J. For. Res.

(2021) 32:649–662
https://doi.org/10.1007/s11676-020-01239-y

ORIGINAL PAPER

The influence of environmental factors on species composition

and distribution in a community forest in Northern Thailand
Siriluck Thammanu1,2 · Dokrak Marod3 · Hee Han4 · Narayan Bhusal1 ·
Lamthai Asanok5 · Pipat Ketdee2 · Noppakoon Gaewsingha6 · Seunghyun Lee4 ·
Joosang Chung1,7

Received: 26 April 2020 / Accepted: 31 July 2020 / Published online: 30 October 2020
© The Author(s) 2020

Abstract Understanding the environmental factors that characterize the difference in biological diversity among the
influence tree species composition is essential for suc- stands. Canonical correspondence analysis (CCA) was used
cessful management of biodiversity and sustainable use of to investigate the environment factors affecting such differ-
community forest resources. This study aims to assess tree ences in biodiversity of the stands. The results showed a high
species composition and distribution in the deciduous Ban diversity of trees in the forest as 197 species, 144 genera,
Mae Chiang Rai Lum Community Forest in Northern Thai- and 62 plant families were recorded. The CCA ordination
land and to analyze the influence of environmental factors identified the environmental factors—the most important
on tree biodiversity in the forest. We conducted a stratified of which were elevation, distance to streams, soil moisture,
systematic sampling of the forest’s total area of 3925 ha, and organic matter, and distance to communities—that signifi-
twenty-five 0.16 ha survey plots were established in three cantly influenced the diversity and distribution of tree spe-
different stands of the deciduous forests to estimate and cies (p < 0.05) in the community forest. Our findings indi-
cate that the implementation of drought reduction measures
Project funding: The work was supported by the Southeast such as building check dams, fire protection, and monitoring
Asian Regional Center for Graduate Study and Research in community forest-product usage would be recommended to
Agriculture (SEARCA) - ASEAN Working Group on Social further biodiversity conservation and the sustainable use of
Forestry Strategic Response Fund (ASRF) under the ASEAN- community forest resources.
Swiss Partnership on Social Forestry and Climate Change and
by the R&D Program for Forest Science Technology (Project
No. 2018113C10-2020-BB01) provided by Korea Forest Service Keywords Community forest management · Species
(Korea Forestry Promotion Institute). biodiversity · Canonical correspondence analysis · Ban
Mae Chiang Rai Lum community forest · Northern
The online version is available at http://www.sprin​gerli​nk.com.
Thailand
Corresponding editor: Yu Lei.

Electronic supplementary material The online version of this

article (https​://doi.org/10.1007/s1167​6-020-01239​-y) contains
supplementary material, which is available to authorized users.

4
* Hee Han Department of Forest Policy and Economics, National
heehan@korea.kr Institute of Forest Science, Seoul 02455, Republic of Korea
5
* Joosang Chung Department of Agroforestry, Maejo University, Phrae
jschung@snu.ac.kr Campus, Phrae 54140, Thailand
1 6
Department of Forest Sciences, Seoul National University, Department of National Parks, Wildlife and Plant
Seoul 08826, Republic of Korea Conservation, Bangkok 10900, Thailand
2 7
Royal Forest Department, Bangkok 10900, Thailand Research Institute of Agriculture and Life Sciences, Seoul
3 National University, Seoul 08826, Republic of Korea
Department of Forest Biology, Faculty of Forestry, Kasetsart
University, Bangkok 10900, Thailand

13
Vol.:(0123456789)
650 S. Thammanu et al.

Introduction Zhao et al. 2015). For example, higher soil sand ratio less-
ens water-holding capacity that can lead to water stress on
In Southeast Asia, Community Forest Management (CFM) trees (Aranguren et al. 1982; Toledo et al. 2012; Zhang et al.
has been recognized as an effective approach to restore 2013), and acidity levels affect the distribution of species
degraded forests, augment the income of the poor, and help and is linked to slope and elevation in lowland tropical for-
meet the needs of local residents for food and medicine ests (Nguyen et al. 2015; Vahdati et al. 2017). Soil moisture
(Blair and Olpadwala 1988). CFM has been applied in pur- also significantly changes the growth patterns of trees in
suit of sustainability and forest biodiversity conservation in drought areas (Fu et al. 2004; Yoshifuji et al. 2006; Asanok
Thailand (Srisutham and Kaewjampa 2010) as the Royal and Marod 2016; Tilk et al. 2017).
Forest Department (RFD) has been promoting CFM since In addition, the organic matter in the soil is relevant to
1987. To date, CFM has been implemented in approximately an analysis of environmental factors and plant communi-
7% of the country’s total forest area, or more than 1.2 million ties in the forest (Sarker et al. 2014; An et al. 2015; Zhang
ha, within roughly 17,400 villages nationwide (Royal Forest et al. 2016). Soil nutrients such as nitrogen, potassium,
Department 2014, 2020). phosphorus, calcium, and magnesium are correlated with
Thailand is one of the most bio-diverse countries in the richness and distribution of plant species in tropical for-
Southeast Asia. Forests cover approximately 32% of Thai- ests (Paulo et al. 2007; Zhang et al. 2013; Tilk et al. 2017).
land’s total land area (Royal Forest Department 2019) and Further, the human impact of resource utilization, road con-
are inhabited by roughly 8% of the world’s plant species struction, and residential development significantly affects
(Office and Natural Resources and Environmental Policy plant diversity (Chen et al. 2014; Måren and Sharma 2018;
and Planning 2009). However, a decline of forest biodiver- Eghdami et al. 2019).
sity has become a critical issue. There are 1442 threatened Overall, the environment’s impact on the biodiversity
plant species in the country, 715 of which have been clas- in community forests is complex as it involves a multitude
sified as vulnerable, 207 as endangered and 18 as critically of factors and considerations that provide limited answers,
endangered (Department of National Parks, Wildlife and presents additional clues and raise new questions that are
Plant Conservation 2017). Participation by local people in crucial to a more thorough and useful understanding of the
the management of forest resources has been identified as relationship. This study sought to increase awareness of
a promising way to address the issue of biodiversity loss in CFM’s role in the furtherance of biodiversity conservation
the country. and to seek additional knowledge to support these efforts. To
Although community forest management can facilitate do so, we identified ecological characteristics, ascertained
biodiversity conservation, there is insufficient information species diversity indices, and investigated pertinent environ-
informing how CFM affects biodiversity and how environ- mental factors impacting three unique forest stands in the
mental factors influence that biodiversity in community community forest. We focused on the following questions:
forests. There are numerous factors that may impact the dis- (1) what is the diversity and composition of tree species in
tribution of trees in community forests. the community forest? (2) what are key environmental fac-
Climatic factors can threaten the extinction of vulner- tors influencing species distribution in the different stands?
able species (e.g. atmospheric pressure, temperature, pre- and (3) what resulting information could prove to be use-
cipitation, solar radiation, etc.) (Körner 2007; Diem et al. ful in CFM to protect species diversity and its benefits to
2018). Climatic factors such as temperature and precipita- livelihoods?
tion are key drivers to control species distribution directly
when they exceed the ecophysiological tolerances of species
and affect photosynthetic activity and biological processes Materials and methods
directly (Rowe 2009). Topographic features such as slope,
aspect, and elevation can impact local climate as well as Study area
soil conditions that in turn have varied effects on vegetation
structure (Zhang et al. 2006; Zhang and Zhang 2007; Zhang The Pa Mae Phrik National Forest Reserve is located in
et al. 2013, 2016). The relative distance from a water source Thailand’s northern province of Lampang. Situated in the
can also affect the composition and distribution of woody southern region of this Forest Reserve is the Ban Mae Chi-
vegetation because of the resulting varying amount of water ang Rai Lum Community Forest (17°22′48″ N–17°27′47″ N
available for growth (Marod et al. 1999; Scalley et al. 2009; and 99°00′47″ E–99°05′48″ E), where in this study was con-
Tavili et al. 2009; Sarvade et al. 2016; Asanok et al. 2017). ducted (Fig. 1).
Physical and chemical soil properties can inform vegeta- This community forest has a total area of 3925 ha, is
tion patterns on a local scale (Oliveira-Filho et al. 1998; at an elevational range of 140–660 m and is composed
Hejcmanovā-Nežerková and Hejcman 2006; Han et al. 2011; of mixed deciduous and dry dipterocarp forest subtypes.

13
The influence of environmental factors on species composition and distribution in a community… 651

Fig. 1  Location map of the Ban Mae Chiang Rai Lum Community Forest in Northern Thailand

Deciduous forests occur in areas with 5–6 month dry and plays a vital role as a source of biodiversity in provid-
seasons, with a mean annual rainfall of 1000–1500 mm ing non-timber forest products (NTFPs) while contributing
in dry dipterocarp forests and 1000–1800 mm in mixed to ecosystem services beneficial to livelihoods. However,
deciduous forest (Bunyavejchewin 1983; Santisuk 1988). unchecked tapping into the forest resources is widespread
Mixed deciduous forests typically grow on moderate fer- and creating a potential existential threat to the forest.
tile loam soil while dry dipterocarp forests have more Northern Thailand in general, and Lampang in spe-
sandy lateritic soil (Bunyavejchewin 1983, 1985; Santisuk cific, have unique climatic features (Diem et al. 2018).
1988). Forest fires are a regular feature of dry season in The climate around the study area features a wet season
deciduous forests (Sukwong and Dhamanittakul 1977; from April to October and a dry season from November to
Bunyavejchewin 1983). A previous study identified Xylia March. Historically, this region also experiences drought
xylocarpa, Schleichera oleosa, Sindora siamensis, Shorea conditions. Temperatures ranged from a minimum of
obtusa, Terminalia mucronata as the dominant species in 31.7 °C (January and November) to a maximum of 37.1 °C
the study area (Royal Forest Department 2017). This com- (March). The mean annual temperature, relative humidity,
munity forest has been locally managed in collaboration and rainfall were 33.6 °C, 76.1%, 1129.4 mm, respectively
with the RFD under a community forest project since 2008 (Thai Meteorological Department 2018).

13
652 S. Thammanu et al.

Field survey vary significantly during the dry season. In addition, the
dry season often featured forest fires. As such, data on tem-
A field survey was conducted in the forest from July to perature, relative humidity and light intensity were collected
October 2018. Results of a 2016 study conducted by the for 92 days during the wet season from July 11 to October
RFD in the same area were used to calculate the sampling 11, 2018.
intensity with a confidence interval of 95%. An estimate of Data regarding 15 environmental factors were collected
the sampling was obtained by using the standard deviation from the 25 sampling plots. Those factors were broadly clas-
from previous surveys (Avery and Burkhart 1983; Asrat and sified into topographic, edaphic, and anthropogenic. The
Tesfaye 2013). The formula was expressed as: topographic factors were elevation (EL), slope (SL), and
aspect (AS), and the data were extracted out of 1:50,000
n = (Z𝜎∕E)2 (1) topographic maps produced by 20 m contour lines using the
where n is the sampling intensity; Z is the z-value for the ArcGIS. The edaphic factors were investigated from samples
confidence interval of 95%; σ is the density of tree species, collected at a depth of 15–20 cm in three random locations
and E is the percent standard deviation of the precision within each 40 m × 40 m plot during the rainy season from
required. June to July 2018. Acidity (pH), organic matter (OM), soil
A total of 25 sampling plots of 40 m × 40 m (0.16 ha) texture [sand (SA), silt (SI), clay (CL)], available phospho-
were surveyed through a systematic sampling method (Asia rus (P), exchange potassium (K), exchange calcium (Ca), and
Network for Sustainable Agriculture and Bioresources soil moisture (SM) in the samples were analyzed. Soil pH
2010). In each plot, trees with a diameter at breast height was measured by 1:1 soil/water ­mol−1 KCL (National Soil
(DBH) ≥ 4.5 cm were identified and measured in every Survey Center 1996). OM was estimated by wet digestion
10 m × 10 m sub-quadrat. Within 10 m sub-quadrats, sam- and titration using Walkley–Black method (Nelson and Som-
plings with DBH < 4.5 cm and height ˃ 1.30 m were recorded mers 1996). Soil texture was determined by Pipette method
in 4 m × 4 m sub-quadrats, whereas seedlings were docu- (Gee and Bauder 1986). P was measured by Bray II (Bray
mented in 1 m × 1 m sub-quadrats within each 4 m sub- and Kurtz 1945), while K and Ca were extracted with 1 M
quadrat (Fig. 2). NH4OAc at pH 7.0 (Thomas 1982), respectively. The soil
moisture analysis was performed by oven drying at 105 °C
(National Soil Survey Center 1996). ArcGIS’s Euclidean
Data collection on environmental factors distance tool was used to calculate the distance to streams
(DS), distance to roads (DR), and distance to communities
Climate data were generated by Hobo U23-001 and UA-002- (DC) from the sampling plot.
64 which operated in six stations located in plots 3, 10, 13,
15, 17, and 24. The monitoring equipment was spread out Data analysis
and placed in these representative plots to provide the opti-
mal coverage to accumulate climate data from throughout The data analysis consisted of examining the biologi-
the study area. Historically, relevant climatic data did not cal diversity of the community forest and exploring

40 m

40 m

10 m × 10 m (Tree) 10 m
4 m × 4 m (Sapling)
1 m × 1 m (Seedling)

10 m

Fig. 2  Sampling plots and sub-quadrats for identification and measurement

13
The influence of environmental factors on species composition and distribution in a community… 653

environmental factors that affect tree species composition Species compositions in the three forest stands were com-
and distribution. pared using the Jaccard similarity index (Magurran 1988).
The forest stands in the community forest were identi- The correlations between environmental factors were
fied by cluster analysis employing importance value index determined through Pearson correlation analysis. In case
(IVI) matrices in each sampling plot. A cluster analysis of a high correlation in which a matrix of two independent
requires pruning of the dendrogram at a level represent- variables has correlation coefficients (r) > 0.7 across all
ing a compromise between the group and the number of environmental factors, only one of the factors was selected
groups. Optimum pruning for the dendrogram was selected for the ordination analysis (Sarker et al. 2014).
by applying the Relative Sorensen Distance and Ward’s The relationship between tree species distribution
Linkage Method (McCune and Grace 2002). and environmental factors was finally assessed using the
To analyze the ecological characteristics of the three canonical correspondence analysis (CCA) (McCune and
stands, we identified mature trees (DBH ≥ 4.5 cm) and cal- Grace 2002). The matrices between the IVI of mature trees
culated the stand density and basal area of individual tree (DBH ≥ 4.5 cm) in each stand and the environmental fac-
species in each stand type. tors were analyzed in the CCA ordination. The correlation
Tree species diversity in each individual stand was significance between the matrices was determined using
analyzed, and a comparison of stand-to stand diversity the Monte Carlo test with 999 permutations. Only the
was examined. A rarefaction approach was implemented environmental factors significantly related (Monte Carlo
in order to compare the diversity from the three different permutation test, p < 0.05) to the mature tree species were
data-source stands (Colwell et al. 2012). Individual based retained. All the statistical calculations were performed
interpolation and extrapolation of Hill numbers (q = 1, 2, using PC.ORD version 5.10 (McCune and Mefford 2006)
3) (Hill 1973) were carried out for each stand and the spe- and the R program version 3.5.1 for Windows software (R
cies diversity among the three forest stands was estimated Development Core Team 2019).
as follows: species richness (q = 0), Shannon diversity
(the exponential of Shannon entropy, q = 1), and Simpson
diversity (the inverse of Simpson concentration, q = 2). Results
The ecological importance of the tree species in each
stand was quantified through the IVI using the equation: Forest stand classification
IVI = R.D + R.F + R.Do (2) The resulting dendrogram reflected 5.83% chaining and was
where R.D is the relative density of the tree species; R.F cut with 25% of the remaining information explained by
is their relative frequency, and R.Do is their relative domi- three stand types: Millettia leucantha-Lagerstroemia duper-
nance. They were calculated as R.D = number of individu- reana stand (MLS), Shorea siamensis-Shora obtusa stand
als of the species × 100/total number of quadrate studies, (SSS), and S. obtusa-S. siamensis stand (SOS) (Fig. 3).
R.F = number of quadrates in which species occurred × 100/
total number of quadrate studies, and R.Do = total basal area Climate data
of species × 100/total basal area of all the species (Curtis and
McIntosh 1951). Relative humidity and light intensity data did not vary sig-
To investigate the environmental factors that affect tree nificantly plot-to-plot, but temperature data was significantly
species in the community forest, a dataset of 15 factors different (p < 0.001). The average temperature during the
was tested for normality using the Shapiro–Wilk test for 92-day collection period varied between 25.09 °C (plot
an alpha level of 0.05. The abnormal data was transformed 17) and 27.37 °C (plot 10). The average relative humidity
to a normal distribution to improve the linearity. In this ranged from 79.19% (plot 15) to 85.10% (plot 17), and the
study, AS, SI, SM, DR, and DC were normally distributed average light intensity ranged from 3011.67 l × (plot 15)
(p > 0.05). Ten factors had values less than the chosen to 4698.48 l × (plot 17). The overall average temperature,
alpha level; log10(y) was used to transform SL, pH, OM, relative humidity, and light intensity throughout the study
CL, K, and Ca, and (y)^3, 1/(y), sqrt(y), and − 1/sqrt(y) area were 26.85 °C, 82.23%, and 3549.09 l ×, respectively
were used to transform SA, EL, DS, and P, respectively. (Fig. 4).
The differences in stand characteristics, such as stand
density, basal area, and the environmental factors among Species composition and diversity
the three stand types were subjected to a one-way analysis
of variance (ANOVA) with a post hoc Tukey’s honestly The field survey recorded 18,567 trees comprising 197 spe-
significant difference (HSD) test (Zhang et al. 2016). cies (129 tree, 99 sapling, and 141 seedling species), 144
genera, and 62 plant families. The Jaccard index showed a

13
654 S. Thammanu et al.

SOS

SSS

MLS

Fig. 3  Classification of the stand types in the deciduous area of the Ban Mae Chiang Rai Lum Community Forest in Northern Thailand.
MLS = Millettia leucantha-Lagerstroemia duperreana, SSS = Shorea siamensis-Shora obtusa, SOS = Shorea obtusa-Sindora siamensis

21.29% similarity (33 species) between the MLS and SSS, in the MLS, followed by the SSS and the SOS, respectively.
a 36.30% similarity (53 species) between the SSS and SOS, The Shannon diversity and Simpson diversity also estimates
and a 17.78% similarity (24 species) between the MLS and are highest in the MLS, the SSS, and the SOS, in descending
SOS. The forest had an average density of 966 trees h­ −1 order (Fig. 5).
and an average basal area of 16.74 m2 ­h−1 (Table S1). The
one-way ANOVA test showed that the average density were
significantly different stand-to-stand (p < 0.01), though aver- Relationship between environmental factors and species
age basal area did not vary significantly. diversity
The five species with the highest IVI in the MLS were M.
leucantha, L. duperreana, M. brandisiana, Antidesma soot- The mean values and standard deviation of the 15 environ-
epense, and Pterocarpus macrocarpus. The most dominant mental factors analyzed in the Ban Mae Chiang Rai Lum
tree species in terms of its IVI in the SSS was S. siamensis, Community Forest are shown in Table S4. The degree of
followed by S. obtusa, X. xylocarpa, T. mucronata, and Dip- soil acidity (pH), organic matter (OM), silt (SI), clay (CL),
terocarpus tuberculatus. S. obtusa was the most common exchange potassium (K), soil moisture (SM), distance to
species in the SOS, and it included S. siamensis, X. xylo- roads (DR), and distance to communities (DC) were greater
carpa, Canarium subulatum, and Ellipanthus tomentosus in the MLS than in the SSS and SOS. The values of elevation
(Table S2). (EL), slope (SL), available phosphorus (P), and exchange
The rarefaction-produced tree species diversity values in calcium (Ca) were highest in the SSS, and in decreasing
different sample sizes of three forest stands are shown in order in the MLS and SOS. The value of aspect (AS), dis-
Table S3. A comparison of extrapolated species diversity tance to streams (DS), and sand ratio (SA) were greater in
shows that species richness was estimated to be the highest the SOS than in the SSS and MLS.

13
The influence of environmental factors on species composition and distribution in a community… 655

Fig. 4  Average a temperature (°C), b relative humidity (%), and c light intensity (lux)

among the stands. The post hoc Tukey’s HSD test iden-
tified that the CL value was different between the MLS
and the SOS, between the SOS and the SSS, and between
the SSS and the MLS. OM and SA differed substantially
between the MLS and the SOS, but they did not differ in
the SSS. Similarly, SL differed between the SOS and the
SSS, but it did not differ in the MLS. K varied when com-
paring the MLS with the SOS and the SSS with the SOS.
A matrix of two independent environmental factors
showed that there were high correlation coefficients
(r > 0.7) between the factors (Table S5). OM was posi-
tively correlated with K (r = 0.932). SA was positively
associated with EL (r = 0.704), but negatively associated
with SI (r = − 0.890), CL (r = − 0.929), DR (r = − 0.746),
and K (r = − 0.724). CL positively correlated to DR
Fig. 5  Comparison of individual-based rarefaction and extrapolation (r = 0.710), but negatively correlated to EL (r = − 0.707).
of species diversity for Hill numbers among three forest stands: spe- EL negatively related to SL (r = − 0.740). DC was posi-
cies richness (q = 0), Shannon diversity (q = 1), and Simpson diversity tively related with DR (r = 0.853). Considering those
(q = 2)
correlations between the environmental factors, the ten
variables, EL, AS, DS, pH, OM, P, Ca, SM, SI, and DC,
Five factors differed significantly among the three stand were selected for further analysis using CCA. In the pre-
types; SL, OM, SA, CL, and K differed from one stand liminary CCA, two poorly correlated factors, Ca and SI,
type to another (p < 0.05). There was no significant dif- were eliminated; the remaining eight factors were used in
ference in EL, AS, DS, pH, SI, P, Ca, SM, DR, and DC the final CCA.

13
656 S. Thammanu et al.

Table 1  Results of the canonical correspondence analysis (CCA) on (r = 0.163), EL (r = 0.125), and P (r = 0.061). The second
the relationship between the distribution of tree species and the envi- axis revealed a negative correlation with DC (r = − 0.513),
ronmental factors
OM (r = − 0.398), DS (r = − 0.281), pH (r = − 0.142), and
CCA results Axis 1 Axis 2 Axis 3 AS (r = − 0.140), but it displayed a positive correlation with
EL (r = 0.810) and SM (r = 0.060). The comparison among
Eigenvalues 0.531 0.363 0.239
the environmental factors showed that pH was positively
Variance in species data
correlated with OM (r = 0.563) and P (r = 0.547) as was OM
% of variance explained 13.5 9.2 6.1
with P (r = 0.604) (Table S5).
Cumulative % explained 13.5 22.7 28.8
The CCA results demonstrate the relationships between
Pearson correlation (species- 0.924 0.917 0.777
environment) environmental factors and species distribution (Figs. 6 and
Monte Carlo test 0.015* – – 7). In Grouping 1, the EL was an important factor in spe-
cies distribution in plots 2, 9, and 11 of the SSS and in plots
*p < 0.05, **p < 0.01, ***p < 0.001 1, 7, 8, and 12 of the SOS. The abundant species in this
grouping were Atalantia monophylla (ATMO), Antidesma
Table 2  Correlation coefficients between the eight environmental ghaesembilla (ANGH), Gluta usitata (GLUS), Bauhinia
factors with the first three axes of the CCA​
bracteata (BABR), Flacourtia indica (FLIN), Memecylon
Environmental Factors Abbreviation Axis 1 Axis 2 Axis 3 edule (MEED), and Flacourtia rukam (FLRU). In Grouping
2, DS was linked to Ochna integerrima (OCIN), Syzygium
Soil acidity pH −0.158 −0.142 0.389
cumini (SYCU), Madhuca dongnaiensis (MADO), Morinda
Organic matter OM −0.546* −0.398* 0.068
elliptica (MOEL), and Tectona grandis (TEGR) in plots 6
Available phosphorus P 0.061 −0.270 0.388
and 10 of the SSS and 3, 13, 21, and 23 of the SOS. This
Soil moisture SM −0.470* 0.060 0.196
suggests that these species can be impacted by their dis-
Elevation EL 0.125 0.810*** −0.268
tance from streams. Regarding Grouping 3, SM, OM, and
Aspect AS 0.163 −0.140 −0.259
DC were closely related to Polyalthia cerasoides (POCE),
Distance to streams DS 0.409* −0.281 −0.125
Zollingeria dongnaiensis (ZODO), Hymenodictyon orix-
Distance to commu- DC −0.402* −0.513** 0.435*
nities ense (HYOR), Zizyphus oenoplia (ZIOE), P. macrocarpus
(PTMA), Mitragyna rotundifolia (MIRO), Lannea coroman-
*p < 0.05, **p < 0.01, ***p < 0.001
delica (LACO), Bridelia retusa (BRRE), and Wendlandia
tinctoria (WETI) in plots 14, 15, and 20 of the MLS and in
The IVI values (i.e. R.D, R.F, and R­ Do) of each of the 129 plot 24 of the SSS. This indicates that they were restricted
mature tree species and the eight explanatory environmental by moisture content, limited by OM, and closely correlated
factors (pH, OM, P, SM, EL, AS, DS, and DC) in the 25 to human disturbance.
sampling plots were subjected to the CCA. The results of the
CCA showed strong correlations between the distribution
of tree species and the environmental factors (Table 1). The
eigenvalues obtained for the first and second axes were 0.531 Discussion
and 0.363, respectively. The Pearson correlation showed that
the environmental factors had a high correlation to the spe- The Ban Mae Chiang Rai Lum Community Forest was
cies for axis 1 (0.924) and axis 2 (0.917). Therefore, these divided into three forest stands: the mixed deciduous MLS
axes were considered as good indicators of relationships and the dry dipterocarp SSS and SOS (Fig. 3). A rarefaction
between the environmental factors and the species distribu- prediction of species diversity (Fig. 5 and Table S3) and
tion in the community forest. The Monte Carlo permutation the wide array of species, genera, and families (Table S1)
test confirmed that the first axis explained the significant recorded via the field survey indicate remarkable diversity,
variation in the community forest (p < 0.05). The signifi- similar to other forests of Northern Thailand (Popradit et al.
cance of the second and third axes was not reported for the 2015; Khamyong et al. 2018). There were no noteworthy
reason that a simple randomization test for these axes could differences in the average basal areas, but average density
bias the p values (McCune and Mefford 2006). varied significantly among the three stand types. There was
The correlations between the environmental factors and a degree of species similarity among the stands whereas
the CCA axes are listed in Table 2. OM (r = − 0.546) had the the forest structures and species compositions were distinct
highest negative correlation coefficient with the first axis, (Table S2), In addition, the importance of the environment
and SM (r = − 0.470), DC (r = − 0.402), and pH (r = − 0.158) factors varied stand-to-stand (Table S4) as did their degree
were also negatively correlated with it. Conversely, DS of correlation (Table S5).
(r = 0.409) had a positive correlation, followed by AS

13
The influence of environmental factors on species composition and distribution in a community… 657

O LAMA LASP

Axis 2
OVI ALSA
MILEP BECO
TEBE
R
DAFO DIMOG
Grouping 1 POOB
SAKO
DACA
FIIN
ATMO
SICE ANGH BASC
ALLE
MIBR EL BERO
BAGL TEAL GLUS
STNE
BABR
SCOL MEED LIGL HOIN
FLIN FLRU GRER
ALOD LAVI SISI
TEMU
DICA
CRCO TECH MOCO
GAPI MACA ELTO AZIN DADO
XYXY CASU
MAST CAPA ROWI
STNU HACO GAOB

SM DIEH
DIOBH DAOL Axis 1
ADVI
HYOR POCE MASI SHOB
HAPE BARA IRMA CAGR APVI
CAJA ZIOE LACO MOEL TEGR PHEM
VICA CASE
MISP ZODO OCIN
WRAR ADPA LADU MIRO BRRE
SYCU
CAGO ANSO ELLA
MIHO
FEAD
BOAN
MILEK
WATR
PTMA
BULA
MADO DS
CHTA SPPI SIMU
CRFO
OM WETI SHSI
VILI LODU
DC DITU

ANACR BUGL
MESU
Grouping 2 VIPI
DACU
CRHU
VIPE
Grouping 3 SEAL
ROSO
LIPO APNI
CATO
DIOBT
PATO
PRLA SEGA

TRBU

GASO QUKE

Fig. 6  The CCA ordination diagram representing the relationship between the distribution of tree species and the environmental factors. Full
names of tree species by abbreviation are listed in Table S6

Our results represent that the heterogeneity of species distribution can be limited by a combination of environmen-
distribution patterns among the different forest stand types tal factors across distinct forests.
could be a response to a variation in the environmental fac- Among the five significant factors (i.e. EL, DS, SM, OM,
tors; distinct environmental conditions in forest stands can and DC), EL appeared to be the most influential on species
be effective indicators of species distribution in the Ban Mae distribution. The IVI of mature trees was positively related
Chiang Rai Community Forest in Northern Thailand. to elevation in the SSS and SOS; elevational gradients pro-
The results of the CCA demonstrated that species com- mote not only species diversification but also soil differen-
position and distribution of the mature trees were related to tiation (Zhang and Zhang 2007; Zhang et al. 2013, 2016).
topographic, edaphic, and anthropogenic factors (Table 1). Though, climatic factors can vary with elevation (Zhang
The environmental factors showed both positive and nega- et al. 2006; Körner 2007), the elevational features of dry dip-
tive impacts on species diversity. Specifically, EL, DS, SM, terocarp in Thailand occur mainly in elevations up to 900 m
OM, and DC were the most evidently linked to species dis- (Bunyavejchewin et al. 2011). At this elevation, there is no
tribution (Table 2). Moreover, environmental factors had drastic change in lowland vegetation similar to that found in
different effects on species distribution between stand types most tropical zones at elevations above 1000 m, including
(Figs. 6 and 7). This indicates that species composition and in the North of Thailand (Küchler and Sawyer 1967). Thus,

13
658 S. Thammanu et al.

Fig. 7  The five most impor-

tant environmental factors
explaining the characteristics SSS-11

Axis 2
of different stand types in the
Grouping 1
CCA ordination: elevation (EL), MLS-5
SSS-9
distance to streams (DS), soil
SOS-7
moisture (SM), organic matter SOS-8
(OM), and distance to com- EL
munities (DC) affecting 25 plots SOS-1
in the three stands (Δ): Millettia SSS-2 SOS-22
SOS-12
leucantha-Lagerstroemia duper-
reana stand (MLS), Shorea SOS-3
siamensis-Shora obtusa stand SOS-13
SOS-23 SOS-21
(SSS), and Shorea obtusa-Sin-
dora siamensis stand (SOS) SM SSS-10 Axis 1
SSS-6
MLS-20
MLS-14
SSS-24 DS
MLS-15
OM
DC
Grouping 2 SOS-25
SSS-19
SSS-4
Grouping 3
SSS-18
SSS-16

SSS-17

the various soil properties unique to elevation could be key correlated whereas SA was positively correlated (Table S5).
to the composition and distribution of tree species in the Poor sandy soil is an important driver for tree species in dry
dipterocarp forest stands this study area (Bunyavejchewin dipterocarp forests (Bunyavejchewin 1983, 1985; Santisuk
et al. 2011). 1988). Therefore, some tree species in the SSS and the SOS
Other tree species associated with dominant species were may be able to grow at higher elevation because of more
also found in the dry dipterocarp stand. This suggests that favorable soil conditions without the limiting climatic condi-
higher elevation promotes species diversity in dry diptero- tions found at extremely high elevation.
carp forests. Teejuntuk et al. (2003) reported that the diver- In this study, it appears that DS and SM influenced spe-
sity of tree species in Doi Inthanon National Park, Northern cies diversity and distribution. The positive correlation
Thailand increased up to an elevation of about 1800 m. Also, between species diversity in the SSS and SOS with the DS
it has been found that tree species in dry dipterocarp forests revealed that diversity is related to water availability. This
can move into higher land to grow with other tree species in indicates that dry conditions may negatively impact spe-
an ecotone zone (Kutintara 1975; Marod et al. 2015) which cies diversity and distribution. Several species in the dry
may be related to a more favorable environment for growth dipterocarp forests are strongly associated with proximity
at slightly higher elevation (Ivanov et al. 2008). to water sources. This suggests that tree species adapt to
Elevational gradients often limit the distribution of tree drought areas that have low soil moisture, high aridity, and
species by decreasing soil moisture and nutrient levels fire disturbance. These are important factors that determine
(Bridge and Johnson 2000). Several studies have reported the species occurrence in dry dipterocarp forests in Thailand
that an increase in species diversity may have an elevational (Rundel and Boonprakob 1995; Marod et al. 2002; Bunyave-
limit above which diversity would become negatively cor- jchewin et al. 2011).
related (Liu et al. 2018). For instance, the resources neces- Similarly, the negative correlation of species diversity
sary for plant growth are limited at extremely high eleva- in the MLS and SSS with SM substantiated the occurrence
tions because of strong winds and shallow soils (Zhang et al. of these forests in dry regions (Marod et al. 1999). Tree
2013). species such as P. macrocarpus (PTMA), P. cerasoides
In this study area, elevation ranged between 140 and (POCE), Z. dongnaiensis (ZODO), H. orixense (HYOR),
660 m. Higher elevation and SI, CL, and K were negatively and L. coromandelica (LACO) were closely associated

13
The influence of environmental factors on species composition and distribution in a community… 659

with SM, which suggests that they were restricted by mois- Plant productivity is positively linked with OM, as
ture levels and sensitive to drought conditions. Generally, reported by Bauer and Black (1994), thus, more OM should
these species had a higher water demand than other tree be made available for plants to increase their productivity.
species in deciduous forests that could adapt and survive Our findings also revealed that OM had a positive correla-
in arid areas (Marod et al. 1999). Previous findings indi- tion with pH and P (Table S5), indicating that trees are not
cate that SM decreased as the distance from a reservoir only affected by OM content but that pH and P are major soil
increased (Sarvade et al. 2016), suggesting that the spe- properties involved in plant growth. In a previous study con-
cies in MLS and SSS were also related to distance from ducted in a tropical forest in Ben En National Park, Vietnam,
a water source. it was reported that an increase in OM could augment P and
Deciduous forests often occur in arid areas with sandy pH, thereby enhancing species composition and distribution
soil; 53% of the soils in mixed deciduous forests and 70% of of plants in the area (Hoang et al. 2011).
those in deciduous dipterocarp forests consist of sand (Myo The dominant species were closely correlated with DC,
et al. 2016). Sandy soils are more susceptible to leaching supporting a previous report that human disturbance is a
which can lead to drought (Aranguren et al. 1982; Zhang threat to plant diversity loss (Millennium Ecosystem Assess-
et al. 2013). Soils with higher sand content have lower water- ment 2005). Not only were dominant species found distrib-
holding capacity, thereby causing plants to compete more for uted in the stands, but shrub species such as W. tinctorial
soil moisture (Toledo et al. 2012). Contrarily, trees grow- (WETI), M. rotundifolia (MIRO), B. retusa and Z. oenoplia
ing in clay soil have more efficient root system and are less (ZIOE) were also distributed near the communities. These
susceptible to drought stress because of the greater capacity species are often used for firewood and construction. As
of the soil for water retention. Trees with deeper roots in such, there is a high risk of them being harvested thereby
sandy soil have a greater chance of surviving as they adapt affecting their distribution. Furthermore, they are pioneer
to reduce drought stress (Wessel 1971). Hence, a deep root species that play a vital role in ecological succession after
system enables plants to adapt to and survive in extremely disturbances (Department of National Parks, Wildlife and
dry conditions, such as being distant from a water supply or Plant Conservation 2007; Asanok et al. 2020). Utilization by
being in an area with limited soil moisture. In this study, our communities may have an impact on the natural restoration
results imply that trees survive and grow well despite the dry of forest ecosystems in the community forest.
conditions in the SSS and SOS; the sufficient depth of root Many studies have shown that the impact on tree spe-
system of trees in the both stands has enabled them to adapt cies in forests could be worse if they were situated closer
to drought stress. However, the existence some species in to a community (Thapa and Chapman 2010; Hoang et al.
the MLS and SSS may be limited by the level of moisture in 2011; Chen et al. 2014; Asanok et al. 2017; Martínez-
soil for their growth. Camilo et al. 2018). These studies reported that trees would
The negative correlation of the dominant species in the be more abundant in less disturbed plots located far from
MLS and SSS with OM and DC, explains that both environ- human settlements. Furthermore, Teejuntuk et al. (2003)
mental factors constrain tree growth and diversity in their discussed that at an elevation less than 1800 m in North-
corresponding forest stands. The growth, species distribu- ern Thailand, the occurrence of ecologically similar species
tion, and composition of trees in tropical areas are influenced increased at higher elevation since forests at higher elevation
by soil nutrients (John et al. 2007; Santiago et al. 2012). are relatively far and inaccessible to man. Lowland forests,
Many studies have shown that OM is a key environmental conversely, have become significantly more fragmented and
factor in plant communities (Zhang and Zhang 2007; Slik degraded because of deforestation through illegal logging,
et al. 2009; Sarker et al. 2014), and that it contributes to the burning, and grazing. Human disturbance is more rampant in
availability of nutrients and water by improving soil struc- areas with easy access, poor monitoring and weak enforce-
ture and physical conditions, increasing the water holding ment of regulations (Måren and Sharma 2018). The apathy
capacity of the soil, and providing a habitat for plant roots of locals to forest management also aggravates the problem
and soil organisms (Carter 2002; Meng et al. 2014). Thus, and contributes to the decline of forest biodiversity.
soils with more OM are more fertile and more favorable for
the optimal growth of trees (Vahdati et al. 2017).
In the study forest, however, there were tree species in Conclusions
the MLS and SSS that were inversely associated with OM,
indicating that this environmental factor was restricting Our study suggests that elevation, distance to streams, soil
their growth. As explained by Grime (1977), if nutrients are moisture, organic matter, and distance to communities are
scarcely available in the soil, then plants will have to adapt the most important factors influencing species composition
to low-resource and nutrient-poor conditions, making natural and distribution in the deciduous forests of the Ban Mae
selection predominant in the process. Chiang Rai Lum Community Forest in Northern Thailand.

13
660 S. Thammanu et al.

Even through tree species in deciduous forests are able to Aranguren J, Escalante G, Herrera R (1982) Nitrogen cycle of tropical
shift to higher elevation to find more favorable conditions perennial crops under shade trees. Plant Soil 67:247–258. https​://
doi.org/10.1007/BF021​82772​
and adapt themselves to grow in drought areas, the limita- Asanok L, Marod D (2016) Environmental factors influencing tree spe-
tion of organic matter, water availability, and soil moisture cies regeneration in different forest stands growing on a limestone
can still impact tree species diversity. This could prove to hill in Phrae Province, Northern Thailand. JFES 32:237–252.
be important and useful information for community forest https​://doi.org/10.7747/JFES.2016.32.3.237
Asanok L, Kamyo T, Norsaengsri M, Salinla-um P, Rodrungruang
management. We recommend that management practices K, Karnasuta N, Navakam S, Pattanakiat S, Marod D, Dueng-
for drought reduction such as building check dams and fire kae P, Kutintara U (2017) Vegetation community and factors that
protection be implemented to protect biodiversity, especially affect the woody species composition of riparian forests growing
in the MLS and SSS. in an urbanizing landscape along the Chao Phraya River, Cen-
tral Thailand. Urban For Urban Green 28:138–149. https​://doi.
In addition, biodiversity conservation would be better org/10.1016/j.ufug.2017.10.013
served by monitoring the utilization of forest resources by Asanok L, Taweesuk R, Papakjan K (2020) Woody species coloniza-
nearby communities. Species propagation, silviculture, and tion along edge-interior gradients of deciduous forest remnants in
transplantation should be initiated and/or expanded, particu- the Mae Khum Mee Watershed Northern Thailand. Int J For Res.
https​://doi.org/10.1155/2020/58673​76
larly in neighboring villages or private farms, to enhance the Asia Network for Sustainable Agriculture and Bioresources (2010)
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