found that, of the 78
studies reported in India, 68 recommended Power-law equation forms,
and about 90% of these equations are based on either D or GBH of the
tree. Similarly, a few studies have used CD as a simple variable and D
and tree height (H) as a compound variable to estimate branch biomass.

6
B. Brahma, A.J. Nath, C. Deb et al. Trees, Forests and People 5 (2021) 100098

Table 3
Details of the regression equations for branch biomass estimation for diverse lifeforms in India.

Age of the Harvested sample

Associated Species Equations stand/tree (yr) size (number) Sources

Bamboos
Bambusa arundinacea BRB= 4.310 + 17.690D2 H 1 25 Chandrashekara, 1996
Bambusa arundinacea BRB= 4.530 + 33.510D2 H 2 25 Chandrashekara, 1996
Bambusa arundinacea BRB= 4.099 + 66.23D2 H 3 25 Chandrashekara, 1996
Bambusa arundinacea BRB= 4.068 + 36.608D2 H >3 25 Chandrashekara, 1996
Trees

Acacia catechu BRB= −0.2451+0.1958D 5 25 Singh et al., 2011

Acacia nilotica BRB= −1.931+0.816 D + 0.164 D2 4.5 to 6.5 75 Rizvi and Ahlawat, 2014
Acacia nilotica BRB= −10.539 + 3.397 lnD 4 20 Singh and Toky, 1995
Acacia nilotica BRB=−17.246 + 4.608 ln D 8 20 Singh and Toky, 1995
Albizia lebbeck BRB=−0.1617+0.1561D 5 25 Singh et al., 2011
Albizia procera BRB=2.12 D2.690 11 32 Newaj et al., 2016
Alnus nepalensis lnBRB =1.455 + 2.216 ln DBH 7 to 56 23 Sharma and Ambasht, 1991
Alnus nepalensis lnBRB = −4.396 + 0.711 lnAge Singh and Yadava, 1994
Alnus nepalensis lnBRB = −6.941 + 1.214 lnD Singh and Yadava, 1994
Anogeissus pendula BRB= −2.133 + 0.6234 BCB 20 Vyas et al., 1978
Barringtonia acutangula BRB= 0.0002D3.629 200 Nath et al., 2017
Casuarina equisetifolia BRB= −4.574+2.7803lnD 2 to 9 90 Vidyasagaran and Paramathma, 2014
Citrus sinensis BRB= (exp(−1.0 + 0.45(lnD))) × 1.08 8 to 30 58 Sahoo et al., 2020
Cinnamomum camphora lnBRB =1.453 + 0.451 lnAge Singh and Yadava, 1994
Cinnamomum camphora lnBRB =0.322 + 0.734 lnD Singh and Yadava, 1994
Dalbergia sissoo BRB= −1.4581+0.7708D 5 25 Singh et al., 2011
Dalbergia sissoo BRB=−0.5876+0.5369D 5 12 Lodhiyal and Lodhiyal, 2003
Dalbergia sissoo BRB=−2.2074+0.7201D 10 12 Lodhiyal and Lodhiyal, 2003
Dalbergia sissoo BRB=4.7404+0.6164D 15 12 Lodhiyal and Lodhiyal, 2003
Dalbergia sissoo BRB=0.026 D2.332 17 42 Newaj et al., 2016
Dendrocalamus strictus BRB=0.00031D4.4420 1 to 4 112 Kaushal et al., 2016
Embilica officinalis BRB=4.262 CD0.941 15 30 Newaj et al., 2016
Embilica officinalis BRB= 0.0007D3.35108 10 Panwar et al., 2014
Eucalyuptus tereticornis BRB=−0.887 + 0.469 ln D 4 20 Singh and Toky, 1995
Eucalyuptus tereticornis BRB=−4.924 + 1.254 ln D 8 20 Singh and Toky, 1995
Gaultheria griffithiana lnBRB = −1.430 + 0.189 lnAge Singh and Yadava, 1994
Gaultheria griffithiana lnBRB = −1.982 + 0.477lnD Singh and Yadava, 1994
Gmelina arborea BRB=0.0102+2.4202 expDBH 1 to 6 75 Swamy et al., 2004
Grewia optiva BRB=0.109D1.523 4 to 23 25 Verma et al., 2014
Hardwickia binata BRB=0.002D3.142 20 30 Newaj et al., 2016
Hevea brasiliensis ln BRB=(exp (2.76+2.07(lnD) × 1.20 6 to 34 67 Brahma et al., 2017
Jatropha curcus BRB= 0.015CD2.063 8 16 Rao et al., 2016
Leucaena leucocephala BRB=−2.293 + 1.126 ln D 4 20 Singh and Toky, 1995
Leucaena leucocephala BRB= −24.459 + 6.272 lnD 8 20 Singh and Toky, 1995
Litchi chinensis BRB=(10.09/[1+(−0.37)exp(−0.35 × CD)]ˆ(1/−0.09)+E 2 to10 30 Naik et al., 2018
Lyonia ovalifolia lnBRB = −2.090 + 0.485 lnAge Singh and Yadava, 1994
Lyonia ovalifolia lnBRB = −5.166 + 1.054 lnD Singh and Yadava, 1994
Mallotus philippensis BRB= −2.28 + 1.22 lnGBH 9 Rana et al., 1989
Neolitsea zeylanica lnBRB =0.206 + 0.383 lnAge Singh and Yadava, 1994
Neolitsea zeylanica lnBRB =0.053 + 0.694 lnD Singh and Yadava, 1994
Pinus kesiya BRB= 0.6212–3.9659D + 6.0668D2 −1.9087D3 40 Baishya and Barik, 2011
Pinus patula BRB= 0.019+0.286D 15 12 Bargali and Singh, 1997
Pinus patula BRB= 0.172+0.278D 18 12 Bargali and Singh, 1997
Pinus patula BRB= −0.099+0.289D 21 12 Bargali and Singh, 1997
Pinus roxburghii BRB=−9.83 + 2.98 lnGBH 16 to 128 26 Chaturvedi and Singh, 1982
Populus deltoides BRB=−0.0935+1.0234D 5 27 Lodhiyal et al., 1995
Populus deltoides BRB=−1.6149+1.1345D 6 27 Lodhiyal et al., 1995
Populus deltoides BRB=−1.0016+1.3047D 7 27 Lodhiyal et al., 1995
Populus deltoides BRB=−5.9209+1.5531D 8 27 Lodhiyal et al., 1995
Psidium guajava BRB=0.009CD2.665 4 to 14 17 Rathore et al., 2018
Pyrularia edulis lnBRB = −7.957 + 1.008 lnAge Singh and Yadava, 1994
Pyrularia edulis lnBRB = −10.554 + 1.760 lnD Singh and Yadava, 1994
Quercus leucotrichophora ln BRB= −0.718 + 1.302 lnGBH 23 Rawat and Singh, 1988
Quercus floribunda ln BRB= −0.987 + 1.377 lnGBH 17 Rawat and Singh, 1988
Quercus dealbata lnBRB = 1.449 + 0.287 lnAge Singh and Yadava, 1994
Quercus dealbata lnBRB = −1.714 + 0.711 lnD Singh and Yadava, 1994
Quercus fenestrata lnBRB = −1.301 + 0.444 lnAge Singh and Yadava, 1994
Quercus fenestrata lnBRB = −2.577 + 0.910 lnD Singh and Yadava, 1994
Quercus griffithii lnBRB = −6.705 + 0.750 lnAge Singh and Yadava, 1994
Quercus griffithii lnBRB = −9.592 + 1.568 lnD Singh and Yadava, 1994
Quercus griffithii lnBRB = −0.463 + 0.482 lnAge Singh and Yadava, 1994
Rhododendron arboreum lnBRB = −3.780 + 0.752 lnAge Singh and Yadava, 1994
Rhododendron arboreum lnBRB = −13.226 + 1.687 lnD Singh and Yadava, 1994
Rhododendron arboreum ln BRB= 1.113 + 0.609 lnGBH 8 Rawat and Singh, 1988
Shorea robusta BRB= −2.04 + 1.50 lnGBH 9 Rana et al., 1989
Styrax serrulatus lnBRB = −2.905 + 0.788 lnAge Singh and Yadava, 1994
(continued on next page)

7
B. Brahma, A.J. Nath, C. Deb et al. Trees, Forests and People 5 (2021) 100098

Table 3 (continued)

Associated Species Equations Age of the Harvested sample Sources

stand/tree (yr) size (number)

Styrax serrulatus lnBRB = −7.325 + 1.351 lnD Singh and Yadava, 1994
Tectona grandis BRB= 0. 0718D2.058 20 to 47 Buvaneswaran et al., 2006
Tectona grandis BRwB= 0.001D3.063 20 to 47 Buvaneswaran et al., 2006
Tectona grandis BRB= 0.0122D2.523 20 to 47 Buvaneswaran et al., 2006
Tectona grandis BRwB= 0.001D3.0634 20 to 47 Buvaneswaran et al., 2006
Tectona grandis lnBRB=6.726+6.000/1+(lnpD2 H/12.89)−13.27 100 Chaturvedi and Raghubanshi, 2015
Tectona grandis BRB= 0.0678D2 –0.7045D+1.5725 1 to 30 33 Jha, 2015
Tectona grandis BRB= 0.156+144.89D2 H 15 to 20 12 Chandrashekara, 1996
Terminalia chebula BRB= 0.0103D2.5388 10 Panwar et al., 2014
Mixed woody species ln BRB = −0.838+2.300 lnD2 460 Chaturvedi and Raghubanshi, 2013
Mixed woody species ln BRB=−0.91 + 1.33 lnGBH 48 Rawat and Singh, 1988
Mixed woody species BRB= −5.21 + 2.08 lnGBH 18 Rana et al., 1989

Note: BRB = Branch biomass, BRwB= Branch wood biomass, CD = Collar Diameter, CBH= Circumference at breast height of the tree, CrW=Crown width, CrD=
Crown diameter, D=Diameter at breast height of tree/Culm (DBH), 𝜌= Wood Density, H= Height of Tree/Clum, E = Model error.

For branch biomass estimation, only four studies were reported to have biomass based on D and GBH for mixed woody species used the Power-
used the Power-law form based on D using more than forty harvest law equation (Rana et al., 1989; Chaturvedi et al., 2012; Chaturvedi and
trees, the minimum dataset recommended for biomass equation devel- Raghubanshi, 2013). However, these studies considered a lower number
opment (Sileshi, 2014). Another three studies that also estimated branch of harvested data set, and used either juvenile or small-diameter trees.

Fig. 2. The number of studies and the number of sample trees harvested for the biomass equation development.

8
B. Brahma, A.J. Nath, C. Deb et al.
3.1.3. Leaf biomass estimation
A total of five biomass equations for bamboos of India have been
reported. Age-specific equations for 1 to >3 yr old Bambusa arundinacea
stands were developed based on the D and H of the culm. Based on D of
112 culms of Dendrocalamus strictus stands of 1 to 4 yr old, a Power-
law regression equation has been reported by Kaushal et al. (2016).
Fifty-nine species-specific Power-law equations have been reported for
twenty-nine different woody tree species of India (Table 4).
The use of regression equations depends on the variables and their
accessibility in the field (Li et al., 2020). Recognizing or measuring the
age, the number of the branch (NB), crown width (CrW), and crown di-
ameter (CrD) of the standing tree can be predictable to some extent that
need sufficient and careful technological efforts. Therefore, the use of
age, NB and CrW, and CrD as predictive variables used in biomass re-
gression equations for leaf may lead to inaccurate assessment. However,
the variables such as D and GBH of the standing trees are easily measur-
able with better precision in the field. These are mostly recommended
to use in non-destructive biomass estimation. Leaf biomass equations
for thirteen tree species viz. Alnus nepalensis, Barringtonia acutangula,
Cinnamomum camphora, Dalberia sisso, Gaultheria griffithiana, Lyonia
ovalifolia, Neolitsea zeylanicus, Pyrularia edulis, Quercus fenestrata,
Q. griffithii, Rhododendron arboreum, Styrax serrulatus and Tectona
grandis (Table 4) have been developed using D and GBH as they were
found to be better predictors for leaf biomass estimation. Similarly,
some efforts have been made to estimate leaf biomass equation for In-
dia’s mixed woody species stand that are either associated with juvenile
trees (Chaturvedi et al., 2012) or small-diameter trees (Chaturvedi and
Raghubanshi, 2013).
3.2. BGB Estimation
BGB accounting for about 20% (Pearson et al., 2017) to 26%
(Cairns et al., 1997) of the total biomass, and is linked to aboveground
biomass dynamics, is a significant carbon pool for many vegetation types
and land-use system. A well-built fraction of the root biomass occurs on
the top 30 cm of the soil surface. Since the BGB contributes significantly
to the carbon pool, an accurate estimation of this pool can contribute
considerably to carbon mitigation and other land-based projects. How-
ever, the assessment of BGB is very tedious and time-consuming. The
loss of roots during excavation and segregation from the soil is an im-
portant issue associated with BGB estimation accuracy. Although the
default ratio of BGB values to the AGB stocks for species and region is
available, the empirical studies preferred species-specific estimation of
BGB for precise reporting on the biomass stock. The BGB of the tree con-
sists of stump roots biomass (SRB), lateral root biomass (LRB), and fine
root biomass (FRB). Incorporating SRB, LRB, and FRB together is also
known as total root biomass (TRB). The TRB also encompasses the FRB,
coarse root biomass (CRB), and taproot (TaRB).
3.2.1. Stump root biomass estimation
Stump root, also known as the primary root, is the belowground por-
tion of the tree’s stump or stem developed as a root under the soil. Stump
root contributed a significant share to the BGB, or root biomass and
ranged from 23‒60% of the total BGB (Jha, 2015; Singh et al., 2011; Das
et al., 2020) and 2‒10% to the total biomass (Jha, 2015; Singh et al.,
2011; Lodhiyal et al., 1995). The literature review found only nine stud-
ies in India that developed nineteen species-specific regression equa-
tions for SRB estimation for ten different tree species (Table 5). These
studies suggest that SRB of the tree can be best described by either D
or GBH of the tree. Among these, 95% of the studies used the Power
function that suggests its reliability in SRB estimation. However, the as-
sociated strength of the dataset in biomass equation development in all
these studies is lower than the recommended number (Sileshi, 2014).
A single study was found on SRB estimation equation for mixed woody
species of India (Rawat and Singh, 1988). These findings reveal that
9
Trees, Forests and People 5 (2021) 100098
the Power-law function based on GBH of the tree can be used in future
references after the multi-fold validation process.
3.2.2. Lateral root biomass estimation
The roots developed horizontally from the primary or stump root are
known as the lateral root, also called as secondary roots. Lateral roots
contribute around 3‒9% of the total tree biomass, whereas its contribu-
tion ranged from 14‒42% to the BGB (Verma et al., 2014; Das et al.,
2011; Jha, 2015; Singh et al., 2011). The present review found twenty
species-specific equations for LRB estimation of eleven different tree
species (Table 5). The findings reveal that the use of D or GBH as a pre-
dictive variable in the Power-law function is the best predictors for LRB
of trees. The LRB estimation regression equation for the mixed woody
species suggested by Rawat and Singh (Rawat and Singh, 1988) can
also be used. However, a multi-fold prior validation of this equation is
strongly recommended before its use.
3.2.3. Fine root biomass estimation
The diameter of the fine roots (FRs) has not been defined uniformly,
and it may range from <2 mm to <5 mm (Cairns et al., 1997; Joslin and
Henderson, 1987; Vogt et al., 1991). However, the size of FRs less than
1 mm diameter has a higher contribution to the total FRs biomass than
the >1 mm diameter roots (Cairns et al., 1997). Therefore, an accurate
estimation of fine root biomass (FRB) in biomass/ecological research
is needed to adequately describe the methodology and efforts. Twenty
species-specific equations for FRB estimation have been reported for ten
different tree species in India. D and GBH of the tree were used as a pre-
dictive variable for 95% and 5% of India’s published research articles.
FRB estimation equation for mixed species in India, however, is scarce.
Rawat and Singh (1988) have recommended the Power function based
on GBH for FRB estimation from forty-eight harvested trees.
3.2.4. Coarse root biomass estimation
The definition of coarse root signifies that the root with diameter of
more than 5 mm is a ‘coarse root.’ The biomass estimation of coarse roots
associated with the extraction, identification, segregation and cleaning
requires sufficient effort and technological support. Therefore, studies
related to root biomass estimation are comparatively lower than the
aboveground parts of the tree. In Indian literature, three seperate coarse
root estimation equations are available for Hevea brasiliensis and Pinus
kesiya (Table 5). An exponential form was developed based on the AGB
from sixty-seven harvested H. brasiliensis trees under 6 to 34 yr old trees
(Brahma et al., 2017). Additionally, another regression equation using D
and H as a compound variable was used in the same study (Brahma et al.,
2017). A linear form of regression equation for Pinus kesiya was also
developed using D from forty harvested trees (Baishya and Barik, 2011).
3.2.5. Tap root biomass estimation
Tap root is the below-ground part of tree bole. A single study car-
ried out for tap root (TaRB) biomass estimation using 4–23 years old
stands with 25 harvested trees of Grewia optiva (Verma et al., 2014).
For estimation of TaRB this study used Power law function based on D.
3.2.6. Total root biomass or belowground biomass estimation
Total root biomass is also known as the belowground biomass com-
prised of SRB, LRB, FRB, CRB and TaRB of the tree. Sixteen differ-
ent studies from Indian literature reported species-specific total root
biomass (TRB) or BGB using regression equations for 18 different species
(Table 5). Tree D is the most recommended variable used for predicting
the biomass of different plant parts (Liu et al., 2017). It is also consis-
tent in India, where nearly 50% of the reported studies have used D as
a predictive variable. At the same time, CD and GBH were the second
and third preferred variables for TRB or BGB estimation, respectively.
The future use of the recommended equations depends on the ease of
equation use, the quality of the equation, and the accessibility of the
predictive variable. The measurement of variables such as H, BaD, CrH,
B. Brahma, A.J. Nath, C. Deb et al. Trees, Forests and People 5 (2021) 100098

Table 4
Details of the regression equations for leaf biomass estimation of bamboo, banana and tree.

Stand/tree Harvested sample

Associated species Equations age (yr) size (number) Sources

Bamboos
Bambusa arundinacea LFB= 1.578 + 5.787D2 H 1 25 Chandrashekara, 1996
Bambusa arundinacea LFB=33.876D2 H - 0.6154 2 25 Chandrashekara, 1996
Bambusa arundinacea LFB=2.102 + 5.601D2H 3 25 Chandrashekara, 1996
Bambusa arundinacea LFB=2.302 + 6.545D2 H >3 25 Chandrashekara, 1996
Dendrocalamus strictus LFB= 0.1902D0.4587 1 to 4 112 Kaushal et al., 2016
Bananas
Musa balbisiana 140 Laskar et al., 2020
LFB=(exp(−7.27+1.12(lnD2Hp))
x 1.23
Trees
Acacia catechu LFB= 0.1098+0.2194D 25 Singh et al., 2011
Albizia lebbeck LFB=−0.1247+0.1853D 25 Singh et al., 2011
Albizia procera LFB= 0.025 D2.237 8 32 Newaj et al., 2016
Alnus nepalensis lnLFB = −4.955 + 0.626 lnAge 7 to 33 47 Singh and Yadava, 1994
Alnus nepalensis lnLFB = −6.165+ 1.085 ln D 7 to 33 47 Singh and Yadava, 1994
Barringtonia acutangula LFB= 0.0013D2.412 200 Nath et al., 2017
Cinnamomum camphora lnLFB =−0.678+ 0.391 ln Age 7 to 33 68 Singh and Yadava, 1994
Cinnamomum camphora lnLFB = −1.871 + 0.654 lnD 7 to 33 47 Singh and Yadava, 1994
Dalbergia sissoo LFB=−0.3337+0.3052D 5 12 Lodhiyal and Lodhiyal, 2003
Dalbergia sissoo LFB=−1.1037+0.3600D 10 12 Lodhiyal and Lodhiyal, 2003
Dalbergia sissoo LFB= 1.7329+0.2238D 15 12 Lodhiyal and Lodhiyal, 2003
Dalbergia sissoo LFB= −0.4501+0.283D 25 Singh et al., 2011
Dalbergia sissoo LFB= 0.041 D 1.845 14 42 Newaj et al., 2016
Embilica officinalis LFB= 0.152 CD1.605 12 30 Newaj et al., 2016
Gaultheria griffithiana lnLFB = −0.955 + 0.122 ln Age 7 to 33 68 Singh and Yadava, 1994
Gaultheria griffithiana lnLFB = −1.330 + 0.310 ln D 7 to 33 47 Singh and Yadava, 1994
Gmelina arborea LFB= 0.0154+ 1.8832D 1 to 6 75 Swamy et al., 2004
Grewia optiva LFB=0.067D1.435 4 to 23 25 Verma et al., 2014
Grewia optiva LFB=0.165D1.357 4 to 23 25 Verma et al., 2014
Hardwickia binata LFB= 0.002 D3.514 17 30 Newaj et al., 2016
Hevea brasiliensis LFB= (exp(−4.41+1.87(lnD)) 6 to 34 67 Brahma et al., 2017
1.25
Jatropha curcus LFB= 0.194 CD0.715 8 16 Rao et al., 2016
Jatropha curcus LFB=0.0158H0.437 8 16 Rao et al., 2016
Jatropha curcus. LFB=0.154NB0.445 8 16 Rao et al., 2016
Jatropha curcus LFB=0.0035CrW1.062 8 16 Rao et al., 2016
Jatropha curcus LFB=1.68E-05 CrD1.655 8 16 Rao et al., 2016
Litchi chinensis LFB = 7.25/ [1 + 0.07 exp 2 to 10 30 Naik et al., 2018
(−0.29 × CD)] ˆ(1/0.02)
+0.418)
Lyonia ovalifolia lnLFB = −3.413 + 0.486 lnAge 7 to 33 47 Singh and Yadava, 1994
Lyonia ovalifolia lnLFB = −6.165 + 1.017ln D 7 to 33 47 Singh and Yadava, 1994
Mallotus philippensis LFB= −3.86+ 1.07lnGBH 9 Rana et al., 1989
Neolitsea zeylanics lnLFB = 1.037 + 0.331 ln Age 7 to 33 68 Singh and Yadava, 1994
Neolitsea zeylanics lnLFB= −1.221 + 0.605 lnD 7 to 33 47 Singh and Yadava, 1994
Pinus patula LFB= −1.120+0.295D’ 15 12 Bargali and Singh, 1997
Pinus patula LFB= 0.365+0.215D’ 18 12 Bargali and Singh, 1997
Pinus patula LFB= 0.063+0.214D’ 21 12 Bargali and Singh, 1997
Pinus roxburghii LFB= −6.11 + 1.87 lnGBH 16 to 128 26 Chaturvedi and Singh, 1982
Populus deltoides LFB= 0.19D1.06 1 to 9 27 Das et al., 2011
Populus deltoides LFB=1.8998+0.7165D 5 27 Lodhiyal et al., 1995
Populus deltoides LFB=−0.6788+0.7978D 6 27 Lodhiyal et al., 1995
Populus deltoides LFB=−0.9837+0.8814D 7 27 Lodhiyal et al., 1995
Populus deltoides LFB=−2.2119+0.9967D 8 27 Lodhiyal et al., 1995
Psidium guajava LFB=0.033CD2.197 4 to 14 17 Rathore et al., 2018
Pyrularia edulis lnLFB = −5.572 + 0.680 ln Age 7 to 33 68 Singh and Yadava, 1994
Pyrularia edulis lnLFB = −7.136 + 1.169 ln D 7 to 33 47 Singh and Yadava, 1994
Quercus floribunda LFB= −1.23+ 1.38lnGBH 17 Rawat and Singh, 1988
Quercus leucotrichophora LFB= −0.98+ 0.85 lnGBH 23 Rawat and Singh, 1988
Quercus dealbata lnLFB = −2.253 + 0.380 lnAge 7 to 33 68 Singh and Yadava, 1994
Quercus fenestrata lnLFB = −0.061 + 0.194 ln Age 7 to 33 68 Singh and Yadava, 1994
Quercus fenestrata lnLFB=−1.069 + 0.386 lnD 7 to 33 47 Singh and Yadava, 1994
Quercus griffithii lnLFB = −6.289+ 0.580 lnAge 7 to 33 68 Singh and Yadava, 1994
Quercus griffithii lnLFB = −2.773 + 0.485ln Age 7 to 33 47 Singh and Yadava, 1994
Quercus griffithii lnLFB = −8.517 + 1.999 lnD 7 to 33 47 Singh and Yadava, 1994
Quercus griffithii lnLFB = −6.459+ 0.984 ln D 7 to 33 47 Singh and Yadava, 1994
Rhododendron arboreum lnLFB = −2.850 + 0.397 ln Age 7 to 33 47 Singh and Yadava, 1994
Rhododendron arboreum lnLFB = −7.860+ 0.892 ln D 7 to 33 47 Singh and Yadava, 1994
Rhododendron arboreum LFB= 1.19+ 0.17 lnGBH 8 Rawat and Singh, 1988
Shorea robusta LFB= −1.74+ 1.18 lnGBH 9 Rana et al., 1989
(continued on next page)

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B. Brahma, A.J. Nath, C. Deb et al. Trees, Forests and People 5 (2021) 100098

Table 4 (continued)

Associated species Equations Stand/tree Harvested sample Sources

age (yr) size (number)

Styrax serrulatus lnLFB = −2.827 + 0.578 lnAge 7 to 33 68 Singh and Yadava, 1994
Styrax serrulatus lnLFB = −6.148 + 0.995 ln D 7 to 33 47 Singh and Yadava, 1994
2.459
Tectona grandis LFB= 0.0037D 20 to 47 – Buvaneswaran et al., 2006
2.1524
Tectona grandis LFB= 0.0116D 20 to 47 – Buvaneswaran et al., 2006
Tectona grandis ln LFB= 2.985+1.029lnD2 100 Chaturvedi and Raghubanshi, 2015
Tectona grandis ln LFB= 6.356+7.280/1+(lnD2 /6.682)−4.706 100 Chaturvedi and Raghubanshi, 2015
Tectona grandis LFB=−0.0025D2 +0.4833D- 1 to 30 33 Jha, 2015
2.3174
Tectona grandis LFB= 74.0D2 H−2.72 15 to 20 12 Chandrashekara, 1996
Mixed woody species lnLFB=−0.687+1.937 lnD2 460 Chaturvedi and Raghubanshi, 2013
Mixed woody species LFB= −4.96+ 1.68 lnGBH 9 Rana et al., 1989
Mixed woody species LFB= −1.11 + 1.04 lnGBH 9 Rawat and Singh, 1988

Note: D’ = Diameter in meter, GBH= Girth at breast height, CB= Culm Biomass, BB= Bole Biomass, BWB= Bole wood biomass, PsS= Pseudo stem, p= wood density,
D= Diameter at breast height of the tree/culm (DBH), CBH= Circumference at breast height of the tree, H= Tree/culm height, CD= Collar diameter of the tree,
𝜌=wood density, CrW=Crown width, CrD= Crown diameter and NB=Number of primary branches.

and CrW generally requires technological support and significant efforts. Equation 1: lnAGB= 3.428 + 0.310 (ln pD2 H) (Chaturvedi et al.,
Therefore, using of such equations associated with H, BaD, CrH, and CrW 2012)
are least recommended for future use. Only four biomass equations for Equation 2: lnABG= −3.206+1.337(ln pD2 H) (Chaturvedi and
Raghubanshi, 2013)
TRB and BGB have been found in the present review. It used data set
Equation 3: AGB=(0.18D2.16 ) × 1.32 (Nath et al., 2019)
of more than 40 harvested trees as suggested for the minimum required Equation 4: AGB= (0.32pD2 H0.75 ) × 1.34 (Nath et al., 2019)
sample size for an efficient biomass equation (Sileshi, 2014). Equation 5: ln AGB= 0.349 + 1.316 ln GBH (Rawat and
Singh, 1988)

All the mixed-species equations (Equation 1, 2, and 4) were based

3.3. Total Biomass estimation
The total biomass of a tree is the sum of AGB and BGB. Indepen-
dent measuring or predicting of the TB may be associated with vari-
ous methodological issues resulting in errors. Predicting TB through a
species-specific equation is a viable option for overcoming methodolog-
ical perturbations with significant precisions. Numerous attempts have
been made to develop TB equations for India’s woody and non-woody
plants. The number of harvested trees has a profound effect on the pre-
cision levels of recommended biomass equations. Among the reported
biomass estimation studies, as many as fifteen studies were found using
harvest tree component data for developing the species-specific equa-
tions (Table 6).
Typically, biomass measurements are performed on the bole,
branches, leaves, and roots, and total biomass is considered as the sum
of these components. Some authors argue that the independent fitting
of equations for the different components is not satisfactory for total
biomass estimation (Sanquetta et al., 2015). The use of simultaneous
equations has recently become a common practice in estimating TB
because it ensures compatibility among predictors of total and compo-
nents of biomass. Fifteen studies have been reported in India to develop
species-specific TB equations for sixteen different tree species (Table 6).
It was found that 48% of the equations used D as the predictor variable,
and Power-law function described over 70% of the relationships. The
use of D over the CD is highly recommended, as the measurement of
the latter is associated with methodological difficulties due to slope po-
sition, and buttresses. The present review reveals that only six studies
have used a minimum of 40 harvested trees for TB equation develop-
ment (Table 6).
3.4. Cross Validation
on the compound variables of 𝜌, D and H. In contrast, Equation 3 and 5
were based on D or GBH as the simple variable.
The test included harvested dataset of 23 samples for T. grandis
(Deb et al., 2016; Sandeep et al., 2015; Chaturvedi and Raghuban-
shi, 2015; Jain and Ansari, 2013), 30 for C. Fistula (Tripathi, 2015),
and 49 for P. kesiya (Baishya and Barik, 2011; Das and Ramakrish-
nan, 1987). The D values for the individual tree species in the equa-
tion test dataset ranged from 0.59 to 48.09 cm (for T. grandis), 3.82 to
30.5 cm (for C. fistula), and 0.33 to 62. 69 cm (for P. kesiya) (Table 1).
Tree height ranged from 1.6 to 25.35 m for T. grandis and 1.8 to 11.2 m
for C. fistula (Table 1), and the collective 𝜌 values for T. grandis and C.
fistula ranged from 0.588 to 0.76 g cm−3 . However, the H and 𝜌 val-
ues for P. Kesiya were not available in the equation test dataset. There-
fore, the RMSE values of equations 1, 2 and 4 were estimated from the
test dataset consisting of 53 and 102 sample trees and for equation 3,
and equation 5 respectively. The average RMSE values were calculated
and summarized in Table 7. However, the equation recommended by
Rana et al. (1989) has not been incorporated in the data validation, as
the equation did not pass the primary screening of the minimum harvest
data requirement.
The RMSE value was lowest for equation 5, followed by equation
3 (218.83) and equation 1 (399.13), respectively. The lowest RMSE
value signifies the best equation (equation 5) among all the five rec-
ommended equations available in the literature. The results revealed
the equation consisting of compound variables (D, 𝜌, and H) are less
empiric in biomass estimation than the single power-law model based
on GBH and D.
3.5. General observations
In the following sections, the general observations that apply to
aboveground, belowground, and total biomass estimation have been dis-
cussed. These fall under three main issues: choice of variables, equation
form, and sample size.

The following five-biomass equations were reported for mixed- 3.5.1. Choice of variables used in biomass equations
species stand biomass estimation in India. The equations were thor- In the various equations that we examined, D has been identified
oughly cross validated with the secondary dataset. as the most reliable variable for biomass prediction. Whereas, CD is

11
B. Brahma, A.J. Nath, C. Deb et al. Trees, Forests and People 5 (2021) 100098

Table 5
Details of the recommended regression equations for belowground biomass estimation.

Stand/ Tree Harvested sample

Associated Species Equations age (yr) size (number) Sources

Bananas
Musa balbisiana Corm=(exp(−3.93+0.715(lnD2 Hp)) x 1.06 149 Laskar et al., 2020

Stump Roots
Acacia catechu SRB=0.081+0.1234D 5 25 Singh et al., 2011
Albizia lebbeck SRB=−0.0162+0.0707D 5 15 Singh et al., 2011
Dalbergia sissoo SRB= −0.6128+0.6231D 5 12 Lodhiyal and Lodhiyal, 2003
Dalbergia sissoo SRB=−2.1194+0.6914D 10 12 Lodhiyal and Lodhiyal, 2003
Dalbergia sissoo SRB=3.4393+0.4626D 15 12 Lodhiyal and Lodhiyal, 2003
Dalbergia sissoo SRB=- 4017+0.2744D 5 25 Singh et al., 2011
Pinus patula SRB= −0.053+0.227D 15 12 Bargali and Singh, 1997
Pinus patula SRB= 0.005+0.225D 18 12 Bargali and Singh, 1997
Pinus patula SRB= −0.845+0.258D 21 12 Bargali and Singh, 1997
Pinus roxburghii SRB= −7.22+2.45 lnGBH 16 to 128 26 Chaturvedi and Singh, 1982
Populus deltoides SRB= 0.23D1.14 1 to 9 27 Das et al., 2011
Populus deltoides SRB=1.7122+0.9519D 5 27 Lodhiyal et al., 1995
Populus deltoides SRB=−0.3404+1.1317D 6 27 Lodhiyal et al., 1995
Populus deltoides SRB=−0.9145+1.4625D 7 27 Lodhiyal et al., 1995
Populus deltoides SRB=−3.7802+1.88910D 8 27 Lodhiyal et al., 1995
Tectona grandis SRB=0.0674D2 –0.8079D+3.7722 1 to 30 33 Jha, 2015
Quercus leucotrichophora SRB= −0.98 + 0.90 lnGBH 23 Rawat and Singh, 1988
Quercus floribunda SRB= −0.25 + 1.11 lnGBH 17 Rawat and Singh, 1988
Rhododendron arboreum SRB= −0.12 + 0.87 lnGBH 8 Rawat and Singh, 1988
Mixed woody species SRB= −0.10+0.95 lnGBH 48 Rawat and Singh, 1988

Lateral roots

Acacia catechu LRB=−0.4909+0.2123D 5 25 Singh et al., 2011

Albizia lebbeck LRB= −0.0614+0.061D 5 15 Singh et al., 2011
Dalbergia sissoo LRB=−0.3962+0.3617D 5 12 Lodhiyal and Lodhiyal, 2003
Dalbergia sissoo LRB=−1.4529+0.4750D 10 12 Lodhiyal and Lodhiyal, 2003
Dalbergia sissoo LRB=2.8214+0.3849D 15 12 Lodhiyal and Lodhiyal, 2003
Dalbergia sissoo LRB= −2723+0.1923D 5 25 Singh et al., 2011
Grewia optiva LRB=0.078D1.786 4 to 23 25 Verma et al., 2014
Pinus patula LRB= −0.225+0.122D 15 12 Bargali and Singh, 1997
Pinus patula LRB= 0.048+0.110D 18 12 Bargali and Singh, 1997
Pinus patula LRB= −0.035+0.114D 21 12 Bargali and Singh, 1997
Pinus roxburghii LRB= −9.16 + 2.59 lnGBH 16 to 128 26 Chaturvedi and Singh, 1982
Populus deltoides LRB= 0.01D1.77 1 to 9 27 Das et al., 2011
Populus deltoides LRB=1.6898+0.4439D 5 27 Lodhiyal et al., 1995
Populus deltoides LRB=−1.1379+0.6466D 6 27 Lodhiyal et al., 1995
Populus deltoides LRB=−1.3171+0.9099D 7 27 Lodhiyal et al., 1995
Populus deltoides LRB=−3.3974+1.0080D 8 27 Lodhiyal et al., 1995
Tectona grandis LRB=0.0583D2 –1.0494D+5.4397 1 to 30 33 Jha, 2015
Quercus leucotrichophora LRB= −0.31 + 0.81 lnGBH 23 Rawat and Singh, 1988
Quercus floribunda LRB= −1.59 + 1.00lnGBH 17 Rawat and Singh, 1988
Rhododendron arboreum LRB= −1.75 + 0.98 lnGBH 8 Rawat and Singh, 1988
Mixed woody species LRB= −2.25 + 1.00 lnGBH 48 Rawat and Singh, 1988

Fine roots

Acacia catechu FRB= −0.0322+0.0147D 5 25 Singh et al., 2011

Acacia catechu FRB= −0.0322+0.0147D 5 25 Singh et al., 2011
Albizia lebbeck FRB= −0.0007+0.0007D 5 15 Singh et al., 2011
Dalbergia sissoo FRB= −0.0948+0.0664D 5 25 Singh et al., 2011
Dalbergia sissoo FRB=−0.1538+0.1413D 5 12 Lodhiyal and Lodhiyal, 2003
Dalbergia sissoo FRB=−0.6120+0.2012D 10 12 Lodhiyal and Lodhiyal, 2003
Dalbergia sissoo FRB=1.3691+0.1649D 15 12 Lodhiyal and Lodhiyal, 2003
Pinus kesiya FRB= 1.547- 8.2181D + 8.9111D2 - 2.55697D3 40 Baishya and Barik, 2011
Pinus patula FRB= −0.201+0.096D 15 12 Bargali and Singh, 1997
Pinus patula FRB= −0.211+0.093D 18 12 Bargali and Singh, 1997
Pinus patula FRB= −0.031+0.112D 21 12 Bargali and Singh, 1997
Pinus roxburghii FRB= −9.10 + 2.07 lnGBH 16 to 128 26 Chaturvedi and Singh, 1982
Populus deltoides FRB= 0.001D1.92 1 to 9 27 Das et al., 2011
Populus deltoides FRB=−0.1757+0.1092D 5 27 Lodhiyal et al., 1995
Populus deltoides FRB= −0.3663+0.1207D 6 27 Lodhiyal et al., 1995
Populus deltoides FRB=−0.3369+0.1392D 7 27 Lodhiyal et al., 1995
Populus deltoides FRB=−0.8636+0.1646D 8 27 Lodhiyal et al., 1995
Quercus leucotrichophora FRB= −1.33+0.50 lnGBH 23 Rawat and Singh, 1988
Quercus floribunda FRB= −1.05 + 0.25 lnGBH 17 Rawat and Singh, 1988
Rhododendron arboreum FRB= −0.01 + 0.41 lnGBH 8 Rawat and Singh, 1988
Mixed woody species FRB= −2.07 + 0.53 lnGBH 48 Rawat and Singh, 1988
(continued on next page)

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B. Brahma, A.J. Nath, C. Deb et al. Trees, Forests and People 5 (2021) 100098

Table 5 (continued)

Associated Species Equations Stand/ Tree Harvested sample Sources

age (yr) size (number)

Coarse roots

Citrus sinensis CRB= (exp(−1.54 + 1.09(lnAGB))) × 1.05 8 to 30 58 Sahoo et al., 2020

Hevea brasiliensis lnCRB=(exp (−0.53+0.64 (ln AGB)) × 1.03 6 to 34 67 Brahma et al., 2017
Hevea brasiliensis lnCRB = (exp(−2.64 + 0.60 (lnD2 H))) × 1.04 6 to 34 67 Brahma et al., 2017
Pinus kesiya
CRB= −12.2742+ 22.8461D-12.3419D2 +2.2129D3
40
Baishya and Barik, 2011
Taproots

Grewia optiva TaRB=0.103D1.612 4 to 23 25 Verma et al., 2014

Total root

Albizia procera TRB=0.031D2.494 11 32 Newaj et al., 2016

Dalbergia sissoo TRB=0.198D1.760 17 42 Newaj et al., 2016
Embilica officinalis TRB=0.622 CD1.313 15 30 Newaj et al., 2016
Gmelina arborea TRB=−1.967+0.0732D 1 to 6 75 Swamy et al., 2004
Hardwickia binata TRB=0.036 D2.337 20 30 Newaj et al., 2016
Litchi chinensis TRB=(3.41/[1 + 0.13 exp(−0.30 × CD)]ˆ(1/0.03)+ 𝜀) 2 to 10 30 Naik et al., 2018
Pinus kesiya TRB= −9.8635+ 17.8955D-8.9558D2 –1.4511D3 40 Baishya and Barik, 2011
Psidium guajava TRB=0.019CD2.492 4 to 14 17 Rathore et al., 2018
Tectona grandis TRB= 0.097D2.023 20 to 47 Buvaneswaran et al., 2006
Tectona grandis TRB= 0.185D2 - 3.747D+51.498 20 to 47 Buvaneswaran et al., 2006

Belowground biomass

Alnus nepalensis ln BGB =0.916 + 0.720 ln D 7 to 46 17 Sharma and Ambasht, 1991

Azadirachta indica
Azadirachta indica
Azadirachta indica
Azadirachta indica
Azadirachta indica
Eucalyptus tereticornis
Casuarina equisetifolia
Jatropha curcus
Litchi chinensis
Populus deltoides
Pinus roxburghii
Quercus leucotrichophora
Quercus floribunda
Rhododendron arboreum
Tectona grandis
BGB=0.003H5.444 3 Mohamed et al., 2018
BGB= 0.131(BaD)1.972 3 Mohamed et al., 2018
BGB= 0.052D2.388 3 Mohamed et al., 2018
BGB= 8.790CrH2.930 3 Mohamed et al., 2018
BGB= 3.686CrW1.508 3 Mohamed et al., 2018
BGB = 0.130D1.89 8 10 Kumar et al., 2019
ln BGB= −3.35+2.4517 ln D 2 to 9 90 Vidyasagaran and Paramathma, 2014
BGB=0.138CD1.158 8 16 Rao et al., 2016
BGB=(3.52/[1–0.35 exp(0.28 × CD)]ˆ(1/−0.10)+ 𝜀) 2 to 10 30 Naik et al., 2018
BGB= 0.17D1.38 1 to 9 27 Das et al., 2011
BGB =−7.015 + 2.469 lnGBH 16 to 128 26 Chaturvedi and Singh, 1982
ln BGB= 0.112 + 0.924 lnGBH 23 Rawat and Singh, 1988
ln BGB= 0.112 + 0.924 lnGBH 17 Rawat and Singh, 1988
ln BGB= 0.942+ 0.506 ln GBH 8 Rawat and Singh, 1988
BGB=0.1257D2 –1.8573D+9.2119 1 to 30 33 Jha, 2015

Note: BGB= Belowground biomass, BaD=Basal diameter, TRB= Total Root biomass, FRB= Fine root biomass, CRB= Coarse root biomass, LRB= Lateral root biomass,
SRB= Stump root biomass, TaRB= Tap root biomass, D=Diameter at breast height (DBH), H= tree height, AGB= Aboveground Biomass, CD= Collar Diameter, GBH=
Girth at breast height, CrW=Crown width, CrD= Crown diameter, CrH= Crown height, 𝜀= Model error.

also shown as the potential independent variable in biomass studies
(Brahma et al., 2017). The present analysis found that 50% of AGB
biomass estimation equations in India are based D (Table 1). Besides,
H and 𝜌 are the best co-variables combined with D to form a compound
variable for biomass estimation (Sileshi, 2014). Measuring the vertical
tree height in forests is difficult due to their extended height and dense
canopy (Larjavaara and Muller-Landau, 2013), and it requires signifi-
cant effort and sophisticated technologies. Laser range finder and cli-
nometers are effectively used to measure tree heights in forests. At the
same time, the former may not be suitable where there is a blockage
between the range finder and the top of the tree (Larjavaara and Muller-
Landau, 2013). Therefore, the use of tree height in regression equation
estimation is considered the most feasible method where the researcher
can measure tree height accurately.
Predicting tree height using D as the dependent variable is widely ac-
cepted in forest ecology (Sileshi, 2014). However, minimal expressions
for the d-H relationship were found in literature published from India.
Since D can be accurately measured in a wide variety of land use, it may
be considered as more reliable when using a single independent variable
to develop biomass estimation. Michaelis-Menten regression equation
(H= aD/(b + D)) was found to be the best equation among the ten tested
equations for Hevea brasiliensis (Brahma et al., 2017) (Table 8).
There were no functions reported for multi-species tree height pre-
diction based on any of the dependent variables. However, the power-
law, logistic and exponential functions based on the culm diameter
were observed for different bamboo species like Schizostachyum dul-
looa, Pseudostachyum polymorphum, Melocanna baccifera respectively
(Singnar et al., 2017), and wild banana Musa balbisiana (Laskar et al.,
2020).
3.5.2. Choice of functions
Various statistical expressions like the linear, Power-law, and expo-
nential functions are the commonly used forms. Describing the biomass
of different plant parts based on the predictive variables is the objective
of biomass equation development. The power-law function has been rec-
ognized as the most reliable function of biomass equations. At least 50%
of the AGB, BGB, and TB equations in the Indian literatures have used
the power functions based on different predictive variables. The use of
simple variables in equation development can reduce the asymmetrical
error (Liu et al., 2017). Moreover, the compatibility of the equation in
future use could play a significant role in equation selection.

13
B. Brahma, A.J. Nath, C. Deb et al. Trees, Forests and People 5 (2021) 100098

Table 6
Details of the recommended regression equations for total biomass (AGB+BGB) estimation.

Harvested sample
Associated Species Equations Age (yr) size (number) Sources

Bananas
Musa balbisiana TB=(exp(−3.49+0.792(lnD2 Hp)) × 1.05 140 Laskar et al., 2020
Palms
Areca catechu lnTB= ln(−1.853) +0.728 lnD2 H 10 to 35 40 Das et al., 2021
Trees
Acacia nilotica TwB= −0.519 - 0.065 D + 0.557 D2 4.5 to 6.5 75 Rizvi and Ahlawat, 2014
Albizia procera TB= 0.102D2.499 11 32 Newaj et al., 2016
Anogeissus pendula TB= −6.684 + 1.2653CBH 20 Vyas et al., 1978
Azadirachta indica TB= 0.017H5.057 3 Mohamed et al., 2018
Azadirachta indica TB= 0.604(BD)1.810 3 Mohamed et al., 2018
Azadirachta indica TB= 0.252(D)2.202 3 Mohamed et al., 2018
Azadirachta indica TB= 28.42(Cr H)2.709 3 Mohamed et al., 2018
Azadirachta indica TB= 12.86(Cr W)1.386 3 Mohamed et al., 2018
Cassia fistula TB= 0.7837(D2 H)- 0.0004 30 Tripathi, 2015
Citrus sinensis TB= (exp(1.0 + 0.307(lnD2 H))) × 1.09 8 to 30 58 Sahoo et al., 2020
Dalbergia sissoo TB=0.904D1.760 17 42 Newaj et al., 2016
Embilica officinalis TB=2.994 (CD)1.285 15 30 Newaj et al., 2016
Gmelina arborea TB=−2.421+4.2551D 1 to 6 75 Swamy et al., 2004
Hevea brasiliensis ln TB=exp (−2.84+0.90 (lnD2 H)) × 1.02 6 to 34 67 Brahma et al., 2017
Hardwickia binata TB=0.158D2.349 20 30 Newaj et al., 2016
Litchi chinensis TB=29.07/[1 + 0.03 × exp(−0.36 × CD]ˆ(1/0.01)+ɛ 2 to 10 30 Naik et al., 2018
Pinus kesiya TB= 0.3686+ 0.3859D+2.2618D2 –0.7785D3 40 Baishya and Barik, 2011
Pinus roxburghii TB= −6.305 + 2.684 lnCBH 16 to 128 26 Chaturvedi and Singh, 1987
Pongamia pinnata TB= 0.8332 (CD)1.818 9 12 Rao et al., 2016
Populus deltoides TB= 1.90D1.23 1 to 9 27 Das et al., 2011
Tectona grandis lnTB = ln(8.165) + (8.165)lnD2 H 418 Deb et al., 2016
Tectona grandis TB= 0.142D2.469 20 to 47 Buvaneswaran et al., 2006
Tectona grandis TB= 0.202D2.353 20 to 47 Buvaneswaran et al., 2006
Tectona grandis TB=0.6246D2 –2.0593D-12.759 1 to 30 33 Jha, 2015

Note: TB= Total biomass, TwB= total wood biomass, D= Diameter at breast height, CD= Collar diameter, H= Tree height, BD= Basal diameter,
CrW=Crown width, CrD= Crown diameter, p= wood density, CBH= Circumference at breast height of the tree, ɛ= Model error.

Table 7
Root mean square error (RMSE) values of data validation analysis.

RMSE
Equations T. grandis C. fistula P. kesiya Average
2
Equation 1 ln AGB= 3.428 + 0.310ln pD H 375.33 422.92 399.13
Equation 2 ln ABG= −3.206+1.337ln pD2 H 5943.29 920.02 3431.66
Equation 3 AGB=(0.18D2.16 ) × 1.32 227.63 207.48 221.38 218.83
Equation 4 AGB=(0.32pD2 H0.75 ) × 1.34 67,540.42 33,996.20 50,768.31
Equation 5 ln AGB= 0.349 + 1.316 ln GBH 209.83 90.79 167.12 155.91
−1 −3
Note: AGB=Aboveground biomass (kg tree ); p=wood density (g cm ); D=Diameter at breast
height (cm); H=tree height (m).

Table 8
Recommended H-D relationship for bamboo, wild banana and woody tree species in India.

SL. No. Species Name Equations Sources

0.65
1 Schizostachyum dullooa H = 438.4D Singnar et al., 2017)
2 Pseudostachyum polymorphum H = 1329.4/(1+exp(2.79–1.73D) Singnar et al. (2017)
3 Melocanna baccifera H = 329.3(exp(0.28D)) Singnar et al. (2017)
4 Musa balbisiana H= −2.28–3.46(exp(0.0417 D) Laskar et al. (2020)
5 Hevea brasiliensis H = 51.1 D/(42.3 + D) Brahma et al. (2017)

Note: D= Diameter at breast height (DBH) and H= tree height.

3.5.3. Sample size
Sample size or the number of trees to be harvested for equation
development is an essential prerequisite for regression equation de-
velopment. However, a sample size of about 40 or more is consid-
ered adequate for developing biomass equations (Sileshi, 2014). In-
adequate sample size may lead to data truncation on the equation
functions. It may not be possible to harvest a higher number of sam-
ple trees for use in equation development pertaining to thier high-
cost and adverse ecosystem issues; a small sample size, on the other
hand, may yield biased equations (Duncansan et al., 2015). A sta-
tistically valid or appropriate number of datasets covering the girth
and age classes can minimize such effects and reduce equation de-
velopment costs. While it may be relatively easier to develop species-
specific biomass equations (mostly in plantations), developing biomass
estimation equations for mixed communities (in forests) is challeng-
ing. The sample number used for different bamboo species ranged
from 90‒180 for Dendrocalamus strictus, Melocanna baccifera, Pseu-
dostachyum polymorphum and Schizostachyum dullooa (Kaushal et al.,
2016; Singnar et al., 2017; Thokchom and Yadava, 2017). Similarly,
a sample number of 51‒170 for Acacia auriculiformis, Ailanthus tri-
physa, Casuarina equisetifolia, Leucaena leucocephala, Mangifera in-
dica, Tectona grandis, Hevea brasiliensis and Citrus sinensis were used

14
B. Brahma, A.J. Nath, C. Deb et al.
for the woody trees (Sandeep et al., 2015; Chaturvedi and Raghuban-
shi, 2015; Kumar et al., 1998; Vidyasagar and Paramathma, 2014;
Sahoo et al., 2020; Brahma et al., 2017; Chaturvedi et al., 1995;
Ganeshamurthy et al., 2016). A larger dataset is preferable for multi-
species biomass regression equations due to the heterogeneous at-
tributes that prevail in the field conditions and minimize the percent
prediction errors. Various researchers have used as high as 470 and 330
harvested trees drawn from different species (with diameter range up to
100 cm) (Nath et al., 2019; Chaturvedi et al., 2012; Chaturvedi and
Raghubanshi, 2013) in India to develop AGB biomass equations for
multi-species stands for juvenile and mature tree stands respectively.
4. Conclusions
We conclude that D is the best predictive variable for AGB and BGB
estimation. The power-law function with an appropriate simple or com-
pound variable is the best function for biomass equation development.
Cross-validation of five multi-species equations showed that the best-
fit equations for AGB estimation are lnAGB= 0.349+1.316lnGBH and
AGB= (0.18D2.16 )х1.32. The equations can be effectively used to pre-
dict the tree biomass of any wood species across a range of conditions
in India.
Author contributions
A.J.N. formulated the research work. B.B., A.J.N. analyzed the data
and wrote the first draft. B.B., C.D. searched and developed the data set.
All authors jointly discussed the results, drew conclusions and finalized
the manuscript. All authors read and approved the final manuscript.
Declaration of Competing Interest
All authors declare no conflict of interest.
Acknowledgements
Financial assistance received from Department of Science and Tech-
nology is highly acknowledged (DST/CCP/MRDP/190/2019).
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