Edaphic and Climatic Factors Affecting the Distribution of Castanopsis tungurrut (Blume) A. DC (Fagaceae) in Cibodas Biosphere Reserve, Indonesia

Abstract

Castanopsis tungurrut (Blume) A. DC. (Fagaceae) is an Indonesian native plant species with limited spatial distribution; it is found only in Java, Kalimantan, and Sumatera in lower montane to submontane forests. This species has been classified as endangered by the IUCN, necessitating collection of more information on its habitat and environmental preferences before embarking on conservation planning. Studies on the species' natural habitat are scarce. This study aims to identify the environmental factors (edaphic + climatic factors) influencing the distribution of C. tungurrut along an altitudinal gradient in the Cibodas Biosphere Reserve. The nested plot method was applied to assess the forest vegetation at an altitudinal range of ca. 750-1800 m asl. Environmental factors were measured using portable equipment and through laboratory analysis. Ordination technique using canonical correspondence analysis (CCA) was utilised to pinpoint the environmental factors influencing the species' distribution based on basal area. CCA showed temperature to be the most limiting factor affecting the distribution. Edaphic factors-cation exchange capacity, content of carbon, nitrogen, phosphorus, and potassium, and soil pH-had less influence. Thus, it can be inferred that dependence of C. tungurrut on temperature determines its distribution pattern in its natural habitat, like Ostodes paniculata and Sloanea sigun. In contrast, the distribution of Castanopsis javanica, Castanopsis argentea, Schima wallichii, Altingia excelsa, Dacrycarpus imbricatus, Cestrum aurantiacum, and Castanopsis acuminatissima was found to be more influenced by edaphic factors than by climatic factors.

Full text

Edaphic and climatic factors affecting the
distribution of Castanopsis tungurrut (Blume) A.
DC. (Fagaceae) in cibodas biosphere reserve,
Indonesia
Dian Ridwan Nurdiana, Inocencio E. Buot, Jr.

THE EGYPTIAN JOURNAL OF BOTANY (EJBO)
The Egypan Journal of Botany (EJBO) is published by the Egypan Botanical Society. Egypt. J. Bot., Vol. 64, Special Issue, pp. 234-246 (2024)
Edaphic and climatic factors affecting the distribution of Castanopsis tungurrut
(Blume) A. DC. (Fagaceae) in cibodas biosphere reserve, Indonesia
Dian Ridwan Nurdiana1,2, Inocencio E. Buot, Jr.2
1Research Center for Ecology and Ethnobiology, Naonal Research and Innovaon Agency (BRIN), Indonesia
2Instute of Biological Sciences, College of Arts and Sciences, University of the Philippines Los Baños, Laguna 4031Philippines
Castanopsis tungurrut (Blume) A. DC. (Fagaceae) is an Indonesian nave plant species with limited
spaal distribuon; it is found only in Java, Kalimantan, and Sumatera in lower montane to
submontane forests. This species has been classified as endangered by the IUCN, necessitang
collecon of more informaon on its habitat and environmental preferences before embarking on
conservaon planning. Studies on the species’ natural habitat are scarce. This study aims to idenfy
the environmental factors (edaphic + climac factors) influencing the distribuon of C. tungurrut
along an altudinal gradient in the Cibodas Biosphere Reserve. The nested plot method was applied
to assess the forest vegetaon at an altudinal range of ca. 750–1800 m asl. Environmental factors
were measured using portable equipment and through laboratory analysis. Ordinaon technique
using canonical correspondence analysis (CCA) was ulised to pinpoint the environmental factors
influencing the species’ distribuon based on basal area. CCA showed temperature to be the most
liming factor affecng the distribuon. Edaphic factors – caon exchange capacity, content of
carbon, nitrogen, phosphorus, and potassium, and soil pH – had less influence. Thus, it can be
inferred that dependence of C. tungurrut on temperature determines its distribuon paern in its
natural habitat, like Ostodes panic ulata and Sloanea sigun. In contrast, the distribuon of
Castanopsis javanica, Castanopsis argentea, Schima wallichii, Alngia excelsa, Dacrycarpus
imbricatus, Cestrum auranacum, and Castanopsis acuminassima was found to be more
influenced by edaphic factors than by climac factors.
Keywords: Castanopsis tungurrut, species composion, altudinal gradient, environmental factors
ARTICLE HISTORY
Submied: Febr uary 20, 2024
Accepted: September 10, 2024
CORRESPONDANCE TO
Dian Ridwan Nurdiana,
Research Center for Ecology and Ethnobiology,
Naonal Research a nd Innovaon Agency
(BRIN), Indone sia
Email: drnurdia na@up.edu.ph
DOI: 10.21608/ej bo.2024.271501.2717
EDITOR
Prof. Monier Abd El-Ghani,
Department of Botany and Microbio logy,
Faculty of Scienc e, Cairo Universi ty, Egypt
Email: moniermoha medabdelghani @gmail.com
©2024 Egypan Botanical Society
INTRODUCTION
The distribuon of plant species is intricately linked to
a multude of factors, including predaon,
compeon, human acvity, and climate, which
shape their survival, growth, and reproducve
success. Specifically, temperature, altude, and water
availability are climac factors that limit species
distribuon (Box, 1995; Buot and Osumi, 2011;
Marnez and Buot, 2018; Villanueva and Buot, 2018;
Caringal et al., 2021). Understanding these factors
and their impact on specific plant species is crucial for
predicng and managing the ecological dynamics of
diverse ecosystems.
Castanopsis tungurrut, commonly known as
‘tungurut’, ‘kalimorot’, ‘tunggeureuk’, or ‘tunggurut’
in Indonesia, is an endangered plant species known
for its ecological significance and unique adaptaons
to specific environmental condions. Its distribuon
is directly affected by environmental condions such
as temperature, humidity, organic carbon (C-organic)
level, total nitrogen (N total), caon exchange
capacity (CEC), and pH (Zuhri and Mutaqien, 2011;
Nurcahyani, 2017). This study examines the
environmental factors (edaphic + climac factors)
influencing its distribuon in the following four
different locaons of the Cibodas Biosphere Reserve:
Cibodas, Cisarua, Selabintana, and Bodogol. This can
help gain insights into broader ecological processes,
potenally improving conservaon efforts for C.
tungurrut and other related plant species in the park.
Harapan et al. (2022) claimed that the most important
variables for predicng the distribuon of C.
tungurrut are elevaon, temperature seasonality, and
precipitaon in the warmest quarter. This species’
distribuon has been limited to an altudinal range of
1400–1800 m asl in Sumatera and 1000–1800 m asl in
Java, and it is mostly absent at lower elevaons due
to human acvies, including agriculture and
selement (Simbolon, 2001; Harapan et al., 2022).
Elevaon has been regarded as the main liming
factor for the distribuon of C. tungurrut. A previous
study on environmental variables influencing species
distribuon in the Philippines revealed that species
diversity decreases at higher altudes and that
vegetaon at lower elevaons is much larger and
taller than at higher elevaons (Buot and Okitsu,
1998). The decrease in species diversity is correlated
with insufficient heat at higher altudes, contribung
to the organic producon that supports tall trees, a
common paern in tropical forests. Notably, the
restricted distribuon range of C. tungurrut might be
correlated with nutrient availability.
Ecologically, C. tungurrut has a high frequency with an
importance value index (IVI) of 10.14, indicang that
the species can adapt to and has a wider tolerance to
environmental condions than other species (Arrijani,

Factors affecng Castanopsis tungurrut' distribuon …
2008). Nevertheless, low regeneraon rates can
threaten the existence of this species in its natural
habitat, as the survival rate was found to be only
33.33% (Handayani et al., 2019).
The low regeneraon rate of the species is closely
related to its low adaptaon to soil nutrients. Several
studies have highlighted the importance of root
morphology, anatomy, and architecture in adapng to
low soil oxygen levels, stress on increase plant
resilience, and the role of soil pH in nutrient
imbalances and constraints to plant growth on acid
soils (Adams, 1981; Puijalon et al., 2008; Pedersen et
al., 2020). Therefore, ability to adapt to the
environment is a significant factor in a species’
existence.
It can be inferred that climac and edaphic factors are
inter-complementary variables influencing the
distribuon of C. tungurrut. Several studies have
revealed the complexity of environmental factors
affecng the distribuon of Castanopsis species.
Cheuck and Fischer (2021) found that climate change
is likely to cause range reducons for some
Castanopsis species, parcularly in marginal tropical
zones, due to fragmented forest cover and a lack of
efficient seed dispersal mechanisms. Wang et al.
(2014) and Yamada and Miyaura (2005) posited that
the topography and nut size of Castanopsis species
contribute to the dominance and spaal genec
structure of the species. Addionally, Wibowo (2006)
suggested that Castanopsis argentea prefers a habitat
with stony soils containing low phosphorus (P)
concentraons.
This study aims to idenfy the environmental factors
affecng the distribuon paern of C. tungurrut
across various altudinal gradients and to elucidate
how specific soil characteriscs and climac variables
contribute to the spaal distribuon and abundance
of the species within the reserve. By understanding
these factors, valuable insights into the ecological
requirements and habitat preferences of the species
will be gained, facilitang informed conservaon and
management strategies for its preservaon.
MATERIALS AND METHODS
Study area
The study site is in Gede Pangrango Naonal Park,
Cibodas Biosphere Reserve, West Java, Indonesia. The
area encompasses three different regencies: Bogor,
Cianjur, and Sukabumi. Cibodas Biosphere Reserve
has a total area of 167,000 hectares and is the highest
mountain complex on Java Island. The study site is
distributed into four different locaons: Cibodas,
Bodogol, Selabintana, and Cisarua. In this study, the
precipitaon rate at a locaon refers to the monthly
precipitaon average during 2010–2021 (Figure 3).
The data showed the lowest precipitaon rate in
August and the highest in February. The precipitaon
rate is closely related to the vegetave and generave
cycles of a tree’s growth. The elevaon gradient at
each locaon varied, ranging from 750 to 1800 m asl
(Bodogol: 750–1100 m asl, Cibodas: 1300–1800 m asl,
Selabintana: 1000–1800 m asl, and Cisarua: 900–1500
m asl), represenng the lower and upper montane
forest of Gede Pangrango Naonal Park. The locaons
are bordered by agricultural land and human
selements at lower elevaons (ca. less than 1000 m
asl) and protected forests at higher elevaons. The
main vegetaon in this forest is Fago-Lauraceous,
referred to by Junghuhn and Miquel as a forest at
1000–2400 m asl in Gede Pangrango Naonal Park,
with common species being Acer laurinum,
Engelhardia spicata, Schima wallichii, Weinmannia
blumei, and the fern Cyathea (Simbolon et al., 2012).
Forest structure analysis
The nested plot method was used for forest structure
analysis. Plots of 20 × 20 m were used for trees, 10 x
10 m for poles, 5 x 5 m for saplings, and 2 x 2 m for
wildings inside each. The plots were ulised with an
elevaon gradient of around 750–1800 m asl, and the
coordinates were recorded using a GPS device
(Garmin eTrex 10). The total number of plots was 41,
represenng four locaons: Bodogol, Cibodas,
Cisarua, and Selabintana. Each locaon had a
different number of plots depending on the
topography, elevaon, and locaon: 10 for Bodogol,
14 for Cibodas, eight for Cisarua, and nine for
Selabintana.
Trees were defined as woody plants with stem
diameters greater than 5 cm and heights greater than
2 metres; poles were those whose stands had
diameters between 2.5 cm and 5 cm; saplings were
the individuals with a height of more than 130 cm and
a diameter at breast height (dbh) of less than 2.5 cm;
wildings were the individuals with heights less than
130 cm (Newton, 1988; Vargas-Rodriguez et al., 2005;
Fathia et al., 2019). All the data recorded were
analysed for abundance, density, frequency, index
diversity, and index evenness.
Basal Area and Relave Basal Area (Hanson and
Churchill, 1961)
Basal area (sq. cm) =  × ()
 and

Nurdiana and Buot, 2024
Egypt. J. Bot. Vol. 64, Special Issue, (2024)
236
Relave basal area (%) =     
      × 100,
where π = 3.14.
Dominance Index (C) (Simpson, 1949)
To assess the dominance within the community, the
dominance index was calculated as follows:
=(/),
where ni is the IVI for each species, and N represents
the total IVI of all species.
Margalef’s Richness Index (R1) (Margalef, 1958)
Margalef’s Richness Index was calculated as
,

()
=
1
R
where S represents the total number of species and n
represents the total number of individuals observed
in the community.
Shannon’s Index or α Diversity (Hʹ) (Shannon and
Weaver, 1963)
The diversity index was determined using the
Shannon–Wiener diversity index method:
(H󰆒)= H󰆒= (plogp)

 or H󰆒=
(ni )
log(ni n)


 ,
where pi represents the proporon of the total
sample belonging to a specific species, piʹs represents
the populaon parameters, log2 is equivalent to 3.322
log10, ni refers to the number of individuals of species
i in the sample, and n represents the total number of
all species.
Evenness Index (E) (Hill, 1973)
The evenness index was calculated as
E = /(/)
󰆓  or E =  
 – ,
where Hʹ refers to Shannon’s index, λ refers to
Simpson’s index, and N1 and N2 represent Hill’s
diversity numbers.
Ecological parameters
The following ecological parameters were measured:
topographic, microclimac, and edaphic factors.
Topographic factors included altude and slope, while
microclimac factors included rainfall, temperature,
and relave humidity. Edaphic factors included the
following chemical properes of soil: CEC; P,
Potassium (K), Calcium (Ca), and Magnesium (Mg)
concentraons; and N total. The slope was measured
using a clinometer.
Microclimac parameters
Temperature, wind speed, light intensity, and
humidity were measured using a portable gauge
(Lutron LM-8010) and a data logger (Benetech GM
1365) at each plot. Rainfall data for the Cibodas
Biosphere Reserve area from 2010 to 2021 (Figure 1)
were obtained from secondary sources using the
website hps://power.larc.nasa.gov/.
Edaphic parameters
To assess edaphic parameters, 24 soil samples were
collected from the four locaons. A modified version
of the soil sampling method proposed by Zhang et al.
(2021) was used. Topsoil samples were collected using
a cylinder core to extract the topsoil layer at a depth
of 0–15 cm or horizons O and A. The collected
samples were weighed, recorded, and analysed. The
analysis covered soil nutrients, including Ca, Mg, K, %
C, % N, and % P, and CEC, following the protocol
suggested by Huluka and Miller (2014). The analysis
was conducted at the Pusat Penelian Tanah dan
Agriklimat (Research Centre for Soil Research and
Agroclimate) and the Indonesian Center for
Biodiversity and Biotechnology (ICBB) Laboratory, PT
Biodiversitas Bioteknologi Indonesia, Bogor.
Addionally, soil pH was measured directly in the field
using a pH meter and soil tester on soil samples
collected from each locaon.
Canonical correspondence analysis
Analysis of the impact of environmental factors,
which comprised edaphic and microclimac factors,
on the distribuon of types was carried out with
ordinaon analysis techniques. This technique is
widely used in the study of modern ecology (Li et al.,
2017). Ordinaon techniques provide an objecve
representaon to determine the relaonship
between environmental factors and type distribuon.
In the ordinaon diagram, the nature of the
relaonships is shown by vectors, with lengths being
proporonal to their importance and direcons
indicang their correlaons with each axis. Canonical
correspondence analysis (CCA) was conducted using
PAST version 4. The basal area and environmental
data were transformed using log (x+1) to consider
zero values and prevent high values from influencing
the ordinaon (Aissat, 2023). The basal area of the
tree stage was used for CCA across all locaons.

Factors affecng Castanopsis tungurrut' distribuon …
Figure 1. Study area in Cibodas Biosphere Reserve, West Java, Indonesia.
RESULTS
Forest structure
The vegetaon analysis of florisc composion of the
study site resulted in 155 species (Table 1). Based on
basal area, the families Fagaceae, Theaceae,
Euphorbiaceae, Elaeocarpaceae, Proteaceae,
Araliaceae, Rhamnaceae, Anacardiaceae,
Podocarpaceae, and Alngiaceae were found to be
the most common in the study area, and S. wallichii
was found to have the highest basal area. The results
of the vegetaon paern analysis across the four
different locaons with an altudinal range of ca.
750–1800 m asl are presented in Table 2. The tree
stage had the highest number of species, while the
pole stage had the lowest. The diversity index was
relavely high for all stages.
Table 2 demonstrates that the forest in the Cibodas
Biosphere Reserve, located at an altudinal range of
ca. 750–1800 m asl, has high diversity. However,
based on the evenness index value (less than 1), it can
be concluded that the distribuon of species in the
area is uneven. The vegetaon structure of the lower
stages was found to differ in composion from the
higher stages (wilding to tree stages). These
differences could be aributed to compeon and
predaon factors in the study area. Regarding the
dominant species composion in the forest,
especially for the pole and sapling stages, invasive
species, namely Cestrum auranacum and Chinchona
pubescens, tended to dominate the area. This is
inseparable from the existence of natural forests with
quinine plantaons and the Cibodas Botanical
Garden, as examined by Zuhri and Mutaqien (2013)
regarding the possibility for plant collecons to
escape from botanical gardens and become exoc in
natural forest areas, such as Cestrum auranacum,
Calliandra calothyrsus, and Cinchona pubescens.
Consequently, in the long term, this can affect the
survival of endemic species and change the natural
vegetaon structure of the Gunung Gede Pangrango
forest.
All zones covered in the study represent the
distribuon range of Castanopsis. Most Castanopsis
species were found across all altudes, except C.
tungurrut, Castanopsis acuminassima, and C.
argentea, which were distributed at elevaons.

Nurdiana and Buot, 2024
Egypt. J. Bot. Vol. 64, Special Issue, (2024)
238
Table 1. The florisc composion of the four sampling locaons. BA is the basal area derived from the diameter at breast height. Dominant
species are indicated by an asterisk (*).
Cibodas
Bodogol
Selabintana
Cisarua
Number of plots
14
10
9
8
Number of spe cies
72
83
54
47
Plot size
Species
Family
Acer laurinum
Sapindaceae
0.214
0.005
Acronychia pedunculata
Rutaceae
0.467
0.056
0.352
Acronychia trifoliolata
Rutaceaea
0.013
0.006
Agathis borneensis
Araucariaceae
0.004
Aglaia sp.
Meliaceae
0.467
Aglaia hiernii
Meliaceae
0.115
Alangium chinense
Cornaceae
0.019
Alangium rotundifolium
Cornaceae
0.017
0.381
Albizia sp.
Fabaceae
0.009
Altingia excelsa
Alngiaceae
4.473*
0.201
0.074
Antidesma tetrandrum
Phyllanthaceae
0.012
0.033
Archidendron clypearia
Leguminosae
0.183
Artocarpus elasticus
Moraceae
0.136
Bonnetia sp.
Bonneaceae
0.028
Brassaiopsis glomerulata
Araliaceae
0.041
Bridelia insulana
Phyllanthaceae
0.056
0.030
Brugmansia suaveolens
Solanaceae
0.003
Calliandra calothyrsus
Fabaceae
0.044
Caryota mitis
Arecaceae
0.079
Casearia coriacea
Salicaceae
0.118
0.120
Castanopsis argentea
Fagaceae
4.881*
0.030
0.275
Castanopsis javanica
Fagaceae
3.478*
0.276
0.376*
0.996*
Castanopsis tungurrut
Fagaceae
1.066
1.125*
0.368
2.709*
Castanopsis acuminatissima
Fagaceae
0.707*
Celtis sinensis
Cannabaceae
0.006
Cestrum aurantiacum
Solanaceae
0.185
Cinchona pubescens
Rubiaceae
0.026
0.129
Cinnamomum rhynchophyllum
Lauraceae
0.093
Claoxylon longifolium
Euphorbiaceae
0.005
Coffea sp.
Rubiaceae
0.006
Coprosma sp.
Rubiaceae
0.002
Croton argyratus
Euphorbiaceae
0.010
Cryptocarya ferrea
Lauraceae
0.011
0.017
Cryptocarya laevigata
Lauraceae
0.096
Dacrycarpus imbricatus
Podocarpaceae
4.232*
0.007
Daphne composita
Thymelaeaceae
0.045
Decaspermum fruticosum
Myrtaceae
0.019
Dendrocnide stimulans
Urcaceae
0.008
0.010
Dipterocarpus hasseltii
Dipterocarpaceae
0.140
Dysoxylum alliaceum
Meliaceae
0.036
0.121
Dysoxylum nutans
Meliaceae
0.062
Dysoxylum parasiticum
Meliaceae
0.152
Dysoxylum excelsum
Meliaceae
0.068
0.003
Ehretia javanica
Boraginaceae
0.006
Elaeagnus latifolia
Elaeagnaceae
0.004
Elaeocarpus acronodia
Elaeocarpaceae
0.142
0.070
0.158
Elaeocarpus petiolatus
Elaeocarpaceae
0.400
Elaeocarpus sp.
Elaeocarpaceae
0.132
0.188*
0.076
Elaeocarpus submonoceras
Elaeocarpaceae
0.047
0.046
Endiandra rubescens
Lauraceae
0.017
Engelhardtia spicata
Juglandaceae
0.513
0.006
Eriosolena composita
Thymelaeaceae
0.022
0.008
Euonymus indicus
Celastraceae
0.014
Eurya acuminata
Pentaphylacaceae
0.034
0.037
0.014
Fagraea blumei
Genanaceae
0.030
Ficus alba
Moraceae
0.007
0.009
0.019
Ficus heterophylla
Moraceae
0.086
0.053
0.006
Ficus ribes
Moraceae
0.084
0.002
0.017
0.041
Ficus sp.
Moraceae
0.056
0.003
Ficus variegata
Moraceae
0.016

Factors affecng Castanopsis tungurrut' distribuon …
Ficus cuspidata
Moraceae
0.002
0.011
Ficus fistulosa
Moraceae
0.012
0.079
Ficus lepicarpa
Moraceae
0.066
Ficus sinuata
Moraceae
0.004
Flacourtia rukam
Salicaceae
0.003
Garcinia forbesii
Clusiaceae
0.184
Glochidion cyrtostylum
Phyllanthaceae
0.196
0.087
Glochidion rubrum
Phyllanthaceae
0.003
0.008
0.030
Gordonia excelsa
Theaceae
0.029
0.012
Gynotroches axillaris
Rhizophoraceae
Helicia robusta
Proteaceae
0.007
Helicia serrata
Proteaceae
0.016
0.002
0.620*
Itea macrophylla
Iteaceae
0.038
0.015
Itea sp.
Iteaceae
0.022
Knema sp.
Myryscaceae
0.065
Laportea sp.
Urcaceae
0.031
Lasianthus stercorarius
Rubiaceae
0.006
Lindera polyantha
Lauraceae
0.310
Lithocarpus elegans
Fagaceae
0.075
0.003
0.011
Lithocarpus indutus
Fagaceae
0.028
0.148
0.016
Lithocarpus pseudomoluccus
Fagaceae
0.144
0.783*
0.004
Litsea resinosa
Lauraceae
0.004
Litsea sp.
Lauraceae
0.019
Litsea garciae
Lauraceae
0.169
Litsea mappacea
Lauraceae
0.043
Macaranga rhizinoides
Euphorbiaceae
0.010
0.027
0.215
Macaranga triloba
Euphorbiaceae
0.040
0.004
Macropanax concinnus
Araliaceae
0.781
0.278
Macropanax dispermus
Araliaceae
0.530
0.397*
0.115
Maesopsis eminii
Rhamnaceae
0.931*
Magnolia liliifera
Magnoliaceae
0.009
Magnolia sumatrana
Magnoliaceae
0.367
Magnolia montana
Magnoliaceae
0.009
0.027
0.024
Mallotus sp.
Euphorbiaceae
0.044
Mangifera indica
Anacardiaceae
0.016
Manglietia glauca
Magnoliaceae
0.261
0.275
Mastixia trichotoma
Cornaceae
0.004
Melastoma
Melastomataceae
0.003
Myristica sp.
Myriscaceae
0.055
Myrtaceae
Myrtaceae
0.006
Neolitsea cassiifolia
Lauraceae
0.197
Neolitsea javanica
Lauraceae
0.137
0.044
0.309
Neolitsea triplinervia
Lauraceae
0.273
Neonauclea excelsa
Rubiaceae
0.002
Neonauclea lanceolata
Rubiaceae
0.119
0.055
0.057
Orophea hexandra
Annonaceae
0.021
Ostodes paniculata
Euphorbiaceae
0.258
0.407*
Pavetta montana
Rubiaceae
0.005
Persea excelsa
Lauraceae
0.158
Persea rimosa
Lauraceae
0.655
0.095
0.060
Phoebe excelsa
Lauraceae
0.424
0.206
Phoebe grandis
Lauraceae
0.083
0.007
0.016
Pinus merkusii
Pinaceae
0.025
Piper sp.
Piperaceae
0.005
Podocarpus neriifolius
Podocarpaceae
0.028
0.091
Polyalthia subcordata
Annonaceae
0.003
0.001
Polyathia sp.
Annonaceae
0.021
Polyosma integrifolia
Escalloniaceae
0.055
Prunus arborea
Rosaceae
0.183
0.409
0.251
Pternandra azurea
Melastomataceae
0.031
Rapanea hasseltii
Primulaceae
0.018
Rauvolfia javanica
Apocynaceae
0.005
Rauvolfia sp.
Apocynaceae
0.089
Saurauia distatosa
Acnidiaceae
0.018
Saurauia nudiflora
Acnidiaceae
0.005
Saurauia pendula
Acnidiaceae
0.178
Saurauia bracteosa
Acnidiaceae
0.029
Schefflera aromatica
Araliaceae
0.149

Nurdiana and Buot, 2024
Egypt. J. Bot. Vol. 64, Special Issue, (2024)
240
Schefflera sp.
Araliaceae
0.049
Schima wallichii
Theacaea
6.131*
1.578*
2.554*
0.450*
Sloanea sigun
Elaeocarpaceae
0.089
0.035
Spondias pinnata
Anacardiaceae
0.529*
Stemonurus secundiflorus
Stemonuraceae
0.002
Sterculia sp.
Malvaceae
0.005
Sterculia sp.
Malvaceae
0.184
Sundacarpus amarus
Podocarpaceae
0.089
Swietenia macrophylla
Meliaceae
0.113
Symplocos cochinchinensis
Symplocaceae
0.011
0.019
0.004
Symplocos costata
Symplocaceae
0.057
0.011
Symplocos fasciculata
Symplocaceae
0.004
0.005
Syzygium antisepticum
Myrtaceae
0.144
Syzygium densiflorum
Myrtaceae
0.021
Syzygium nervosum
Myrtaceae
0.099
Syzygium pycnanthum
Myrtaceae
0.071
0.020
Syzygium rostratum
Myrtaceae
0.783
0.500
0.143
Syzygium sp.
Myrtaceae
0.047
Syzygium racemosum
Myrtaceae
0.004
0.035
0.049
Turpinia sphaerocarpa
Staphyleaceae
0.356
0.101
0.078
Turpinia montana
Staphyleaceae
0.004
0.005
Vernonia arborea
Asteraceae
0.074
0.155
Viburnum sambucinum
Adoxaceae
0.032
0.016
Villebrunea scabra
Urcaceae
0.427
0.056
0.296
0.296
Wendlandia densiflora
Rubiaceae
0.004
Wenlandia sp.
Rubiaceae
0.146
Wenmannia blumei
Cunoniaceae
0.025
Plot size = 20 × 20 m.
Notably, Castanopsis diversity decreased with
increasing altude. This is a common phenomenon in
which the number of species, species composion,
structure, physiognomy, and tree architecture change
as the elevaon increases (Simbolon et al., 2012).
Factors influencing the distribuon of Castanopsis
The ordinaon technique was used to assess the
relaonship between dominant trees and
environmental factors (edaphic and climac) (Figure
2). Environmental factors consisted of three climac
factors and six edaphic factors. C. tungurrut was found
to be associated with moderate temperatures and
lower altudes; its distribuon was more driven by
increased temperature and decreased altude than
by edaphic factors. Other Castanopsis species
recorded in this study, including Castanopsis javanica,
C. argentea, and C. acuminassima, showed
variaons in their relaonship with environmental
factors. C. argentea and C. acuminassima were
observed to be closely influenced by pH, while C.
javanica was found to be more influenced by edaphic
factors (N, K, P, and Ca concentraons, CEC, and pH)
and altude. Like C. javanica, S. wallichii, and
Dacrycarpus imbricatus were likely more influenced
by edaphic factors than by climac condions. The
remaining species, including Ostodes paniculate,
Sloanea sigun, Cestrum auranacum, and Alngia
excelsa, were influenced by temperature and pH.
Upon invesgang the soil properes and
microclimac condions, we observed variaons in
environmental factors at different elevaons (Table
4).
DISCUSSION
Environmental factors influencing C. tungurrut and
associated vegetaon at Cibodas Biosphere Reserve
Table 1 presents variaons observed in species
composion per locaon, dominant species, and
basal area values of C. tungurrut at the Cibodas
Biosphere Reserve. The distribuon of C. tungurrut
and its correlaon with environmental factors
(edaphic and climac factors) is illustrated in Figure 2.
The edaphic and climac factors covered in this study
varied with altudinal gradient (Table 4). Temperature
tended to decrease with increasing altude, while the
edaphic factors, consisng of CEC and Mg, Ca, K, P, N,
and C concentraons, varied at different altudes.
This variaon could be aributed to the slope
variaon and decomposion rates per altude.
Addionally, the precipitaon rate at the study site
showed dry and wet months, with relavely high
precipitaon from December to March and lower
precipitaon in April (Figure 3).
C. tungurrut and nine dominant species were tested
in relaon to environmental variables using CCA
(Figure 2). A total of nine environmental variables

Factors affecng Castanopsis tungurrut' distribuon …
Table 2. General characteriscs of vegetaon structure based on the density of species at various life stages at Cibodas Biosphere Reserve.
Scienfic name
Life stage
Density
Total Number of Species
Diversity Index
Evenness Index
Schima wallichii Choisy
Tree
56.09
154
4.3
0.48
Villebr unea scabra (Blume) Wedd.
Tree
43.29
Macropanax dispermus (Blume) Kunze
Tree
32.3
Syzygium rostratum (Blume) DC.
Tree
31.09
Castanopsis tungurrut (Blume) A. DC
Tree
23.17
Macropanax concinnus Miq.
Tree
21.34
Turpinia sphaerocarpa Hassk.
Tree
19.51
Castanopsis javanica (Blume) A. DC
Tree
18.29
Elaeocarpus sp
Tree
16.46
Acronychia pedunculata (L.) Miq.
Tree
15.85
Cestrum aurantiacum Lindl.
Pole
39
105
4.2
0.65
Lasianthus stercorarius Blume
Pole
36.5
Polyalthia subcordata (Blume) Blume
Pole
36.5
Turpinia sphaerocarpa Hassk.
Pole
31.7
Syzygium rostratum (Blume) DC.
Pole
29.26
Villebr unea scabra (Blume) Wedd.
Pole
26.8
Cinchona pubescens Va h l
Pole
24.39
Schima wallichii Choisy
Pole
24.39
Macropanax undulatus ( Wall.ex G. Donn) Seem
Pole
21.95
Magnolia liliifera (L.) Baill.
Pole
21.95
Lasianthus stercorarius Blume
Sapling
390
135
4.3
0.5
Cestrum aurantiacum Lindl.
Sapling
370
Magnolia liliifera (L.) Baill.
Sapling
260
Freycinetia insignis Blume
Sapling
230
Polyalthia subcordata (Blume) Blume
Sapling
220
Bartlettina sordida (Less.) R.M. King & H.Rob.
Sapling
200
Psychotria montana Blume
Sapling
170
Cinchona pubescens Va h l
Sapling
150
Symplocos cochinchinensis (Lour.) S Moore
Sapling
150
Acronychia pedunculata (L.) Miq.
Sapling
140
Lasianthus laevigatus Blume
Wilding
3719.5
111
3.9
0.5
Elatostema strigosum Hassk.
Wilding
2560.9
Psychotria montana Blume
Wilding
1707.3
Cyrtandra picta Blum e
Wilding
1158.5
Trevesia sundaica Miq.
Wilding
1158.5
Strobilanthes cernua Blume
Wilding
914.6
Cestrum aurantiacum Lindl.
Wilding
853.6
Piper baccatum C.DC.
Wilding
792.6
Dichroa febrifuga L o u r.
Wilding
731.7
Ficus sp.
Wilding
731.7
Table 3. Eigenvalues for axes of the ordinaon diagram.
Axis
1
2
3
4
5
6
7
8
9
Eigen value
0.5442
0.3127
0.1804
0.08146
0.05784
0.02982
0.02275
0.004598
1.28E-07
Table 4. Canonical coefficient and intra-set correlaon of experimental variables with two axes of canonical correspondence analysis (CCA).
Axis variable
Canonical coefficient
Correlaon coefficient
Axis 1
Axis 2
Axis 1
Axis 2
Slope
-0.16
-0.47
0.12
-0.54
CEC
-0.63
0.46
-0.68
0.42
C (%)
-0.5
0.47
-0.57
0.52
N (%)
-0.4
0.45
-0.56
0.52
K (cmol/kg)
-0.2
0.66
-0.43
0.3
Ca (cmol/kg)
-0.08
0.51
-0.36
0.36
Alt
-0.3
0.63
-0.31
0.71
Tem p
0.5
-0.59
0.43
-0.54
pH
0.14
0.32
0.14
0.51

Nurdiana and Buot, 2024
Egypt. J. Bot. Vol. 64, Special Issue, (2024)
242
were analysed relave to 10 dominant tree species,
resulng in nine axes, with a total variance of 44.1%
and 25.5% for axis 1 and axis 2, respecvely. The
eigenvalues for axes 1 and 2 were 0.5542 and 0.3127,
respecvely. Based on the orthogonal projecons, C.
tungurrut is more likely to be found in areas with low
pH, low Ca, low K, low altude, low N, low C, low CEC,
high temperature, and high slope. The species was
found to have the strongest posive correlaon with
temperature (0.54) and a strong negave correlaon
with CEC (-0.49).
Based on the CCA ordinaon diagram (Figure 2),
temperature can be defined as an important factor
influencing the distribuon of C. tungurrut. According
to Körner et al. (2016), plants have well-defined
threshold responses to temperature, which reveal
unique abnormalies in cell funcon within a
constrained temperature range, affecng the plant’s
survival, growth, and ability to regenerate. The result
that temperature is the dominant climac driver of C.
tungurrut distribuon conforms to the findings of
previous studies by Harapan et al. (2022), Fathia et al.
(2019), Kusmana and Suwandhi (2019), Santhyami et
al. (2021), and Wibowo (2006). They revealed that
temperature and altude were the dominant factors
influencing the distribuon of C. tungurrut in
Sumatera, while in West Java, including Mount
Galunggung, Pakenjeng Garut and Mount Gede
Pangrango, the altude in the submontane zone
(1000–1400 m asl) and slopes > 40% were observed
to be the liming environmental factors for the
distribuon of C. tungurrut. However, this finding is
discordant with Paramita et al. (2022) and Sosilawaty
et al. (2022), who found that C. tungurrut can also be
found in peat swamp forests in Kalimantan, which had
not been recorded before. Usually, C. tungurrut is
found in lower to submontane forests. It is indicated
that C. tungurrut can be tolerant of higher
temperatures but is intolerant of lower temperatures,
which are common at higher altudes. In this regard,
Schindlbacher et al. (2010) claimed that, compared to
lower elevaon locaons, soil organic maer
decomposion at higher elevaon forests is more
suscepble to climate change because it will be
impacted in a more sensive (cooler) temperature
range.
On the other hand, a few dominant species found at
the study site, including C. javanica, C. argentea, S.
wallichii, A. excelsa, and D. imbricatus, were observed
to be driven by edaphic factors rather than climac
factors. In this study, C. javanica was found to be more
sensive to lower C, Ca, N, and K content, CEC, pH,
and altude. In addion, C. argentea and C.
acuminassima were observed to be influenced by
pH, while O. paniculate and S. sigun were found to be
closely influenced by moderate to high temperatures.
Furthermore, previous studies by Hilwan and Irfani
(2018), Putri (2021), and Wibowo (2006) on C.
argentea have revealed different results indicang
that the distribuon of C. argentea is influenced by
total N, altude, CEC, temperature, and P content. In
submontane forests and the surrounding forest
(Telaga Warna Nature Reserve) in Java Mountain
Forest, C. argentea dominated at 1100–1400 m asl
and gradually diminished in number at higher
elevaons and at regions with higher P content.
Wibowo (2006) found that C. argentea and other
dominant species (A. excelsa, Laportea smulans, C.
tungurrut, and C. javanica) are not dependent on soil
type, except S. wallichii.
Compared to other species found in this study, C.
tungurrut was observed to be driven largely by
temperature rather than edaphic factors, with a
correlaon of 0.54 (p > 0.05, not significant). Previous
studies by Toledo et al. (2012) and Zhang et al. (2016)
revealed that climac factors are stronger drivers of
species distribuon in subtropical karst forests than
edaphic factors. It is noteworthy that C. tungurrut can
opmise soil nutrients, which enables the species to
survive in nutrient-limited environments, exhibit
tolerance to temperature changes, and form
underground microbial associaons, which have not
been covered in this study.
Effect of environmental factors on C. tungurrut
regeneraon
The regeneraon of C. tungurrut is correlated with
environmental factors. It was found that edaphic and
climac factors contributed to the populaon and
distribuon of the species in its natural habitat (Table
2). We found that the tree stage of C. tungurrut was
significantly influenced by temperature (Figure 2). The
temperature gradient influenced the germinaon
capacity and restoraon of populaon in the natural
habitat, as described in previous studies in semi-
sunny slope habitats (Song et al., 2016; Zhao et al.,
2021). In addion, temperature could shi the
distribuon of Fagaceae, as described in the
distribuon paerns of Castanopsis echinocarpa and
Castanopsis calathiformis in tropical montane regions
of Southwest China and C. tungurrut in Mount Geulis,
West Java, in relaon to their tolerance to low
temperature and soil moisture (Du and Huang, 2008;
Song et al., 2021; Lukman et al., 2022). Temperature

Factors affecng Castanopsis tungurrut' distribuon …
Figure 2. Ordinaon diagram of canonical correspondence analysis (CCA) showing C. tungurrut and nine selected dominant tree species with nine
environmental variables (slope; CEC; C, N, K, and Ca concentraons; altude; temperature; and pH). CT – Castanopsis tungurrut, CJ – Castanopsis
javanica, CA – Castanopsis argentea, SW – Schima wallichii, AE – Alngia excelsa, DI – Dacrycarpus imbricatus, CAu – Cestrum auranacum, CAc
– Castanopsis acuminassima, OP – Ostodes paniculata, SSi – Sloanea sigun. The highest total variances of axis 1 and axis 2 are 44.1% and 25.5%,
respecvely. Total inera is 1.2327462.
Table 5. Average values of environmental factors across various altudinal gradients.
Altitude
Edaphic
Climatic
CEC
pH
P
N
C
Mg
Ca
K
Humidity
Slope (%)
Tem p . (°C)
700–800
26,2
4.50
4.7
0.42
6.33
0.02
0.12
0.09
99.85
57.50
21.84
800–900
26.20
4.50
4.70
0.42
6.33
0.04
0.12
0.04
99.81
25.00
20.92
900-–1000
31.44
5.40
3.15
0.67
11.36
0.03
0.12
0.05
99.97
35.50
20.40
1000–1100
40.72
5.00
3.00
0.84
14.10
0.02
0.06
0.03
99.92
44.50
19.10
1100–1200
28.62
6.23
16.83
0.83
13.80
2.32
14.13
1.07
95.23
17.33
19.33
1200–1300
39.41
5.73
15.07
1.28
20.25
1.51
5.43
0.18
99.54
45.00
18.09
1300–1400
42.86
5.93
45.41
0.95
16.83
0.39
187.09
35.55
99.87
24.40
18.34
1400–1500
28.65
5.98
13.99
0.93
14.39
0.51
8.96
8.45
99.67
18.86
17.59
1500–1600
41.29
6.00
19.74
1.55
26.75
0.84
133.78
25.39
99.92
19.00
16.99
1600–1700
62.88
5.20
41.80
2.20
42.23
0.23
175.38
84.83
99.94
25.20
16.12
1700–1800
42.20
6.15
56.91
1.29
25.00
0.19
83.31
40.25
99.84
27.50
15.70
S.D.
10.8
0.63
18.9
0.5
10.3
0.7
75.6
27
1.3
13
1.9
Figure 3. Monthly precipitaon average in Gede Pangrango Naonal Park, Cibodas Biosphere Reserve, Indonesia, during 2010–2021.

Nurdiana and Buot, 2024
Egypt. J. Bot. Vol. 64, Special Issue, (2024)
244
is considered as a predominant factor in the
distribuon of C. tungurrut because nutrient
availability is closely related to slope declivity and
temperature. Nutrient availability and temperature
tend to decrease with increasing altude,
contribung to soil decomposion.
CONCLUSION
The distribuon of C. tungurrut is driven largely by
temperature rather than edaphic condions
(macronutrients). The tree stage of C. tungurrut tends
to distribute at higher altudes (1500 m asl), with
temperatures around 16°C as the lowest temperature
limit, and can tolerate higher temperatures at lower
elevaons. Environmental factors significantly affect
the distribuon paern of C. tungurrut. The species is
well adapted to low soil nutrion in warm
temperatures. However, the selected dominant
species in the study area responded differently to
environmental factors. Most dominant species were
more significantly impacted by edaphic factors than
by climac factors.
ACKNOWLEDGMENTS
We wish to thank the SEARCA for the research grant
and Instute of Biological Sciences, CAS, University of
the Philippines Los Baños for the permit. We are very
grateful to Mr. Nudin, Mr. Ujang Rustandi, Mr.
Rustandi, Mr. Emus, and Mr. Cahyadi staff member
Cibodas Botanic Garden, Naonal Research and
Innovaon Agency (BRIN) for field work assistance.
We also acknowledge with gratude the support from
Gunung Gede Pangrango Naonal Park staff member
Mr. Ae, Mr. Dayat and Mr. Komar for their support.
This study was carried out under Gunung Gede
Pangrango Naonal Park permit reference no
SI37/BBTNGGP/Tek.2/06/2022.
REFERENCES
Adams F (1981) Nutritional imbalances and constraints to
plant growth on acid soils. Journal of Plant Nutrition 4:
81–87.
Aissat L (2023) Diversity and Ant Community Composition
in Some Algerian Islands. Ekologia Bratislava 42(1),
17–25. https://doi.org/10.2478/eko-2023-0003
Arrijani (2008) Vegetation structure and composition of the
montane zone of Mount Gede Pangrango National
Park. Biodiversitas Journal of Biological Diversity 9(2):
134–141. https://doi.org/10.13057/biodiv/d090212
Box EO (1995) Factors Determining Distributions of Tree
Species and Plant Functional Types. Vegetatio 121(1):
101–116.
Buo t I and Osumi K (2011) Land use type pattern and woody
species composition near human disturbed landscapes
on mount makiling, Luzon Island. American Journal of
Environmental Sciences 7(4): 306–315.
https://doi.org/10.3844/ajessp.2011.306.315
Buot I and Okitsu S (1998) Vertical distribution and
structure of the tree vegetation in the montane forest
of Mt. Pulog, Cordillera mountain range, the highest
mountain in Luzon Is., Philippines. Vegetation Science
15(1): 19–32.
Caringal AM, Buot I and Villanueva ELC (2021) Endemic
Philippine teak (Tectona philippinensis Benth. & Hook.
f.) and associated flora in the coastal landscapes of
Verde Island Passage, Luzon Island, Philippines.
Current Science 120(6): 1057–1065.
https://doi.org/10.18520/cs/v120/i6/1057-1065
Cheuk ML and Fischer GA (2021) The impact of climate
change on the distribution of Castanopsis (Fagaceae)
species in south China and Indo-China region. Global
Ecology and Conservation 26: 1-14.
https://doi.org/https://doi.org/10.1016/j.gecco.2020.
e01388
Du Y, and Huang Z (2008) Effects of seed mass and
emergence time on seedling performance in
Castanopsis chinensis. Forest Ecology and
Management 255: 2495–2501.
https://api.semanticscholar.org/CorpusID:84917181
Fathia AA, Hilwan I, and Kusmana C (2019) Species
Composition and Stand Structure in sub-montane
Forest of Mount Galunggung, Tasikmalaya, West Java.
IOP Conference Series: Earth and Environmental
Science 394(1):1-9. https://doi.org/10.1088/1755-
1315/394/1/012012
Handayani A, Lailaty IQ, and Astutik S (2019) Evaluasi
Kesintasan dan Pertumbuhan Beberapa Jenis Pohon
Lokal di Area Restorasi Cagar Biosfer Cibodas. Jurnal
Pengelolaan Sumberdaya Alam Dan Lingkungan
(Journal of Natural Resources and Environmental
Management 9(3): 541–548.
https://doi.org/10.29244/jpsl.9.3.541-548
Harapan TS, Nurainas, Syamsuardi, and Taufiq A (2022)
Identifying the potential geographic distribution for
Castanopsis argentea and C. tungurrut (Fagaceae) in
the Sumatra Conservation Area Network, Indonesia.
Biodiversitas 23(4): 1726–1733.
https://doi.org/10.13057/biodiv/d230402
Hilwan I, and Irfani E (2018) Pola Penyebaran Dan
Regenerasi Jenis Saninten (Castanopsis argentea
Blume) Di Resort Selabintana, Taman Nasional Gunung
Gede Pangrango." Distribution Pattern and
Regeneration of Saninten (Castanopsis argantea
Blume) in Selabintana Resort, Gunung Gede". Journal
of Tropical Silviculture 9(1): 53–59.
https://doi.org/10.29244/j-siltrop.9.1.53-59
Huluka G, and Miller R (2014) Particle size determination by
hydrometer method. Southern Cooperative Series
Bulletin 419: 180–184.
Körner C, Basler D, Hoch G, Kollas C, Lenz A, Randin CF,
Vitasse Y, and Zimmermann NE (2016) Where, why and
how? Explaining the low‐temperature range limits of

Factors affecng Castanopsis tungurrut' distribuon …
temperate tree species. Journal of Ecology 104(4):
1076–1088.
Kusmana C, and Suwandhi I (2019) Diversity of plant species
and the presence of Invasive Alien Species (IAS) in the
Sub-Montane Forest at Pakenjeng Region, Southern
Part of Garut, West Java. IOP Conference Series: Earth
and Environmental Science 399(1):,1-10.
Li W, Cui L, Sun B, Zhao X, Gao C, Zhang Y, Zhang M, Pan X,
Lei Y, and Ma W (2017) Distribution patterns of plant
communities and their associations with
environmental soil factors on the eastern shore of Lake
Taihu, China. Ecosystem Health and Sustainability 3(9):
1-11.
https://doi.org/10.1080/20964129.2017.1385004
Lukman AH, Parikesit P, Hadikusumah HY, and Rahmat A
(2022) Tree diversity and forest structure of tropical
forest in Mount Geulis, Cianjur. Journal of Global
Forest and Environmental Science 2(2) :10–19.
Martinez M, and Buot IE (2018) Altitudinal Distribution of
Some Hoya species in the Philippines. The Thailand
Natural History Museum Journal 12(1): 9–18.
Newton PN (1988) The structure and phenology of a moist
deciduous forest in the Central Indian Highlands.
Vegetatio 75(1–2): 3–16.
https://doi.org/10.1007/BF00044621
Nurcahyani R (2017) Pola Penyebaran dan Karakteristik
Tempat Tumbuh Castanopsis javanica dan Castanopsis
tungurrut di Hutan Gunung Galunggung Tasikmalaya.
Master thesis, IPB University, Indonesia.
Paramita S, Suwasono RA, Lasmito, Setyasih I, Ariyanto,
Mulyadi R, and Sulistioadi YB (2022) Vegetation
diversity of Hemaq Beniung Customary Forest, West
Kutai, East Kalimantan. Jurnal Penelitian Kehutanan
Wallacea 11(2): 111–123.
https://doi.org/10.18330/jwallacea.2022.vol11iss2pp
111-123
Pedersen O, Sauter M, Colmer TD, and Nakazono M (2020)
Regulation of root adaptive anatomical and
morphological traits during low soil oxygen. The New
Phytologist 229: 42-49.
Puijalon S, Piola F, and Bornette G (2008) Abiotic stresses
increase plant regeneration ability. Evolutionary
Ecology 22: 493–506.
Putri DM (2021) Autekologi Castanopsis argentea (Blume)
A.DC. di Kawasan Cagar Alam Telaga Warna,
Kabupaten Bogor. Master thesis, IPB university,
Indonesia.
Santhyami S (2021) Tree Community Structure and
Aboveground Carbon Stock of Sacred Forest in
Pasaman, West Sumatera. BIOTROPIA - The Southeast
Asian Journal of Tropical Biology 28(3): 253–262.
https://doi.org/10.11598/btb.2021.28.3.1416
Schindlbacher A, de Gonzalo C, Díaz-Pinés E, Gorría P,
Matthews B, Inclán R, Zechmeister-Boltenstern S,
Rubio A, and Jandl R (2010) Temperature sensitivity of
forest soil organic matter decomposition along two
elevation gradients. Journal of Geophysical Research
115: 1-10.
Simbolon H (2001) The growth dynamics on tree species of
Fagaceae family in a tropical montane rain forest of
West Java, Indonesia. Berita Biologi 5(6) : 659–666.
Simbolon H, Suzuki E, and Susanti R (2012) Some Vegetation
types studied with the same methodology in
Indonesia. In "Ecological Research Monographs",
Springer. Japan. p 71-92
Song X, Cao M, Li J, Kitching RL, Nakamura A, Laidlaw MJ,
Tang Y, Sun Z, Zhang W, and Yang J (2021) Different
environmental factors drive tree species diversity
along elevation gradients in three climatic zones in
Yunnan, southern China. Plant Diversity 43(6) : 433–
443.
https://doi.org/https://doi.org/10.1016/j.pld.2021.04
.006
Song X, Nakamura A, Sun Z, Tang Y, and Cao M (2016)
Elevational Distribution of Adult Trees and Seedlings in
a Tropical Montane Transect, Southwest China.
Mountain Research and Development 36(3) : 342–354.
https://doi.org/10.1659/MRD-JOURNAL-D-15-
00109.1
Sosilawaty, Jaya A, Rotinsulu JM, Hastari B, Hidayat N, and
Sianipar E (2022) Effect of Drainage Channels on
Vegetation Diversity of Tropical Peatswamp Forest of
Sebangau National Park, Indonesia. Journal of
Experimental Biology and Agricultural Sciences 10(1):
48–63. https://doi.org/10.18006/2022.10(1).48.63
Toledo M, Peña‐Claros M, Bongers F, Alarcón A, Balcázar J,
Chuviña J, Leaño C, Licona JC, and Poorter L (2012)
Distribution patterns of tropical woody species in
response to climatic and edaphic gradients. Journal of
Ecology 100(1): 253–263.
Vargas-Rodriguez YL, Vázquez-García JA, and Williamson G
B (2005) Environmental correlates of tree and
seedling-sapling distributions in a Mexican tropical dry
forest. Plant Ecology 180(1) : 117–134.
https://doi.org/10.1007/s11258-005-3026-9
Villanueva ELC, and Buot IE (2018) Vegetation analysis along
the altitudinal gradient of Mt. Ilong, Halcon range,
Mindoro island, Philippines. Biodiversitas 19(6): 2163–
2174. https://doi.org/10.13057/biodiv/d190624
Wang Z, Lian J, Ye W, Cao HL, and Wang Z (2014) The spatial
genetic pattern of Castanopsis chinensis in a large
forest plot with complex topography. Forest Ecology
and Management 318 : 318–325.
Wibowo C (2006) Hubungan Antara Keberadaan Saninten
dengan Beberapa Sifat Tanah: Kasus di Taman
Nasional Gunung Gede - Pangrango, Jawa Barat. Ph.D.
Dissertation, IPB University, Indonesia.
Yamada H, and Miyaura T (2005) Geographic variation in
nut size of Castanopsis species in Japan. Ecological
Research 20(1) : 3–9. https://doi.org/10.1007/s11284-
004-0006-9
Zhang, C., Li, X., Chen, L., Xie, G., Liu, C., & Pei, S. (2016).
Effects of topographical and edaphic factors on tree
community structure and diversity of subtropical
mountain forests in the Lower Lancang River Basin.
Forests 7(10). https://doi.org/10.3390/f7100222

Nurdiana and Buot, 2024
Egypt. J. Bot. Vol. 64, Special Issue, (2024)
246
Zhang J, Chen H, Fu Z, and Wang K (2021) Effects of
vegetation restoration on soil properties along an
elevation gradient in the karst region of southwest
China. Agriculture, Ecosystems and Environment,
320(3):1-13
https://doi.org/10.1016/j.agee.2021.107572
Zhao Z, Liu Y, Jia H, Sun W, Ming A, Pang S, An N, Zhang JH,
Tang C, and Dong S (2021) Influence of Slope Direction
on the Soil Seed Bank and Seedling Regeneration of
Castanopsis hystrix Seed Rain. Forests 12(500): 1-13.
https://api.semanticscholar.org/CorpusID:234830889
Zuhri M and Mutaqien Z (2013) The Spread of Non-native
Plant Species Collection of Cibodas Botanical Garden
into Mt. Gede Pangrango National Park. Journal of
Tropical Life Science 3(2) : 74–82.
https://doi.org/10.11594/jtls.03.02.01
Zuhri M, and Mutaqien Z (2011) Perubahan Komposisi
Vegetasi dan Struktur Pohon pada Plot Meijer( 1959-
2009 ) Di Gunung Gede [Change in vegetation
compostion and tree structure in Meijer plot (1959-
2009) on Mount Gede]. Buletin Kebun Raya 14(1).