sustainability
Article
Relative Humidity, Soil Phosphorus, and Stand Structure
Diversity Determine Aboveground Biomass along the Elevation
Gradient in Various Forest Ecosystems of Pakistan
Shahab Ali 1 , Shujaul Mulk Khan 1,2, * , Zeeshan Ahmad 1 , Abdullah Abdullah 1 , Naeemullah Kazi 3 ,
Ismat Nawaz 4 , Khalid F. Almutairi 5 , Graciela Dolores Avila-Quezada 6 and Elsayed Fathi Abd_Allah 5
Citation: Ali, S.; Khan, S.M.; Ahmad,
Z.; Abdullah, A.; Kazi, N.; Nawaz, I.;
Almutairi, K.F.; Avila-Quezada, G.D.;
Abd_Allah, E.F. Relative Humidity,
Soil Phosphorus, and Stand Structure
Diversity Determine Aboveground
Biomass along the Elevation Gradient
in Various Forest Ecosystems of
Pakistan. Sustainability 2023, 15, 7523.
https://doi.org/10.3390/su15097523
Academic Editor: Jasmin
Mantilla-Contreras
Received: 25 February 2023
Revised: 23 April 2023
Accepted: 25 April 2023
Published: 4 May 2023
Copyright: © 2023 by the authors.
Licensee MDPI, Basel, Switzerland.
This article is an open access article
distributed under the terms and
conditions of the Creative Commons
Attribution (CC BY) license (https://
creativecommons.org/licenses/by/
4.0/).
Sustainability 2023, 15, 7523. https://doi.org/10.3390/su15097523
1 Department of Plant Sciences, Quaid-i-Azam University Islamabad, Islamabad 45320, Pakistan
2 Pakistan Academy of Science, Islamabad 45320, Pakistan
3 Sindh Wildlife Department, Karachi 76090, Pakistan
4 Department of Biosciences, Islamabad Campus, COMSATS University Islamabad, Islamabad 45550, Pakistan
5 Plant Production Department, College of Food and Agricultural Sciences, King Saud University,
P.O. Box 2460, Riyadh 11451, Saudi Arabia
6 Facultad de Ciencias Agrotecnológicas, Universidad Autónoma de Chihuahua, Chihuahua 31350, Mexico
* Correspondence: smkhan@qau.edu.pk
Abstract: The direct effects of relative humidity and soil on aboveground biomass (AGB) versus the
indirect effects mediated by stand structural diversity remain unclear in forest ecosystems across
large-scale elevation gradients. Forest inventory data containing 15,260 individual trees and 104 tree
species from 200 forest plots were collected. The result shows that the relative humidity, elevation,
and Coefficient of Variation of Diameter at breast height (CVD) significantly influence AGB in the
Tropical Thorn Forest (TTF). Regarding elevation, CVD was positive and significant, and relative
humidity and SR negatively impacted AGB in sub-tropical broad-leaved forests (STBLF). In moist
temperate mixed forests (MTMF), soil phosphorus and CVD have a significant positive impact, while
relative humidity, elevation, and SR negatively influence AGB. Elevation and CVD have positive,
while SR and soil phosphorus have a negative and insignificant effect on AGB in Dry Temperate
Conifer Forests (DTCF). Soil phosphorus and relative humidity positively affected AGB (β = 0.021),
while elevation, CVD, and SR negatively affect AGB in dry temperate, pure pine forests (DTPPF).
Relative humidity and soil phosphorus have a positive direct effect on AGB in multi-species forests.
The current study suggests that AGB primarily depends on relative humidity, soil phosphorus, and
elevation in different forest types.
Keywords: forest inventory; environmental factors; stand structure complexity; regional scale;
structural equation modeling
1. Introduction
The influence of species richness (SR) and stand structure diversity (SSD) on above-
ground biomass (AGB) has been widely studied and found to be significant. Species
richness and SSD contribute to overall ecosystem productivity and health and directly
impact AGB. High species richness and stand structure diversity are important for main-
taining long-term ecosystem sustainability and resilience [1,2]. SR is considered a part of
stand structure complexity (SSC), but variations in tree diameter and height, combined or
alone, are primarily categorized as SSC in the forest ecosystem [3]. The ecological tools
for linking the various biotic, i.e., biodiversity, stand structure, and abiotic, i.e., relative
humidity and soil factors concerning aboveground biomass, have been comparatively well
evaluated [2,4–8]. Direct and indirect effects on AGB, SR, and SSC across regional scale
ecological gradients persist less contested in natural forest ecosystems [2,4,5].
Nevertheless, ecological theories and regional-level experimental studies indicate that
climatic conditions mainly determine AGB [9–12]. Especially in the natural forest ecosystem,
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Sustainability 2023, 15, 7523 2 of 16

higher SR and AGB have usually experienced in regions with high precipitation and climatic
water availability [13,14]. Diversity might greatly enhance AGB production due to the
enabling effect in severe ecological conditions instead of a productive environment due
to the sturdy antagonistic effect in inferior species forest ecosystems [15]. Furthermore,
the fertility soil hypothesis assumes the plant can grow competently under high nutrient
availability, leading to significant competition [16]. Nevertheless, most tropical forest
species are distributed on deficient nutrients and wet soil [17,18]. In addition, it is well-
assumed in the natural forest ecosystem that conservative and acquisitive species are
situated on the opposite excesses of nutrient-axis soil, e.g., nutrient-deficient and rich
soil, respectively [19–22]. However, in this condition, it is reasonable to assume that SR
enhances AGB and demographic processes, i.e., growth and enlistment underneath suitable
climatic conditions instead of under excellent fertile soil conditions in a natural tropical
forest ecosystem [2,13,23]. Through assistance mechanisms and niche distinction multi-
layered stand structure, theories have been suggested to enhance light capturing and its
utilization within and amongst the constituent species in a community [24]. However, SR
and species diversity within a forest stand or community recognize frequent ecological
mechanisms and might increase or decrease AGB directly, i.e., niche complementarity, mass
ratio and competitive elimination effects [13,25–27]. Furthermore, SR or species diversity
and SSC endorse one another to achieve maximum AGB or storage of carbon over the
effect of niche complementarity in a natural forest ecosystem [26,28,29]. Nevertheless, the
direct independent predictors of AGB might be SSC, and species diversity withstands
structure as a better predictor [30–32]. However, the multivariate association among SR,
SSC, and AGB remains debated. This paper documents the direct and indirect effects of
relative humidity and soils on SR, SSC, and AGB in different natural forest types along
regional-scale elevation gradients.

Hypothesized Model
The key objective of the current study was to define the combined effect of relative
humidity, soil via SR, and coefficient of diameter variations, such as breast height (CVD)
on AGB, across different forest types and species mixtures along regional-scale elevation
gradients across Pakistan (Figure 1). The forest inventory dataset comprised 15,260 in-
dividuals belonging to 104 tree species from 200 forest plots in this study. These plots
were classified into five local forest types and two groups of single- and multiple-species
stand forests based on species composition. We used structural equation modeling, a
powerful integrative approach to forest inventory data [33]. We hypothesized that climatic
factors (relative humidity), soil (phosphorus) stand structure complexity, and regional-scale
elevation gradients have a robust positive effect on AGB in multi-species forest types com-
pared to single-species stand forest types. Secondly, we hypothesized that the relationship
among CVD, SR, relative humidity, and soil directly depends on the forest’s stand type,
while the multi-species forest leads to higher AGB through higher relative humidity water
availability and higher soil fertility factors and size of the individual trees (Figure 1).
23, 15, x FOR PEER REVIEW
Sustainability 2023, 15, 7523 3 of 16 3 of 16

Figure
Figure 1. A theoretical 1. A theoretical
model modelthe
for evaluating forhypothesized
evaluating theassociation
hypothesized association
among amongSoil,
elevation, elevation, Soil,
relative humidity, and SR on AGB in a regional
relative humidity, and SR on AGB in a regional forest of Pakistan. forest of Pakistan.

2. Materials and Methods

2. Materials and Methods
2.1. Study Area
2.1. Study Area The current study was conducted across five different forest types in Pakistan.
(1) Tropical
The current study was conductedthorn forest across
(2) Sub-tropical
five differentbroad-leaved forestin
forest types (3)Pakistan.
Dry temperate
(1) conifer
forest (4) Moist temperate mix forest (5) Dry temperate, pure pines forest (Figure 2). The
Tropical thorn forest (2) Sub-tropical broad-leaved forest (3) Dry temperate conifer forest
tropical thorn forest ecosystem of the Kirthar is located in the mountainous range of south-
(4) Moist temperate mix forest (5) Dry temperate, pure pines forest (Figure 2). The tropical
western Sindh. The area has been categorized as a protected category II by the international
thorn forest ecosystem
union of forthe
theKirthar is located
conservation in the
of nature mountainous
(IUCN). range
It covers the of mountainous
rocky southwestern ranges from
Sindh. The area has been
north to categorized as aby
south, separated protected
flat valleys category
[34]. The II sub-tropical
by the international
broad-leavedunionforest of Mar-
for the conservationgallaof Hills
nature (IUCN).
National ParkIt covers
is locatedtheatrocky mountainous
the western ranges
extremities of thefrom north
foothills of the lesser
2
to south, separated by flat valleys
Himalayas, covering[34]. Theofsub-tropical
an area 126.05 km [35]. broad-leaved forest of
The dry temperate Margalla
conifer forest lies in Dir
Hills National Park upper, with approximately
is located at the western 2000 masl. The monthly
extremities mean minimum
of the foothills and maximum
of the lesser Him- temper-
atures have been recorded as 11.22 ◦ C and −2.39 ◦ C [36]. The moist temperate mixed forest
alayas, covering an area of 126.05 km [35]. The dry temperate conifer forest lies in Dir
2
is located in Murree and Ayubia in the temperate region of the sub-Himalayan Mountains.
upper, with approximately 2000 masl. The monthly mean minimum and maximum tem-
The mountains of Murree and Ayubia reach an altitude of about 5000 to 8200 feet [37]. The
peratures have been recorded as 11.22 °C and −2.39 °C [36]. The moist temperate mixed
dry temperate, pure pine forest located in the Suleiman Range District Shirani covers an
forest is located in Murree
area of 260 km and Ayubia
2 and containsin the
theworld’s
temperate largestregion of the
Chilghoza sub-Himalayan
forest (Pinus gerardiana). It is an
Mountains. The mountains of Murree and Ayubia reach an altitude
extension of the Hindu Kush mountain at the junction of the three of about 5000 to 8200borders of
provincial
feet [37]. The dry Balochistan,
temperate, pure Khyberpine forest located
Pakhtunkhwa, andinPunjab
the Suleiman Range
[38] (Figure 2). TheDistrict Shi-
detailed description,
rani covers an area i.e.,offorest
260 types,
km2 and contains
location, and the world’s
latitude largest Chilghoza
and longitude forest (Pinus
climatic condition, stand type and
main
gerardiana). It is an or dominant
extension species
of the Hindu are Kush
given in Table 1. at the junction of the three
mountain
provincial borders of Balochistan, Khyber Pakhtunkhwa, and Punjab [38] (Figure 2). The
detailed description, i.e., forest types, location, and latitude and longitude climatic condi-
tion, stand type and main or dominant species are given in Table 1.
 Sustainability 2023, 15, x FOR PEER REVIEW  4  of  16 
 
Sustainability 2023, 15, 7523 4 of 16

 
Figure 2. GIS-generated map of the studied forests across Pakistan.
Figure 2. GIS-generated map of the studied forests across Pakistan. 

2.2. Forest Inventory

Table 1. Forest type, elevation, stand type, mean annual temperature, mean annual precipitation, 
We randomly sampled 40 (20 × 20 m) plots in each forest from March 2020 to
and main or dominant species of the studied forest types. 
September 2020. Plots were established at one km or 20 m altitudinal variation. All
Mean Annual  Mean Annual 
three plant individuals with a ≥1 cm diameter at breast height (DBH) were noted
Elevation 
S/No  Forest Type  Stand Type  Coordinates 
within each plot. The Temperature  Precipitation 
tree  DBH was recorded Dominant Species 
through measuring tape. The recorded
m.a.s.l 
(°C)  (mm)   
tree plants (≥1 cm DBH) present in the plots were identified by taxonomists and
Prosopis glandulosa Torr., 
confirmed at http://www.efloras.org/flora_page.aspx?flora_id=5, (accessed on 12
October 2020) http://www.theplantlist.org/tpl1.1/search, Prosopis juliflora (Sw.) DC., 
(accessed on 12 October
670 10ˈ  to 670 
2020). GPS (geographical positioning system) and Acacia modesta Wall., 
elevation values were also noted for
Sub-tropical  Conserved  55ˈ  E longi-
each plot on the x-axis. Ziziphus nummularia 
1  thorn  natural for- tudes to 250  56–302  33.8  245.3 
(Burm.f.) Wight & Arn.,   
Forest  est  13ˈ  to 260 12ˈ 
Salvadora oleoides Decne.,   
N latitude 
Combretum molle R.Br. ex 
G.Don 
Sustainability 2023, 15, 7523 5 of 16

Table 1. Forest type, elevation, stand type, mean annual temperature, mean annual precipitation, and main or dominant species of the studied forest types.

Elevation Mean Annual Mean Annual

S/No Forest Type Stand Type Coordinates Dominant Species
m.a.s.l Temperature (◦ C) Precipitation (mm)
Prosopis glandulosa Torr., Prosopis juliflora
(Sw.) DC.,
670 10| to 670 55| E
Sub-tropical thorn Conserved natural Acacia modesta Wall., Ziziphus
1 longitudes to 250 13| 56–302 33.8 245.3
Forest forest nummularia (Burm.f.) Wight & Arn.,
to 260 12| N latitude
Salvadora oleoides Decne.,
Combretum molle R.Br. ex G.Don
Acacia modesta Wall., Cassia fistula L.,
33,0400 –33,0440 N Justicia adhatoda L., Carissa spinarum L.,
Sub-tropical Conserved natural longitude to Mallotus philippensis (Lam.) Müll.Arg.,
2 555–1117 27.8 1572.1
broad-leaved forest forest 33,0550 –73,0200 E Dodonaea viscosa (L.) Jacq., Bauhinia
latitude variegata L., Albizia lebbeck (L.) Benth.,
Celtis australis L.
Pinus wallichiana A.B.Jacks., Diospyros
virginiana L.,
Aesculus indica (Wall. ex Cambess.)
33◦ 520 to 33◦ 590 N
Moist temperate Hook., Populus alba L., Cedrus deodara
3 Old age mix forest and 73◦ 240 to 73◦ 310 1249–2892 17.8 1596.1
mix forest (Roxb. ex D.Don) G.Don, Castanea
E
dentata (Marshall) Borkh., Abies pindrow
(Royle ex D.Don) Royle, Quercus dilatata
A.Kern.
Pinus roxburghii Sarg., Abies pindreow
(Royle ex D.Don) Royle,
Dry temperate Old age conifer 350–280 N latitude to
4 1040–2566 23.4 1371.8 Pinus wallichiana A.B.Jacks., Picea
conifer forest forest 720–200 E longitude
smithiana (Wall.) Boiss.,
Cedrus deodara (Roxb. ex D.Don) G.Don
Dry temperate pure 310–36 N latitude
Old age pure Pinus
5 Pinus gerardiana and 690–59 E 1841–2282 25.9 299.0 Pinus gerardiana Wall. ex D.Don
gerardiana forest
forest longitude
Sustainability 2023, 15, 7523 6 of 16

2.3. Statistical Analysis Predictor and Predicted Variable

Our hypothetical model (Figure 1) comprises six variables: elevation, mean relative
humidity, soil phosphorus, SR, CVD, and AGB. Climatic data (relative humidity) were noted
for each plot using a digital humidity meter [2]. The current study used soil phosphorus (P)
as a soil factor. Soil phosphorus was determined by following the protocol of [39,40]. We
used the Shannon’s species diversity index because the index deals with species evenness
and SR of the community, i.e., plots, and predicts better AGB in the natural forest [26], and,
similarly, for quantifying the coefficient variation of DBH for each plot [41]. Pantropical
allometric equations were used (AGB = 0.0673 (ρ dbh × h) 20.976) for tropical forests, and
each tree’s AGB estimation was based on tree height, diameter at breast height (dbh), and
species wood density [42].
For Pines Forest, we used the below equation;

Crown Biomass

WC = 0.1377 (D)1.4873 × (L)0.4052

WS = 0.0600 (D)0.7934 × 0.7934 (L)1.8005

Total Biomass = WC + WS
where WC = woody crown, WS = woody stem.

2.4. Statistical Analysis

The first question to answer is whether the fixed factors, which include relative
humidity, elevation, soils, CVD, and SR, or the random factors that affect, i.e., types of
forests single-species, species mixture, and climatic factors, explain higher variability in
aboveground biomass. Piecewise structure equation modeling (PSEM) deals with the
random and fixed effects in a single equation [33]. This study used two piecewise structural
equation models to test the primary conceptual model as a random effect of species mixture
and forest type. The above hypothetical model described: (1) The effect of soil, relative
humidity, CVD, and SR on AGB along a large regional scale elevation gradient; (2) AGB is
affected by climatic factors, soil, and DBH; (3) Elevation, SR, and soil effect in different types
of forest and mixture species forest on the AGB. We determined the Pearson correlation and
removed the insignificant relationship in the equation to obtain a good fit model. We used
random effect PSEM for species mixture and excluded the model’s fixed effect to avoid the
problem of model circularity. To evaluate the model fit of PSEM, we used Fisher statistics.
Moreover, we applied the directional separation test in the CFA in the case of poor fit or
over the fitted model; it included or excluded any significant or insignificant path, though
in our study, the case was not a strong one.
Therefore, the second question is the relationship between relative humidity, soil, SR,
CVD, and AGB along different regional-scale elevation gradients in different forest types.
We divided the collective dataset into five local forest types; for each conceptual model,
we used linear structural equation modeling (SEM) techniques (Figure 1). To investigate
the 3rd question, to determine whether the relationship between relative humidity, soil,
SR, CVD, AGB, and elevation fluctuates along with single-species or multi-species forest
types, we again divided the data into two groups, single-species and multi-species forests,
then applied linear SEM. Four fit indices, i.e., we determined the Acai information criteria
(AIC), comparative fit index (CFI), the goodness of fit index (GFI), and root mean square
residual (SRMR) via linear SEM model fit. The goodness of fit model is based on CFI > 0.90,
GFI > 0.90, and SRMR < 0.08 [43]. We applied the same models for all seven equations.
Here, we mainly focused on the relationship amongst the variables, not only the goodness
of fit models. The direct effect was identified among dependent and independent variables
through a direct path. The multiplication of dependent and independent measured the
Sustainability 2023, 15, 7523 7 of 16

models’ indirect effect and mediator variables. For the current analysis, R codes previously
developed by [44] were used.
We calculated the simple linear regression for each conceptual model among pooled
and binary variables (forest type and species mixture). We converted all the variables into
continuous form to estimate linear SEM analysis values through a natural logarithm. We
transformed them to standardize and satisfy linearity and normality assumptions. We
estimated the Pearson correlation coefficient of all the independent variables and took
only those variables in the model with a significant relationship. We also checked the
multicollinearity among the variables through the VIF test; there was no such issue of
multicollinearity in our case.

3. Results
In the sub-tropical thorn forest (TTF), we found the impact of soil phosphorus had a
negative and significant on AGB (β = −0.849), and relative humidity had an insignificant
positive effect on AGB (β = 1.352). In contrast, elevation had a significant positive effect
on AGB (β = 3.126). The effect of a stand structure, i.e., coefficient of variation of DBH
(CVD), was recorded as positive and insignificant (β = 0.087), and species richness (SR) had
a negative and insignificant effect on AGB (β = −0.823) (Figure 3A & Table S1).
In the sub-tropical broad-leaved forest (STBLF), the impact of soil phosphorus
had a positive and significant effect on AGB (β = 0.048), and relative humidity had an
insignificant negative effect on AGB (β = −0.001), while elevation had a significant
positive impact on AGB (β = 0.025). Conversely, relative humidity and elevation had
an indirect (through CVD & soil phosphorus) positive effect on AGB. The effect of
CVD was recorded as positive and significant (β = 0.046), and SR had a negative and
insignificant effect on AGB (β = −0.004) (Figure 3B & Table S2).
In the moist temperate mixed forest (MTMF), the impact of soil phosphorus had
a positive and significant effect on AGB (β = 0.484), and relative humidity had an
insignificant negative effect on AGB (β = −0.229), while elevation had a significant
negative effect on AGB (β = −0.161). Conversely, relative humidity and elevation had
an indirect (through CVD & soil phosphorus) positive effect on AGB. The effect of CVD
was recorded as positive (β = 0.062), and SR had a negative and insignificant effect on
AGB (β = −0.046) (Figure 3C & Table S3).
In the dry temperate conifer forest (DTCF), the impact of soil phosphorus had a negative
and significant effect on AGB (β = −0.849), and relative humidity had a positive effect on AGB
(β = 1.352). Similarly, elevation had a significant positive effect on AGB (β = 3.126). Conversely,
relative humidity had a positive indirect effect on AGB through soil phosphorus and CVD,
while elevation had a negative indirect effect on AGB through soil phosphorus, CVD, and SR.
The effect of CVD was recorded as positive (β = 0.087), and SR had a negative and insignificant
effect on AGB (β = −0.823) (Figure 3D & Table S4).
In the dry temperate, pure pine forest (TTPPF), the impact of soil phosphorus had a
positive effect on AGB (β = 0.023), and relative humidity had a significant negative effect on
AGB (β = 0.021), while elevation had an insignificant negative effect on AGB (β = −0.045).
The effect of CVD was recorded as positive and insignificant (β = 0.017), and SR had a
negative and insignificant effect on AGB (β = −0.001) (Figure 3E & Table S4). The detail of
the descriptive analysis is given in Supplementary Table S5.
We compared direct (dark green bar) and indirect (maroon bar) effects derived from
structural equation models of elevation, relative humidity, soil, CVD, and SR on AGB
and regional forest types. We took standardized coefficient plus and minus standard
error. Abbreviations: elevation (Elev), soil phosphorus (P), relative humidity (mrh), species
richness (SR), and coefficient of variation of DBH (CVD) (Figure 4).
Results of the linear structural equation model of single tree forests showed that
elevation, soil phosphorous, and CVD had a positive effect on AGB, i.e., (β = 0.255),
(β = 0.084), (β = 0.046), while relative humidity had a negative effect on AGB,
Sustainability 2023, 15, 7523 8 of 16

Sustainability 2023, 15, x FOR PEER REVIEW 
−0.001). At the same time, relative humidity had a positive indirect effect8 on
i.e., (β = of  16 
AGB
 
through CVD, and soil, i.e., (β = 0.69), and (β = 0.073), respectively (Figure 5A & Table S7).

 
Figure 3. The relation among elevation, relative humidity, soil, DBH, and aboveground biomass in 
Figure 3. The relation among elevation, relative humidity, soil, DBH, and aboveground biomass
(A) Sub-tropical thorn forest, (B) Sub-tropical broad-leaved forest, (C) Moist temperate mixed forest, 
in (A) Sub-tropical thorn forest, (B) Sub-tropical broad-leaved forest, (C) Moist temperate mixed
(D) Dry temperate conifer forest, and (E) Dry temperate, pure pine forest. Note: The black and or-
forest, (D) Dry temperate conifer forest, and (E) Dry temperate, pure pine forest. Note: The
ange bold lines represented positive and negative significant impacts, respectively (p < 0.05). In con-
black and
trast,  the orange
dashed bold lines
black  and represented
orange  line  positive and
shows  the  negative
positive  significant
and  impacts, respectively
negative  insignificant  impact 
(p amongst the predictors (p > 0.05). We used a standardized regression coefficient for each path. The 
< 0.05). In contrast, the dashed black and orange line shows the positive and negative
model fits indices for tropical thorn, i.e., tropical thorn forest. CFI; 0.84; GFI; 0.90; SRMR; 0.105; AIC; 
insignificant impact amongst the predictors (p > 0.05). We used a standardized regression
379.131.  Sub-tropical  broad-leaved  forest.  CFI;  0.981  GFI;  0.936  SRMR;  0.094 AIC;  119.305.  Moist 
coefficient for each path. The model fits indices for tropical thorn, i.e., tropical thorn forest.
temperate mix forest, CFI; 0.913 GFI; 0.978 SRMR; 0.092 AIC; 222.596. Dry temperate conifer forest, 
CFI; 0.84; GFI; 0.90; SRMR; 0.105; AIC; 379.131. Sub-tropical broad-leaved forest. CFI; 0.981 GFI;
CFI; 0.981 GFI; 0.936 SRMR; 0.094 AIC; 119.305. Dry temperate, pure, CFI; 0.840 GFI; 0.900 SRMR; 
0.936 SRMR; 0.094 AIC; 119.305. Moist temperate mix forest, CFI; 0.913 GFI; 0.978 SRMR;
0.105 AIC; 379.131. 
0.092 AIC; 222.596. Dry temperate conifer forest, CFI; 0.981 GFI; 0.936 SRMR; 0.094 AIC; 119.305.
We compared direct (dark green bar) and indirect (maroon bar) effects derived from 
Dry temperate, pure, CFI; 0.840 GFI; 0.900 SRMR; 0.105 AIC; 379.131.
structural equation models of elevation, relative humidity, soil, CVD, and SR on AGB and 
Sustainability 2023, 15, x FOR PEER REVIEW 9 of 16

regional forest types. We took standardized coefficient plus and minus standard error.
Sustainability 2023, 15, 7523 9 of 16
Abbreviations: elevation (Elev), soil phosphorus (P), relative humidity (mrh), species rich-
ness (SR), and coefficient of variation of DBH (CVD) (Figure 4).

Sustainability 2023, 15, x FOR PEER REVIEW 10 of 16

Figure
Figure 4.
4. The
Thedirect
directand
andindirect
indirecteffect
effectofofthe
theindependent
independentvariable
variableononaboveground
abovegroundbiomass.
biomass.

Results of the linear structural equation model of single tree forests showed that ele-
vation, soil phosphorous, and CVD had a positive effect on AGB, i.e., (β = 0.255), (β = 0.084),
(β = 0.046), while relative humidity had a negative effect on AGB, i.e., (β = −0.001). At the
same time, relative humidity had a positive indirect effect on AGB through CVD, and soil,
i.e., (β = 0.69), , and (β = 0.073), respectively (Figure 5A & Table S7).
In multiple tree species forest, we saw that elevation, soil phosphorous, and relative
humidity had a positive effect on AGB, i.e., (β = 0.255), (β = 0.302), (β = 0.04), while CVD
had a negative effect on AGB, i.e., (β = −0.010). Relative humidity and elevation had a
positive indirect effect on AGB through soil phosphorous and CVD, i.e., (β = 0.21), (β =
0.169) (Figure 5B & Table S8).
The result showed that there are strong relationships among all variables. The two
red values represented negative and significant relations between elevation, and relative
humidity. In comparison, other variables showed strong positive and significant relation-
ships among the variables (Figure 6).

Figure 5.
Figure 5. (A)
(A)Link
Linkamong
among elevation, relative
elevation, humidity,
relative soil,soil,
humidity, CVD, and aboveground
CVD, and abovegroundbiomass in sin-in
biomass
gle-species forests,
single-species (B)(B)
forests, TheThe
link among
link amongelevation, relative
elevation, humidity,
relative humidity, soil, DBH,
soil, DBH,and
andaboveground
aboveground
biomass in a multi-species forest of Pakistan. Note: The black and orange bold lines represent posi-
biomass in a multi-species forest of Pakistan. Note: The black and orange bold lines represent positive
tive and negative significant impact, respectively (p < 0.05), while the dashed black and orange lines
and negative significant impact, respectively (p < 0.05), while the dashed black and orange lines show
show positive and negatively insignificant impact among the predictors (p > 0.05). The model fit
positive andsingle-species
indices for negatively insignificant
forest type impact among
CFI; 0.805 GFI;the predictors
0.921 SRMR; (p > 0.05).
0.087 AIC;The model
618.877 andfitmodel
indicesfitfor
single-species
indices for theforest type CFI;forest
multi-species 0.805type
GFI;CFI;
0.9210.921
SRMR;
GFI;0.087 AIC;
0.965; 618.877
SRMR; and
0.071 model
AIC; fit indices for the
4792.035.
multi-species forest type CFI; 0.921 GFI; 0.965; SRMR; 0.071 AIC; 4792.035.
We compared direct (dark green bars) and indirect (maroon bar) effects derived from
In multiple
structural treemodels
equation speciesofforest, we saw
elevation, that elevation,
relative humidity, soil
soil,phosphorous,
and CVD on AGB and relative
along
humidity had a positive effect on AGB, i.e., (β = 0.255), (β = 0.302), (β = 0.04), while
regional-scale elevation gradients of single and multi-species forest types in Pakistan. We
CVD
took ahad a negativecoefficient
standardized effect on (Figure
AGB, i.e., (β = −0.010). Relative
6). Abbreviations: humidity
Elev, elevation; andphospho-
P, soil elevation
rus; much, relative humidity; SR, SR, and CVD.

Sustainability 2023, 15, 7523
Figure 5. (A) Link among elevation, relative humidity, soil, CVD, and aboveground biomass in sin-
gle-species forests, (B) The link among elevation, relative humidity, soil, DBH, and aboveground
biomass in a multi-species forest of Pakistan. Note: The black and orange bold lines represent posi-
tive and negative significant impact, respectively (p < 0.05), while the dashed black and orange lines
show positive and negatively insignificant impact among the predictors (p > 0.05). The model 10 offit
16
indices for single-species forest type CFI; 0.805 GFI; 0.921 SRMR; 0.087 AIC; 618.877 and model fit
indices for the multi-species forest type CFI; 0.921 GFI; 0.965; SRMR; 0.071 AIC; 4792.035.

had We
a positive
comparedindirect
directeffect
(darkon AGBbars)
green through soil phosphorous
and indirect andeffects
(maroon bar) CVD, i.e., (β =from
derived 0.21),
(β = 0.169) (Figure 5B & Table S8).
structural equation models of elevation, relative humidity, soil, and CVD on AGB along
The resultelevation
regional-scale showed that there are
gradients strongand
of single relationships among
multi-species all types
forest variables. The twoWe
in Pakistan. red
values
took represented negative
a standardized coefficientand significant
(Figure relations between
6). Abbreviations: elevation,P,and
Elev, elevation; soilrelative
phospho-hu-
midity. In comparison, other variables showed
rus; much, relative humidity; SR, SR, and CVD. strong positive and significant relationships
among the variables (Figure 6).

Figure 6. The direct and indirect effect of an independent variable on aboveground biomass.
Figure 6. The direct and indirect effect of an independent variable on aboveground biomass.
We compared direct (dark green bars) and indirect (maroon bar) effects derived from
Overall, we tested our proposed hypothesis and found our hypothesis acceptable be-
structural equation models of elevation, relative humidity, soil, and CVD on AGB along
cause soil factors,
regional-scale climatic
elevation factors, of
gradients and structural
single complexity forest
and multi-species along types
regional-scale eleva-
in Pakistan. We
tion gradients have a strong positive effect on AGB in multi-species forest types compared
took a standardized coefficient (Figure 6). Abbreviations: Elev, elevation; P, soil phosphorus;
to single-species
much, forest types.
relative humidity; SR, Similarly,
SR, and CVD.the relationship among CVD, SR, relative humid-
ity, and soil P we
Overall, is directly dependent
tested our proposed onhypothesis
the stand type
and of the forest.
found Still, the multi-species
our hypothesis acceptable be-
forest leads to higher AGB through higher relative humidity water availability,
cause soil factors, climatic factors, and structural complexity along regional-scale higher soil
elevation
fertility factors, and the size of the individual trees (Figure 7).
gradients have a strong positive effect on AGB in multi-species forest types compared to
single-species forest types. Similarly, the relationship among CVD, SR, relative humidity,
and soil P is directly dependent on the stand type of the forest. Still, the multi-species forest
Sustainability 2023, 15, x to
leads FOR PEER REVIEW
higher AGB through higher relative humidity water availability, higher soil fertility11 of 16

factors, and the size of the individual trees (Figure 7).

Figure
Figure 7. Pearson 7. Pearson
correlation correlation
among among explanatory
explanatory variables. Abbreviations:
variables. Abbreviations: Elev, elevation;
Elev, elevation; P, soil P, soil
phosphorus; mrh, relative humidity; SR, Species Richness, and CVD, Coefficient Variation of DBH.of DBH.
phosphorus; mrh, relative humidity; SR, Species Richness, and CVD, Coefficient Variation

4. Discussion
The current study examines soil and relative humidity’s direct and indirect effect on
AGB via SR and stand structure complexity along regional-scale elevation gradients
across tropical, subtropical, and temperate forests. The current study shows that relative
Sustainability 2023, 15, 7523 11 of 16

4. Discussion
The current study examines soil and relative humidity’s direct and indirect effect on
AGB via SR and stand structure complexity along regional-scale elevation gradients across
tropical, subtropical, and temperate forests. The current study shows that relative humidity
affects AGB indirectly along regional-scale elevation gradients. Moreover, soil affects
AGB directly and indirectly via stand structure complexity. In addition, stand structure
complexity directly increase AGB along regional-scale elevation gradients.

4.1. Effect of Relative Humidity and Soil on AGB in Individual Forest Type
In SEM, the AGB is essentially and significantly affected by soil (phosphorus) rather
than relative humidity. In the present study, the direct effect of relative humidity and soil is
also supported by other researchers’ findings that relative humidity and soil predominantly
regulate AGB on a large scale [14]. The indirect findings of the current study are also
supported by [2,45]. AGB variation is mostly affected by relative humidity and soil along
large-scale ecological gradients. In addition, the insignificant direct effect of relative
humidity on AGB in our SEM is strongly supported by the previous study, i.e., that AGB is
insignificantly directly affected by climatic water availability [46,47]. We also found that
elevation has an insignificant effect on AGB in overall and single forest types. Nevertheless,
the present study also highlighted that when elevation increases, AGB decreases; this might
be because the soil fertility decreases with an increase in elevation, and wind pressure also
increases; hence decline occurs in AGB. The present study is critically supported by the
findings of other researchers accordingly. We found the opposite relation of AGB with
elevation gradients [48,49]. In the cited literature, soil fertility becomes a more limiting
factor than plant needs at higher elevations, decreasing forest productivity and biomass
accumulation [50,51]. Commonly decreasing soil nutrients has been recommended to
decrease tree height and biomass production with elevation [52,53].
We investigated the effect of relative humidity and soil along the elevation gradient
in tropical thorn forests, subtropical broad-leaved forests, moist-temperate mix forests,
dry-temperate coniferous forests, and dry-temperate pine forests. We found that relative
humidity had an insignificant effect on AGB while indirectly, through stand structure
complexity, positive and significant effects were noted on AGB in individual forest types.
Yet, soil’s positive and direct effect on AGB was noted, along with a significant indirect
effect on AGB through stand structural complexity, though the SR had an insignificant
direct impact on aboveground biomass. The previous literature supported these results,
i.e., the elevation is strongly correlated and harms AGB in China’s tropical and temperate
forests, where the soil fertility and climatic variables vary [54]. Our study found that AGB
was strongly correlated with soil in all forest types and had a positive and significant effect
on AGB directly and indirectly. The finding of our study aligns with the results of the
Guyanese tropical rainforest, where AGB and productivity strongly correlated with a high
concentration of phosphorus [55,56]. The current study found that overall elevation has a
considerable positive effect on AGB and production in the tropical thorn forest, followed
by the subtropical broad-leaved forest, dry-temperate pine forest, dry-temperate coniferous
forest, and multi-species forests.
However, elevation has an insignificant and adverse impact in moist temperate mixed
and single-species forests, while, in our study, the forest type and species mixture in the
SEM indicate a positive and insignificant effect on elevation. Our result is supported
by other researchers’ findings that elevation, directly and indirectly, affects aboveground
biomass, even in the sub-tropical forest with small elevation gradients affected by AGB [54].
It is mostly due to temperature and rainfall varying with elevation changes [57,58]. Nev-
ertheless, elevation has an insignificant positive effect in the moist-temperate mix forest;
similarly, in the single-species forest type, insignificant and negative effects were observed.
The findings of our study were supported by the findings of other researchers stating that
an increase in elevation leads to a decline in soil fertility. Hence, plants need more fertile
soil for growth, and, as a result, decreases occur in AGB [48,50,51,59–61].
Sustainability 2023, 15, 7523 12 of 16

4.2. Direct and Indirect Effects in Single and Multi-Species Forest Type
Our study of the SEM shows that elevation and relative humidity have negative and
insignificant effects on AGB in the single-species forest. The negative relationship between
AGB and elevation may refer to a statement from previous studies that an increase in
altitudinal gradients leads to a lower soil water content and lower temperature. As a
result, AGB and production decrease [62]. In addition, the areas at higher altitudes have
a shorter growing season, leading to lower AGB [45,63]. In contrast, the positive effect of
the altitudinal gradient in the tropical-thorn forest, and the sub-tropical forest, in which
the AGB strongly and positively correlates with altitudinal gradients, increases AGB with
an increase in altitude up to a limit. The other studies also found that biomass varies
with elevation in lower altitudes from 1000 to 1500 m [64–66]. In comparison, it shows
an insignificant positive effect on AGB through a stand’s structural complexity and soil.
The low-land forest-type soil is significantly correlated with large trees [67]. Soil positively
correlates with stand structure complexity and insignificantly affects AGB in the single-
species forest types. The positive relationship between soil and stand structure complexity
is due to the interaction among the plants in a community that covers all the available
spaces [68–71]. On the other hand, the soil is positively and significantly correlated with
aboveground biomass in multi-species forest types.

5. Conclusions
We concluded that the single-species forest relative humidity and elevation have a
negative and insignificant effect on AGB. Moreover, stand structure diversity indirectly
affects AGB, while soil has a positive impact through stand structural complexity. We also
concluded that the multi-species forest, relative humidity, and soil phosphorus positively
affect AGB. Soil phosphorus affects AGB directly and indirectly through stand structural
diversity. Our study suggested that relative humidity, elevation, and soil primarily influ-
ence the AGB. CVD played an essential role in AGB, while SR did not significantly affect
AGB productivity in different forest types.
This study suggests that aboveground biomass is mainly determined by stand struc-
tural complexity, followed by positive indirect effects on climatic water availability and
soil fertility via stand structural complexity. Species diversity has a nonsignificant direct
impact on aboveground biomass but a significant positive indirect effect via stand struc-
tural diversity. This study also suggests high species diversity is essential for conserving
biodiversity and maintaining stand structural complexity for high forest functioning.

Supplementary Materials: The following supporting information can be downloaded at: https:
//www.mdpi.com/article/10.3390/su15097523/s1, Table S1. Summary relationship of the linear
SEM for founding the relationship amongst AGB, CVD, soil (P), SR, mrh and elevation in sub- tropical
thorn forest of Pakistan. The significant relationship is indicated by bold (p < 0.05). Table S2. The
summary relationship of the linear SEM for founding the relationship amongst AGB, CVD, soil (P),
SR, mrh and elevation in subtropical broad leaved forest of Pakistan. The significant relationship
indicate by bold. Which p < 0.05. Table S3. The summary relationship of the linear SEM for founding
the relationship amongst AGB, CVD, soil (P), SR, mrh and elevation in moist temperate mix forest
of Pakistan. The significant relationship indicates by bold. Which p < 0.05. Table S4. The summary
relationship of the linear SEM for founding the relationship amongst AGB, CVD, soil (P), SR, mrh
and elevation in dry temperate conifer forest of Pakistan. The significant relationship indicates
by bold. Which p < 0.05. Table S5. The summary relationship of the linear SEM for founding the
relationship amongst AGB, CVD, soil (P), SR, mrh and elevation in dry pure pines forest of Pakistan.
The significant relationship indicates by bold. Which p < 0.05. Table S6. Descriptive statistics of
the explanatory variables. Table S7. The summary relationship of the linear SEM for founding the
relationship amongst AGB, CVD, soil (P), SR, mrh and elevation in single species forest of Pakistan.
The significant relationship indicates by bold. Which p < 0.05. Table S8. The summary relationship of
the linear SEM for founding the relationship amongst AGB, CVD, soil (P), SR, mrh and elevation in
Multi species forest of Pakistan. The significant relationship indicates by bold. Which p < 0.05.
Sustainability 2023, 15, 7523 13 of 16

Author Contributions: S.A.: Investigation, Methodology, Software, Writing—original draft. S.M.K.:

Supervision, Conceptualization, project administration, Writing—review & editing. Z.A.: Formal
analyses, Methodology, Writing—review & editing, A.A.: review & editing. N.K.: Methodology,
Facilitation in fieldwork, I.N.: Methodology, Writing—original draft, K.F.A.: Funding acquisition,
Writing—review & editing, G.D.A.-Q.: Funding acquisition, Writing—review & editing, E.F.A.:
Funding acquisition, Writing—review & editing. All authors have read and agreed to the published
version of the manuscript.
Funding: The authors would like to extend their sincere appreciation to the Researchers Supporting
Project Number (RSPD2023R561), King Saud University, Riyadh, Saudi Arabia. The authors are
also thankful to WWF Pakistan for providing financial assistance under the project “Linking Forest
Diversity, Structure and Functions along the Climate and Soil Conditions across Pakistan”.
Institutional Review Board Statement: Not applicable.
Informed Consent Statement: Not applicable.
Data Availability Statement: Data will be available upon special request from the first and corre-
spondence authors.
Acknowledgments: This paper is a part of PhD study conducted by the first author Shahab Ali at
Plant Ecology and Conservation Lab., Quaid-i-Azam University Islamabad, Pakistan. The authors
would like to extend their sincere appreciation to the Researchers Supporting Project Number
(RSPD2023R561), King Saud University, Riyadh, Saudi Arabia. The authors are also thankful to the
WWF Pakistan for providing financial assistance for the field work through their approved project
titled “Linking Forest Diversity, Structure and Functions along the Climate and Soil Conditions
across Pakistan”. The authors also thank Shahfahad Ali Shah, Department of Economics, Quaid-
i-Azam University of Islamabad, Pakistan for providing assistance in the statistical analyses and
data interpretation.
Conflicts of Interest: The authors declare no conflict of interest.

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