REINWARDTIA
Vol. 22. No. 1. pp: 1‒25
DOI: 10.55981/reinwardtia.2023.4399
1
VARIATION IN THE COMPOSITION AND STRUCTURE OF NATURAL LOW-
LAND FORESTS AT BODOGOL, GUNUNG GEDE PANGRANGO NATIONAL
PARK, WEST JAVA, INDONESIA
Received September 1, 2022; accepted January 3, 2023
ASEP SADILI
Department of Biology, Faculty of Mathematics and Natural Sciences, Universitas Indonesia. Kampus UI Gedung E
Level 2, Jln. Lingkar Kampus Raya, Pondok Cina, Beji, Depok 16424, Indonesia.
Research Center for Ecology and Ethnobiology, National Research and Innovation Agency (BRIN), Jln. Raya Jakarta-
Bogor Km. 46, Cibinong, Bogor 16911, Indonesia.
Email: asep016@brin.go.id. https://orcid.org/0000-0002-9019-8094.
ANDI SALAMAH
Department of Biology, Faculty of Mathematics and Natural Sciences, Universitas Indonesia. Kampus UI Gedung E
Level 2, Jln. Lingkar Kampus Raya, Pondok Cina, Beji, Depok 16424, Indonesia.
Email: salamah@sci.ui.ac.id. https://orcid.org/0000-0002-4074-8342.
EDI MIRMANTO
Research Center for Ecology and Ethnobiology, National Research and Innovation Agency (BRIN), Jln. Raya Jakarta-
Bogor Km. 46, Cibinong, Bogor 16911, Indonesia.
Email: emirmanto@yahoo.com. https://orcid.org/0000-0001-7121-9980.
KUSWATA KARTAWINATA
Integrative Research Center, The Field Museum, 1400 Lake Share Drive, Chicago, IL, 60605, USA.
Email: kkartawinata@gmail.com. https://orcid.org/0000-0001-9656-8095.
ABSTRACT
SADILI, A., SALAMAH, A., MIRMANTO, E. & KARTAWINATA, K. 2023. Variation in the composition and
structure of natural lowland forests at Bodogol, Gunung Gede Pangrango National Park, West Java, Indonesia.
Reinwardtia 22(1): 1‒25. — An analysis of the composition and structure of lowland natural forests was carried out in
Bodogol, Gunung Gede Pangrango National Park (GGPNP). The two study plots (P1CS and P2CS) were located on
Cisuren and one plot (P3CP) on Cipadaranten hill. We recorded 107 species and 48 families with an average basal
area of 19.73 m
2
/ha, and an average density of 348 trees/ha. The species richness was poorer than those of the typical
lowland rainforests of Kalimantan and Sumatra but comparable to those of the montane forests of Java. The IUCN-
Red Listed species were Castanopsis argentea and Castanopsis tungurrut (critical) and Saurauia
bracteosa (vulnerable). Based on the two dominant species, the forests can be designated as the Maesopsis eminii-
Syzygium acuminatissimum association and Syzygium acuminatissimum-Lithocarpus korthalsii association. Maesopsis
eminii was dominant in P1CS (IV= 56.46%) and P3CP (IV=55.94%), while Syzygium acuminatissimum in P2CS (IV=
43.67%). Maesopsis eminii was a strongly aggressive and invasive species, that endangered the purity of the natural
forest GGPNP, therefore, it must be eradicated. Vertically, P2CS and P3CP consisted of four strata, while P1CS had
three strata. This one-hectare study can be considered as a minimal area to reflect the floristic representation of
lowland forest and submontane forest.
Key words: Association, forest structure, Gunung Gede Pangrango National Park, lowland forests, minimal area,
species composition, species richness.
ABSTRAK
SADILI, A., SALAMAH, A., MIRMANTO, E. & KARTAWINATA, K. 2023. Variasi komposisi dan struktur hutan
pamah alami di Bodogol, Taman Nasional Gunung Gede Pangrango, Jawa Barat, Indonesia. Reinwardtia 22(1): 1‒
25. — Analisis komposisi dan struktur hutan pamah alami dilakukan di Bodogol, Taman Nasional Gunung Gede
Pangrango (TNGGP). Kajian dilakukan di dua petak (P1CS dan P2CS) di bukit Cisuren, dan satu petak (P3CP) di
bukit Cipadaranten. Dalam tiga petak tersebut tercatat 107 jenis dan 48 suku, dengan luas area dasar rata-rata 19,73
m
2
/ha dan kerapatan rata-rata 348 pohon/ha. Kekayaan jenis di lokasi penelitian ini lebih rendah daripada di hutan
hujan pamah Kalimantan dan Sumatra, tetapi sebanding dengan di hutan pegunungan Jawa. Berdasarkan IUCN Red
List Castanopsis argentea dan Castanopsis tungurrut tercatat sebagai jenis kritis dan Saurauia bracteosa sebagai jenis
rentan. Berdasarkan dua jenis dominan hutan di petak penelitian dapat disebut sebagai asosiasi Maesopsis eminii-
Syzygium acuminatissimum dan asosiasi Syzygium acuminatissimum-Lithocarpus korthalsii. Maesopsis eminii
dominan di P1CS (IV=56,46%) dan P3CP (IV=55,94%), sementara Syzygium acuminatissimum dominan di P2CS
(IV=43,67%). Maesopsis eminii adalah jenis invasif yang sangat agresif sehingga membahayakan kemurnian hutan
pegunungan alami TNGGP, oleh karena itu harus diberantas. Secara vertikal, struktur P2CS dan P3CP terdiri dari
empat strata, sementara P1CS tiga strata. Hasil penelitian ini menyimpulkan bahwa luas satu hektar dapat dianggap
sebagai area minimum yang dapat mencerminkan representasi floristik hutan pamah bagian atas dan hutan pegu-
nungan.
Kata kunci: Asosiasi, area minimum, Bodogol, hutan pamah, kekayaan jenis, komposisi jenis, struktur hutan, Taman
Nasional Gunung Gede Pangrango.
https://dx.doi.org/10.55981/reinwardtia.v22i1.4399
https://orcid.org/0000-0002-9019-8094
https://orcid.org/0000-0002-4074-8342
https://orcid.org/0000-0001-7121-9980
https://orcid.org/0000-0001-9656-8095
REINWARDTIA 2 [VOL.22
INTRODUCTION
Studies of flora, composition, and structure of
forests and other vegetation of GGPNP (Gunung
Gede Pangrango National Park) have been
undertaken since the early 19th century (Steenis et
al., 1972, 2006). The GGPNP has been a key for
research on flora, vegetation, and fauna (Karta-
winata & Sudarmonowati, 2022; Rozak et al.,
2016; Steenis et al., 1972, 2006). The first botani-
cal exploration and investigation in Indonesia
were carried out in 1777 in the Gede Pangrango
twin mountains (now within the GGPNP) by Carl
Pehr Thunberg, a Swedish naturalist and student
of Carollus Linnaeus, the father of taxonomy.
He was the first European naturalist who explored
the area. He published the species he collected
in Florula Javanica in 1825 (Kartawinata, 2010;
Kartawinata & Sudarmonowati, 2022; Steenis et
al., 1972, 2006).
The Cibodas Botanical Garden, established on
the slopes of Mt. Gede, in 1852 was the research
station for studies of mountain flora and fauna in
Indonesia. It is part of the GGPNP region
stretching from the lower montane forest at 1,400
m asl (above sea level) to subalpine vegetation on
the top of Mt. Pangrango at 3,019 m asl. Earlier
investigators, who undertook vegetation studies,
included Junghuhn (1845, 1853-1854), Seifriz
(1923, 1924), and Docters van Leeuwen (1933).
Steenis-Kruseman (1953) made a complete biblio-
graphy of the studies in Mt. Gede-Pangrango and
Bogor Botanical Garden. After the 2
nd
World War,
vegetation studies were carried out by many
investigators, including Abdulhadi et al. (1998),
Arrijani (2008), Arrijani et al. (2008), Meijer
(1959), Rollet et al. (1976), Rozak et al. (2016),
Sadili & Alhamd (2012), Sadili et al. (2009),
Sriyanto (1987), Yamada (1975; 1976a, 1976b,
1977), and Zuhri & Mutaqien (2013).
The GGPNP was divided into seve-
ral resorts (regions), including the Bodogol Resort
(BR) for management purposes. BR covers
planted forests, natural upper lowland forests with
an elevation of < 1,000 m asl, and montane forests
with an elevation > 1,000 m asl (Sadili et al.,
2008; Sadili & Alhamd, 2012; Sadili, 2014;
Junaedi et al., 2020). We know very little about
the structure and composition of the lowland
forests in GGPNP and to date only Helmi et al.
(2009), who had investigated a one-hectare plot of
the upper lowland forest at Bodogol. Therefore,
more studies in the lowland forests are necessary
to complement the montane and subalpine forest
data for better scientifically based management of
the GGPNP (Brearley et al., 2019). Species
of Dipterocarpaceae can be found in the Bodogol
lowland forest, including A nisoptera costata,
Vatica sp., Dipetrocarpus hasseltii, D. retusus, D.
gracilis (Ismail et al., 2000; Helmi et al., 2009;
Junaedi et al., 2020). These species are listed as
vulnerable by the IUCN (IUCN, 2013).
Data on the structural characteristics and species
composition of the GGPNP’s lowland forest are
limited to a one-hectare plot near the Research
Station at Bodogol (Helmy et al., 2009). Here we
study the composition and structure of the lowland
forests at Cisuren and Cipadaranten hill ranges in
Bodogol Resort to support data-driven park mana-
gement, including ecological restoration of disturb-
ed forests and degraded lands within the GGPNP
domain.
MATERIALS AND METHODS
Study site
The GGPNP is located in the Bogor, Cianjur,
and Sukabumi Regencies (Kabupaten), West Java,
at 6°10’‒6°51’ South and 106°51’‒107°02’ East.
The study was carried out in May 2022 in Bodogol
Resort, in the administrative region of the Benda
Village, Cicurug District, Sukabumi Regency,
West Java (Fig. 1). The Bodogol resort covered
natural forest areas on the Cisuren and Cipada-
ranten hill ranges, extending from the lowland area
on the western side of the GGPNP towards the top
of Mt. Pangrango. It also included tree plantations
of rasamala (Liquidambar excelsa), damar (Agathis
dammara), and pinus (Pinus merkusii). We select-
ed subjectively the study sites in the lowland
natural forest with slight disturbances at the
Cisuren and Cipadaranten mountain ridges. Three
study plots of one hectare (100 m × 100 m) were
established on the partially flat sections of the
ridges, with steep to very steep slopes (55%‒75%)
on the sides, at elevations of < 1,000 m asl. Plot
1 (designated as P1CS) and plot 2 (P2CS) were
located in the Cisuren hill range, while plot 3
(P3CP) was in the Cipadaranten hill range.
The terrain on the sides of the three plots was
steep slopes leading to river valleys. The Cisuren
river was located in the northern and Tangkil river
on the southern sides of the P1CS and P2CS. The
upper stream of the Cisuren river lies on the
western side of the P3CP and the Cipadaranten
river is on the eastern side of the P3CP. Table 1
summarizes the data on the three plots, including
Global Positioning System (GPS) position, eleva-
tion, the thickness of litter, air humidity, soil
acidity (pH), and disturbances in the three plots at
Bodogol, GGPNP.
Each one-hectare plot was divided into 10 m ×
10 m subplots to measure trees with diameters at
breast height (DBH) ≥ 10 cm. The diameters of all
trees were measured at 1.3 m above the ground.
The total height of each tree was estimated by eye.
The position of each tree was mapped out using the
X and Y coordinates. Each tree was identified in
the field and voucher specimens were collected for
identification at the Herbarium Bogoriense, BRIN
(National Research and Innovation Agency) in
Cibinong Bogor. The measurements of density,
dominance (basal area), and frequency followed
SADILI et al.: Variation composition and structure of natural lowland forest
2023] 3
Fig. 1. Map of the Gunung Gede Pangrango National Park showing the study site at Bodogol Resort
(Redrawn and modified from the map of BBTNGGP 2020).
Cox (1967) and Mueller-Dombois & Ellenberg
(1974, 2016). The construction of the species-area
curve followed Sadili et al. (2018), which was
based on species data from 100 subplots nested
within the plot, i.e. 100 m
2
(10 m × 10 m), 400 m
2
(20 m × 20 m), 900 m
2
(30 m × 30 m),..... 10,000
m
2
(100 m × 100 m).
RESULTS
General habitat conditions around the study
plot
The study plots were located in slightly disturbed
natural forests. Natural disturbances, such as
landslides were absent in P1CS and P2CS. The
canopy cover in P1CS and P2CS varied from 40%
to 75%. Many subplots with open canopies were
found in P1CS (Fig. 7) and P2CS (Fig. 8). The
gaps were filled up with shrubs, climbers, and
herbs, including Bambusa sp., Calamus sp.,
Dinochloa scandens, Dendrocnide stimulans, Meli-
cope latifolia, Ficus sp., Pinanga sp., and Piper
aduncum. In P3CP gaps in the subplots (Fig. 9)
and those resulting from fallen trees were present,
in which shrubs and herbs grew, including Cyathea
sp., Calliandra houstoniana var. calothyrsus,
Etlingera sp., Piper aduncum, and Selaginella
willdenowii.
Species composition
A total of 107 tree species with DBH ≥ 10 cm
and 37 families (Appendix 1), with an average
basal area of 19.73 m
2
/ha, and an average density
of 348 trees/ha, were recorded in the lowland
forests of GGPNP as represented by the three
study plots. Table 2 shows the summary of the
lowland forest vegetation data as recorded in the
three study plots. Maesopsis eminii in P1CS and
P3CP had the highest density (D= 69 trees/ha), ba-
sal area (BA= 2.68 m
2
/ha) and absolute frequ-
ency (F= 12.61%), while in P2CS Syzygium acu-
minatissimum had the highest density (61 trees/
ha), BA (3.81 m
2
/ha), and absolute frequency (F=
12.46%). Based on IV analysis, the dominant
species in P1CS and P3CP were Maesopsis eminii
with IV= 56.46% (P1CS) and IV= 55.93% (P3
CP), while in P2CS was Syzygium acuminatissi-
mum (IV= 43.67%) (Table 3 and Table 4).
Ten species with the highest density (D) and
highest basal area (BA) (Table 3), highest absolute
frequency (F), and highest importance value (IV)
(Table 4) in each plot varied. Among them,
however, three species were always recorded as
the highest in three plots, i.e., Lithocarpus
korthalsii, Syzygium acuminatissimum, and
Lithocarpus pseudomoluccus. Species present in
the P1CS, P2CS, and P3CP but with low values of
REINWARDTIA 4 [VOL.22
D, F, BA, and IV were listed in Group 1 in
Appendix 1, while other species present in one or
two plots were listed in Groups 2‒7 in Appendix
1.
The relationship between the number of species
and BA varied (Fig. 2). The highest number of
species occurred in the BA class of 0.01‒0.09 cm
2
and the highest appeared in P1CS (71 species).
Species with BA of 0.40‒0.49 m
2
appeared only in
P3CP (2 species), while BA > 0.50 occurred in
P2CS (2 species) and P3CP (4 species). The rela-
tionship between the number of trees and stem
diameters also varied (Fig. 3). The highest number
of trees appeared in the 10‒19.99 cm diameter
class and the highest was in P2CS (223 trees).
Trees with a diameter class of 80‒89.9 cm
occurred only in P2CS and P3CP (2 trees,
respectively).
The species-area curves in the three plots
differed as can be seen in Fig. 4. The curves
appeared to begin flattening at 6,400 m
2
(0.64 ha)
in P1CS, and 10,000 m
2
(1 ha) in P2CS and P3CP.
At the points of inflection, the number of species
ranged between 50 and 70. Based on density, the
species composition of P1CS, P2CS, and P3CP
were slightly similar, where the index of similarity
between P1CS, and P2CS was 57% and between
P3CP and combined P1CS, P2CS was 54%.
Of the total 107 species (Appendix 1), we
recorded nine exotic species (8.41%), which were
Maesopsis eminii and Cecropia peltata occurring
in all the plots, Calliandra houstoniana var. calo-
thyrsus, Bellucia pentamera, and Swietenia
mahagoni in P3CP, and Indonesian species but not
native to GGNPP (Dracaena angustifolia, Falca-
taria falcata, Pinus merkusii, and Artocarpus
heterophyllus) were slightly similar, where the
index of similarity between P1CS, and P2CS was
57% and between P3CP and combined P1CS,
P2CS was 54%.
Table 5 shows that six families contained five or
more species in each plot. Lauraceae, Fagaceae,
and Euphorbiaceae occurred in all plots, while
Moraceae in two plots (P2CS and P3CP), and
Meliaceae in one plot (P3CP). Lauraceae (11
species) in P1CS was the largest family and other
families contained less than five species, each
(Appendix 1).
Table 1. GPS position, elevation, litter thickness, air humidity, soil acidity (pH), and disturbances in the
three plots at Bodogol GGPNP.
Plot
GPS position
Elevation
(m)
Litter
Mean air
humidity
(%)
Mean
pH
Disturbance
Latitude (S) Longitude
(E)
P1CS -06
o
46'52.6" 106
o
51.09.4" 713 Thick 90 7 None
P2CS -06
o
46'53.1" 106
o
51.23.1" 767 Thick 89 7 None
P3CP -06
o
46'40.2" 106
o
51.41.1" 843 Thin 87 6.8 Slight (human activi-
ties along forest trails)
No
Plot
Species
Family
D (ha)
BA (ha)
Index
Diversity (H’) Evennes (E)
1 P1CS 59 25 283 13.79 3.33 0.82
2 P2CS 64 27 385 24.83 3.52 0.85
3 P3CP 74 29 376 20.57 3.61 0.83
Mean 66 27 348 19.73 3.49 0.83
Table 2. Summarized data show the number of species, the number of families, Density (D), Basal Area
(BA Index (H’), and Evenness Index (E) in three plots at the lowland natural forest Bodogol, GGPNP.
SADILI et al.: Variation composition and structure of natural lowland forest
2023] 5
Table 3. Ten species with the highest density (D) and basal area (BA) in three plots at the lowland natural
forests in Bodogol, GGPNP.
No Species Density (trees/ha) Basal area (m
2
/ha)
P1CS P2CS P3CP P1CS P2CS P3CP
1 Syzygium acuminatissimum 19 61 21 1.20 3.81 1.46
2 Maesopsis eminii 69 7 51 2.68 0.69 6.54
3 Lithocarpus
pseudomoluccus
16 22 13 1.41 1.85 0.49
4 Schima wallichii 6 11 20 0.64 2.74 1.43
5 Lithocarpus korthalsii 11 25 16 1.55 2.32 0.78
6 Macaranga denticulata 9 12 12 0.17 0.27 0.23
7 Castanopsis tungurrut 12 4 11 0.59 0.26 0.34
8 Neoscortechinia kingii 10 17 4 0.27 0.72 0.10
9 Sandoricum koetjape 1 12 8 0.36 0.36 0.47
10 Pometia pinnata 7 8 7 0.34 0.21 0.41
11 Nephelium juglandifolium 6 25 4 0.29 1.53 0.22
12 Ficus padana 7 1 1 0.15 0.02 0.10
13 Aglaia elliptica 2 2 5 0.09 0.22 0.54
14 Castanopsis argentea 1 4 2 0.02 0.15 0.42
15 Beilschmiedia madang 7 6 0.31 0.22
16 Neonauclea lanceolata 7 0.32
17 Donella lanceolata 1 0.37
18 Syzygium antisepticum 13 31 0.37 1.24
19 Didymocheton nutans 14 7 0.79 0.42
20 Symplocos acuminata 8 9 0.18 0.12
21 Chisocheton ceramicus 2 3 0.97 0.07
22 Elaeocarpus angustifolius 2 2 0.71 0.06
23 Spondias pinnata 3 0.98
24 Calliandra houstoniana
var. calothyrsus
28 0.33
REINWARDTIA 6 [VOL.22
Forest structure
Forest profile diagrams (Fig. 6) for all plots
were constructed following the method of
Kartawinata et al. (2004) as applied by Rahma et
al. (2016). The profile diagrams show the tree
heights that jointly form the vertical stratification
(A Stratum to D Stratum), which varied from one
plot to another. The ten main tree species with the
highest density composing A Stratum to C Stra-
tum are presented in Table 6. Fig. 5 shows the
relationship between the diameters and heights of
trees within the three plots revealing that most
trees were concentrated in the 10‒20 cm diameter
class with heights of < 20 m (C Stratum). In P1CS
no trees were forming the emergent A Stratum
(height > 34 m). In P2CS, five trees of five
species constituted the A Stratum and in P3CP
there was only one tree formed the emergent A
stratum (Fig. 5 and Fig. 6).
The distribution of the exotic species,
Maesopsis eminii, in each plot was scattered
(Figs. 7‒9, in red). The density of Maesopsis
eminii in P1CP (69 trees/ha) was higher than that
in P2CS (7 trees/ha) and P3CP (51 trees/ha) (Table
3). Observations during the field study revealed
that the fruits of Maesopsis eminii were consumed
daily by Owa Jawa (Hylobates moloch), Lutung
Jawa (Trachypithecus auratus), Surili (Presbytis
comata) and Tupai (Tupaia sp.).
DISCUSSION
Species Composition
Of the total 107 species recorded in the three
plots (Appendix 1), 48 species (44.86%) were not
registered in the Gunung Gede Pangrango flora
(Sunarno & Rugayah, 1992), and in Helmi et al.
(2009), whose study site was located in the same
forest area. They are here considered as the
lowland forest species which were confirmed
Table 4. Ten species with the highest absolute frequency (AF) and the highest importance value (IV) in
three plots at the lowland natural forests in Bodogol, GGPNP.
No Species Absolute Frequency (F= %) Importance Value (IV= %)
P1SC P2CS P3CP P1SC P2CS P3CP
1 Maesopsis eminii 12.61 1.87 10.58 56.46 6.48 55.93
2 Syzygium acuminatissimum 7.66 12.46 4.49 23.10 43.67 17.15
3 Lithocarpus korthalsii 3.60 6.85 4.81 18.74 22.68 12.84
4 Lithocarpus
pseudomoluccus
5.41 6.23 3.53 21.32 19.40 9.34
5 Neoscortechinia kingii 4.05 4.05 1.28 9.55 11.38 2.81
6 Schima wallichii 2.25 2.49 4.49 9.03 16.38 16.77
7 Castanopsis tungurrut 4.95 1.25 2.88 13.50 3.33 7.46
8 Macaranga denticulata 3.15 2.80 2.56 7.56 7.01 6.88
9 Nephelium juglandifolium 2.70 5.92 0.96 6.90 18.58 3.07
10 Pometia pinnata 3.15 2.49 2.24 8.09 5.40 6.09
11 Cecropia peltata 1.35 0.31 5.13 2.65 1.53 5.44
12 Sandoricum koetjape 0.45 3.43 2.24 3.42 8.01 6.68
13 Syzygium antisepticum 2.80 5.77 7.67 20.05
14 Didymocheton nutans 3.12 1.92 9.94 5.83
15 Neonauclea lanceolata 3.15 7.98
16 Calliandra houstoniana
var. calothyrsus
5.13 14.18
SADILI et al.: Variation composition and structure of natural lowland forest
2023] 7
through comparison with Backer & Bakhuizen van
den Brink (1963-1968), Djarwaningsih et al.
(2010), Ismail et al. (2000), Kartawinata (1977),
Rozak et al. (2016), and Yusuf (2004). They
contributed to the increase of the current GGNPP
floristic diversity of 1,103 species (Kartawinata &
Sudarmonowati, 2022), which should now total
1,151 species.
The present study, along with Helmy et al.
(2009) and Sadili (2014), has described the
transitional forest area between the lowland and
montane forests with tree species diversity that
was not known when GGPNP was established in
1980. It is interesting to note that species of
Dipterocarpaceae were reported to occur spa-
ringly in Bodogol (Ismail et al., 2000; Helmi et
al., 2009; Junaedi et al., 2020) but they were
not found in the present study plots, possibly
because of unsuitable microclimate and located
far away from the seed producing mature trees
(Purwaningsih, 2004).
Ten species with the highest density (D), basal
area (BA) (Table 3), absolute frequency (F), and
important value (IV) (Table 4) varied. Based on
the dominance as reflected by the highest IV
analysis the forest in P1CS and P3CP, on the one
hand, may be called Maesopsis eminii-Syzygium
acuminatissimum association, with the character
species that were confined to the two plots as
listed in Group 3 in Appendix 1. The forest in
Fig. 2. The relationship between the number of species and BA classes in three plots at the lowland
natural forests in Bodogol, GGPNP.
Fig. 3. The relationship between density (D) and diameter class in three plots at the lowland forest in
Bodogol, GGPNP.
REINWARDTIA 8 [VOL.22
Fig. 4. Species-area curves in three plots of one hectare each in the lowland natural forests in Bodogol,
GGPNP.
Table 5. The number of families having five or more species in the three plots at the lowland natural
forests Bodogol, GGPNP.
No Family Number of species
P1CS P2CS P3CP
1 Lauraceae 11 6 7
2 Fagaceae 6 7 6
3 Euphorbiaceae 5 6 8
4 Moraceae 6 6
5 Meliaceae 6
Fig. 5. Scattered diagram of diameter classes and tree heights in the three plots at the lowland natural
forests at Bodogol, GGPNP.
SADILI et al.: Variation composition and structure of natural lowland forest
2023] 9
Fig. 6. Profile diagrams of the forest on the plots constructed by plotting the height of each tree and
sequential tree position from tree no. 1 in the 1st subplot up to the last tree in the 100
th
subplot in all plots in
the lowland natural forests at Bodogol, GGPNP, using the method of Kartawinata et al. (2004).
Tree
Height
(m)
Tree
Height
(m)
Tree
Height
(m)
REINWARDTIA 10 [VOL.22
Table 6. The number of species, number of trees, and ten species with the highest density in the A, B, and C
strata in the three plots in the lowland natural forests at Bodogol, as graphically presented in Fig. 9.
Plot
Stratum
Number Ten species with the highest density (trees/ha)
Species Tree
A
(Height > 30 m)
-
-
-
P1CS
B
(Height: 20‒30 m)
23
50
Maesopsis eminii (8), Syzygium acuminatissimum
(7), Lithocarpus korthalsii (7), Lithocarpus
pseudomoluccus (5), Schima wallichii (2),
Neoscortechinia kingii (2), Castanopsis tungurrut
(2), Neonauclea lanceolata (2), Adina trichotoma
(1), Liquidambar excelsa (1)
C
(Height: 4‒19.9 m)
55
233
Maesopsis eminii (61), Syzygium acuminatissimum
(2), Lithocarpus pseudomoluccus (12),
Castanopsis tungurrut (10), Neoscortechinia kingii
(10), Macaranga deltoidea (8), Beilschmiedia
madang (7), Ficus padana (7), Nephelium
junglandifolium (6), Pometia pinnata (6)
P2CS
A
(Height: > 30 m)
5
5
Chisocheton ceramicus (1), Lithocarpus korthalsii (1),
Lithocarpus pallidus (1), Schima wallichii (1),
Dalrympelea sphaerocarpa (1)
B
(Height: 20‒30 m)
33
107
Maesopsis eminii (8), Syzygium acuminatissimum
(7), Lithocarpus korthalsii (7), Lithocarpus
pseudomoluccus (5), Schima wallichii (2),
Neoscortechinia kingii (2), Castanopsis tungurrut
(2), Neonauclea lanceolata (2), Adina trichotoma
(1), Liquidambar excelsa (1)
C
(Height: 4‒19.9 m)
55
273
Calliandra houstoniana var. calothyrsus (31),
Syzygium antisepticum (27), Maesopsis eminii
(22), Syzygium acuminatissimum (13), Schima
wallichii (13), Lithocarpus korthalsii (12),
Macaranga deltoidea (12), Castanopsis tungurrut
(11), Symplocos acuminata (10), Lithocarpus
pseudomoluccus (9)
P3CP
A
(Height: > 30 m)
1
1
Maesopsis eminii (1)
B
(Height: 20‒30 m)
24
76
Syzygium acuminatissimum (25), Lithocarpus
korthalsii (14), Nephelium junglandifolium (9),
Lithocarpus pseudomoluccus (8) Schima wallichii
(7), Neoscortechinia kingii (4), Beilschmiedia
madang (3), Didymocheton nutans (3),
Sandoricum koetjape (3), Spondias pinnata (3)
C
(Height: 4‒19.9 m)
69
299
Syzygium acuminatissimum (36), Nephelium
juglandifolium (16) Lithocarpus pseudomoluccus
(13), Macaranga deltoidea (12), Neoscortechinia
kingii (12), Didymocheton nutans (11),
Lithocarpus korthalsii (11), Syzygium antisepticum
(10), Dendrocnide stimulans (9), Myristica sp. (9)
SADILI et al.: Variation composition and structure of natural lowland forest
2023] 11
P2CS, on the other hand, may be designated as
Syzygium acuminatissimum-Lithocarpus korthalsii
association, characterized by species that were
confined to P2CS, as listed in Group 6 in
Appendix 1. Before the Maesopsis eminii invasion,
the Syzygium acuminatissimum-Lithocarpus kor-
thalsii association could have been prevalent in
the lowland forests at Bodogol. In these
associations, the following species were registered
in the IUCN Red List (Effendi et al., 2022;
Wihermanto, 2007) as critical (Castanopsis argen-
tea and C. tungurrut) and vulnerable category
(Saurauia bracteosa).
The number of species along the basal area
classes (Fig. 2) and diameter classes (Fig. 3)
showed that the highest number of species
occurred in the lowest BA class (0.01‒0.09 m
2
)
and diameter class (10.0‒19.9 cm). It follows that
the number of species decreased as the BA and
diameter increased. The BA is related to the
diameter. The above reverse-J pattern is typical for
the tropical rainforest and indicated that the forest
regenerated well and was in a dynamic state
(Ogawa et al., 1965; Richards, 1996; Mirmanto,
2014).
The species richness and tree density in our
plots were comparable to those obtained from
various studies in the montane forests of Mount
Gede-Pangrango and Mount Halimun in West
Java, Foja Mountains in the upper montane forest
of Papua, but lower than those of the studies in
Sumatra (Table 7). From Table 7 it is evident that
the montane forests, even the pristine forest un-
touched by human activities in the Foja Mountains
in Papua (Sadili et al., 2018), and the transitional
lowland-montane forests at Bodogol had a low
species diversity of 59‒70 species in one hectare,
compared to very species-rich undisturbed lowland
forests at Kalimantan (Sheil et al., 2010), and Su-
matra (Kartawinata et al., 2004), where the number
of species per hectare was 205 and 182, respec-
tively. Studies in Sulawesi by Brambach et al.
(2017) in several plots with a total area of 3.1 ha in
the Lore Lindu National Park at 700–2,400 m asl
and by Trethowan et al. (2019) in several plots
with a total area of 2.5 ha in lowland forest areas at
Morowali, Wawonii and Bualemo, show the mean
species richness of 9/ha and 113/ha, respectively,
which are comparable to the result of the present
study. It may be implied that it is a norm for any
montane forest to have low species diversity. The
richness of a plot may be due to disturbance by
facilitating species dependent on disturbance to
associate very closely with late-successional speci-
es (Connell, 1978; Sheil & Burslem, 2003; Slik et
al., 2008).
The occurrence of low species richness is
reflected also in a species-area curve. The species-
area relationship is essential to ecology (Plotkin et
al., 2000) and it can also be used to approximate
species extinction resulting from habitat loss (May
et al., 1995; Pimm & Raven, 2000) and to assess
patterns of species diversity in different forest
types (Ashton, 1965; Mueller-Dombois & Ellen-
berg, 1974, 2016). The curve was constructed to
indicate species diversity in relation to the increas-
ing size of the area (Mueller-Dombois & Ellen-
berg, 1974, 2016) within the plot. Fig. 4 shows the
species-area curves in three plots with an area of
one hectare each. The species-area curves in the
three plots differed. The curves began to flatten at
6,400 m (0.64 ha) in P1CS and at 10,000 m (1 ha)
in P2CS and P3CP, which indicated that at the
points of inflection, the number of species ranged
between 50 and 70. It is in contrast to the species-
area curves of undisturbed lowland forests in
Kalimantan and Sumatra. In Kalimantan, Kartawi-
nata et al. (2008) showed that the curve still steadi-
ly rose to indicate no sign of leveling at the area of
10.5 ha with a number of species of 552 and in
Sumatra. Kartawinata et al. (2004) noted the same
pattern with no indication of flattening of the curve
at one hectare and 182 species.
Table 2 shows the Diversity Index (DI) with a
mean value of H’= 3.49 may be considered high,
comparable to H’= 3.39 at higher elevations (1,000
m asl) of the GGNP forest (Zuhri & Mutaqien,
2011) but higher than the diversity index at the
montane forest at the Mt. Ceremai National Park
with H’= 2.66‒2.77 (Purwaningsih & Yusuf,
2008). The diversity index is considered high if
H’ > 3 and H’ in the tropical forests is 1.5–3.5 and
rarely H’ > 4 (Greig-Smith, 1983). The Evenness
Index (E) is used as the indicator of prevalence in
the forest community. In the present study plots E
values were high (0.82‒0.85), which were indi-
cating that all the species were even relatively
(Ludwig & Reynold, 1988; Oosting, 1956).
Exotic species, other than Maesopsis eminii,
(i.e. Bellucia pentamera, Calliandra houstoniana
var. calothyrsus, Cecropia peltata, and Swietenia
mahagoni) and Malesian species but not native to
GGPNP (i.e., Artocarpus heterophyllus, Dracaena
angustifolia, Falcataria falcata, and Pinus
merkusii) were contaminating but not aggressively
invading the natural tree communities, as indicated
by their very low densities and basal areas
(Appendix 1). They should be, however,
monitored and prevented from spreading more
widely and even eradicated.
Maesopsis eminii, which was introduced to
Indonesia for the first time in 1920, should receive
special attention because this species aggressively
and rapidly invaded submontane rain forests and
became the dominant species in Eastern Tanzania
(Schabel & Latiff, 1997). In Bodogol it dominated
the lowland forest, as indicated in the study plots
(Appendix 1, Figs. 7‒9, and Table 4), it reached
the highest IV in P1CS (56.46%) and P3CP
(55.93%), while in P2CS it was one of the 11 main
REINWARDTIA 12 [VOL.22
Fig. 7 The scattered and dispersed tree distribution of trees with DBH ≥ 10 in the P1CS at the lowland
natural forest Bodogol, GGPNP.
Fig. 8. The scattered and dispersed tree distribution of trees with DBH ≥ 10 in the P2CS at the lowland
natural forest Bodogol, GGPNP.
SADILI et al.: Variation composition and structure of natural lowland forest
2023] 13
species with IV= 6.48%. Naturally, the species
occurred in Africa from lowland tropical rain
forests to savannas, extending to montane forests
at the altitude of 1,800 m asl and dispersed by
birds (especially hornbills), rodents, and monkeys
(Schabel & Latiff, 1997). It is a fast-growing
species with a growth rate of 1.5‒5.5 cm per year
in diameter and 1‒3 m per year in height (Schabel
& Latiff, 1997) and could grow well at the
altitudes of 100‒900 m asl within full sunshine. It
was planted in community forests and plantations
in West Java in 1960 by the Forestry Department
(Samsoedin et al., 2016). Informants (pers. comm.,
2022) stated that it was planted in the community
forests in Bodogol village in the 1970s.
In Bodogol, many primate species (Owa Jawa-
Hylobates moloch, Lutung Jawa-Trachypithecus
auratus, and Surili-Presbytis comata) and rodents
(Tupaia sp.) living in the GGPNP vigorously
consumed the fruits of M. eminii (Helmi et al.,
2009) and later dispersed seeds germinated and
developed into large trees elsewhere in the interior
of natural forests as indicated in the results of the
present study as far as 1,000 m from the natural
forest edges. In Africa, it is an amazingly long-
lived pioneer species that could live up to 150
years (Schabel & Latiff, 1997). Considering the
above growth and behavioral characteristics of the
species and the current dominating existence in
Bodogol inland natural forests it would be not
surprising that M. eminii will become a dominant
species in the upper montane forests and even in
the subalpine forests of GGPNP. A drastic action
to eradicate M. eminii from the natural forest of
GGPNP and the adjacent areas of GGPNP should
be seriously undertaken without delay to save the
continuing existence of natural forests in GGPNP.
Falcataria falcata and Pinus merkusii are
native to Indonesia and were present in the study
plots but at low density (Appendix 1), hence not
aggressive invaders. They were planted by the
Forest Department in the vicinity of the GGPNP
and those in Bodogol have been incorporated into
the GGPNP. The presence of A rtocarpus hetero-
phyllus and Dracaena angustifolia could have
been due to human activities that inadvertently
carry the propagules into the natural forests (An-
dila & Warseno, 2019; Samsoedin et al., 2016).
Species having high commercial value is
Liquidambar excelsa (Sutomo & Sari, 2019). It is
native to Bhutan, Assam, Myanmar, Peninsular
Malaysia, Sumatra, and Java. Growing at the ele-
vation of 550‒1,700 m asl (Vink, 1957). In P1CS
it had D= 3/ha, BA= 0.07 m
2
and IV= 2.89% (Ap-
pendix 1). In Bodogol, it was recorded by Ismail
et al. (2000), Helmi et al. (2009), Gunawan et al.
(2011), Sadili & Alhamd (2012), and Sadili
(2014). In Cibodas, grew on the submontane to
montane forests (Meijer, 1959; Yamada, 1975;
Abdulhadi et al., 1998; Arrijani et al., 2006;
Rozak et al., 2016). Local people informed us that
seedlings and saplings of A . excelsa were hard to
find in the natural forests at Bodogol. Similarly,
the planted forest of A . excelsa in Bodogol, which
according to informants in the village (pers.
comm., 2022) was planted in 1918. Sadili (2014)
Fig. 9. The scattered and dispersed tree distribution of trees with DBH ≥ 10 in the P3CP at the lowland
natural forest Bodogol. A relatively large gap (dark green colour) resulting from fallen trees was present.
REINWARDTIA 14 [VOL.22
Table 7. Comparison of the number of species and number of trees in one-hectare plots in montane forests
of Java and Papua and the lowland forests of Sumatra and Kalimantan.
Elevation
(m)
Plot size
(ha)
Density
(trees/ha)
Number
of species
Source Locality
WEST JAVA
Gunung Gede pangrango National Park
Bodogol P1CS 713 1 283 59 Present study
P2CS 767 1 385 64 Present study
P3CP 843 1 376 74 Present study
Mean 713-843 1 348 66 Present study
Bodogol 806 1 352 70 Helmi et al.
(2009)
Cibodas 1600 1 427 57 Yamada (1975)
Gunung Halimun Salak National Park
Gunung Kendeng 1000 1 406 64 Suryanti (2006)
Gunung Malang 1000 1 421 69 Suryanti (2006)
Gunung Panenjoan 1000 1 405 69 Suryanti (2006)
Purabakti 900 1 441 69 Yusuf (2004)
KALIMANTAN
Malinau 200 1 759 205 Sheil et al.
(2010)
SUMATRA
Batang Gadis National
Park
660 1 583 182 Kartawinata et
al. (2004)
Bukit Lawang 297 1 453 216 Polosakan
(2001)
PAPUA
Foja Mountains 1710 1 693 59 Sadili et al.
(2018)
SADILI et al.: Variation composition and structure of natural lowland forest
2023] 15
found only two saplings (diameter= 8.62 cm and
9.39 cm), indicating that the species had a poor
regenerating capability. An officer of the Halimun-
Salak National Park (pers. comm., 2022) informed
that seedlings and saplings of A . excelsa could be
found only in secondary growth on small
landslides near mature trees. Qualitative
observation revealed that the absence of
regeneration in the old A . excelsa planted in
Bodogol could be related to the unsuitable
elevation of less than 1,000 m asl for growth and
reproduction. It could be inferred that planted A .
excelsa had a low ability to sustain itself with the
implication that the species may not be a good
candidate to be used in ecological restoration but
acceptable for rehabilitation of degraded lands.
Native tree species that are potentially good for
ecological restoration is S. wallichii. It is a variable
species, consisting of nine subspecies and five
varieties (Bloembergen, 1952), including S. walli-
chii subsp. noronhae. In Java, it is spreading from
300 m to 2,600 m asl, which comprised two
varieties (Backer & Bakhuizen van den Brink,
1963). Many pioneer species inhabiting the low-
land to montane forests are amazingly long-lived
(Whitmore, 1986) and S. wallichii could be placed
in this category, as can be seen, it lived in the
primary montane forest of GGPNP (Kartawinata &
Sudarmonowati, 2022; Yamada, 1974, 1976). In
the present study, it occurred in all plots (Table 3,
Table 4, and Appendix 1). Vertically (Table 6)
filled up the B stratum (height 20‒30 m) and C
stratum (height 4‒20 m). Gunawan et al. (2011)
noted that its distribution was clustered and based
on high IV it was predicted that S. wallichii could
become dominant in the A gathis dammara planta-
tion. In the Kawah Ratu (Halimun-Salak National
Park), Hilwan & Rahman (2021) reported good
regenerations of S. wallichii. Rozak et al. (2016) in
GGPNP, S. wallichii occurred from the submon-
tane to the subalpine forests.
Lauraceae, Fagaceae, and Euphorbiaceae are
common families in the submontane forests
(Steenis et al., 2006; Purwaningsih & Polosakan,
2016), hence, the forest is often designated as
Lauro-Fagaceous forest (Steenis & Shippers-
Lammertse, 1965; Steenis et al., 2006). In the
present study, families in the submontane forest
with a high number of species (> 5) were Laurace-
ae, Fagaceae, and Euphorbiaceae (Table 5). Eu-
phorbiaceae contained the highest number of
species (eight species) in P3CP, which was related
to high adaptability to the environment (Kartawi-
nata, 1977; Riswan, 1987). Based on the number of
species, in Malesia, Euphorbiaceae is listed as one
of the four big family groups (Whitmore, 1995).
The existence of Euphorbiaceae in three plots
could be implied that the habitat was once
disturbed, such as fallen trees creating gaps, which
in turn stimulated seeds to germinate.
Structure
Species distribution in each plot varied
(Appendix 1). The number recorded in three plots
accounted for 29 species (27.10%), in two plots 31
species (28.97%), and only in one plot 47 species
(43.93%). It is implied that each species had
different characteristics in its regeneration and sur-
vival. The species that had the widest distribution
were tolerant to the varied environment of dif-
ferent forests (Group 1 in Appendix 1). Magnolia
montana (IV= 0.62%) and Neesia altissimo (IV=
0.62%) were species with the lowest IV and occur-
red only in P3CP, which could be due to the
presence of the highest number of exotic species in
the plot. It needs further research to clarify the
phenomenon.
Forest profile showing the vertical structures
(Figs. 5 and 6), containing ten species with varied
highest diversity in each stratum of each plot
(Table 6). The A stratum (height > 30 m) occurred
in P2CS (5 trees/ha) and P3CP (1 tree/ha). Further-
more, the correlation between diameter class and
tree height of < 20 m (C Stratum), and in general
diameters are correlated with tree height, implying
that the bigger the diameters the taller the trees,
hence the forest canopy. Fig. 5, illustrating the
profile of the P2CS, revealed the presence of trees
with small diameters but with very tall trunks,
implying that the trees optimally utilized the light
along with the others. This phenomenon could be
observed in Lithocarpus korthalsii. Figs. 7–9 show
the uneven distribution of all trees in the plots. The
distribution of Maesopsis eminii was concentrated
on the more or less flat ridges of the hills and
decreased towards the sloping sides. It should be
pointed out that in some subplots with open
canopies and in such openings, many species of
young trees, shrubs and lianas grew profusely,
including Bambusa sp., Calamus sp., Dendrocnide
stimulans, Dinochloa scandens, Melicope latifolia,
Ficus sp., Pinanga sp., and Piper aduncum. Forest
structure in Bodogol is comparable to those at
1,600 m asl (Kartawinata & Sudarmonowati,
2022; Yamada, 1975).
Successional status of the forest
We checked the species listed in Appendix 1 to
identify the successional status of species using the
following references, Kramer (1926, 1933),
Backer & Bakhuizen van den Brink, Jr. (1963‒
1968), Steenis et al. (1972), Riswan (1984),
Whitmore (1986), Sriyanto (1987), Sunarno &
Rugayah (1992), Keβler et al. (2000), and
Kartawinata & Sudarmonowati (2022). Appendix
1 recorded the following 32 secondary forest spe-
cies in the plots i.e. Liquidambar excelsa,
Artocarpus elastica, Bellucia pentamera,
Blumeodendron tokbrai, Glochidion rubrum var.
rubrum, Calliandra houstoniana var. calothyrsus,
Castanopsis argentea, C. tungurrut, Cecropia pel-
REINWARDTIA 16 [VOL.22
tata, Decaspermum fruticosum, Dendrocnide
stimulans, Melicope latifolia, Ficus tinctoria
subsp. gibbosa, Ficus fistulosa, Ficus padana,
Ficus variegata, Polyspora excelsa, Knema
laurina, Litsea noronhae, Maesopsis eminii,
Mallotus paniculatus, Neonauclea lanceolata,
Homalanthus populneus, Falcataria falcata, Pinus
merkusii, Podocarpus neriifolius, Pometia pinata,
Schima wallichii, Sterculia chrysodasys, Swietenia
mahagoni, Strobocalyx arborea and Oreocnide
rubescens. The fact that they constituted 29.9% of
the total species recorded in the three plots,
combined with many open subplots and the
presence of gaps in the three plots (Figs. 7‒9),
pointed to the somewhat secondary nature of the
lowland forests of Bodogol. It was resulted from
human disturbances and natural disruptions such
as landslides and thunderstorms, which occasion-
ally took place in the area. Structural charac-
teristics showing low forest heights (Fig. 5 and
Table 6) support this contention. It can be inferred
that the forests in the plots were regenerating after
disturbances in the past.
CONCLUSION
The forests investigated contained a total of 107
species recorded in three one-hectare plots.
Lowland species were prevalent, thus forming
the transition between lowland and lower montane
forests, with relatively poor species richness. It
was much poorer than the typical lowland rain
forest of Kalimantan and Sumatra but comparable
to those of the montane forests of Java. Based
on two dominant species, the forest could be
designated as the Maesopsis eminii-Syzygium acu-
minatissimum association and Syzygium acumina-
tissimum-Lithocarpus korthalsii association. Be-
fore M. eminii invasion the Syzygium acuminatis-
simum-Lithocarpus korthalsii association could
have been prevalent in lowland forests at
Bodogol. The trees native to Mt. Gede-Pangrango
recorded in the present studies are the potential
species for ecological restoration undertaken
around the GGPNP.
This study strongly confirmed that the one-
hectare area could be considered a minimal area to
reflect the representativeness of the floristic
composition. The dominance of M. eminii, point-
ed to a strong aggressive, and invasive species,
which in no time will frighteningly take over the
dominance of the lowland to subalpine forests of
GGPNP. It, therefore, is very urgent to totally
eradicate it at all costs from the natural forests,
the plantations, and community forests adjacent to
the GGPNP. Structurally and floristically the fo-
rest represented a developing and regenerating
disturbed forest, and very low frequency and
density in the majority of the species reflected
species heterogenity. Investigations, similar to the
present study, should be extended to other un-
described natural lowland forests of other resorts
for better management, including ecological resto-
ration.
ACKNOWLEDGEMENTS
We are grateful to the Head of the Research
Center for Ecology and Ethnobiology, National
Research and Innovation (BRIN) for the research
permission. We also thank the Head of the
GGPNP, the Head of the Bodogol Resort of
GGPNP, Director of Scientific Collection Mana-
gement – BRIN, and other staff members of the
GGPNP for the provision of various facilities.
Our special gratitudes go to Pak Ae and Pak
Syarif for their remarkable assistance in the field.
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REINWARDTIA 20 [VOL.22
Appendix 1. Species present in the plots of the lowland natural forests at Bodogol GGPNP with their
parameters on Density (D=trees/ha), Basal area (BA= m
2
), and Importance Value (IV= %) The NSR and
NH are considered lowland species following Backer & Bakhuizen van den Brink (1963-1968).
Note; L= lowland; M=Montane; A=Alien; NSR=Non Sunarno & Rugayah (1992); NH=Non Helmi et al.
(2009):
No
Species
Family
P1CS P2CS P3CP
Note D
(Ha)
BA
(m2/
ha)
IV (%) D
(Ha)
BA
(m2/
ha)
IV (%) D
(Ha)
BA
(m2/
ha)
IV (%)
1 2 3 4 5 6 7 8 9 10 11 12 13
GROUP 1
1 Maesopsis eminii
Engl.
Rham. 69 2.68 56.46 7 0.69 6.48 51 6.54 55.93 L.M.A.N
H
2 Syzygium
acuminatissimum
(Blume) DC.
Myrt. 19 1.20 23.10 61 3.81 43.67 21 1.46 17.15 M.NH
3 Lithocarpus
pseudomoluccus
(Blume) Rehder
Fag. 16 1.41 21.32 22 1.85 19.40 13 0.49 9.34 M.NH
4 Lithocarpus
korthalsii (Endl.)
Soepadmo
Fag. 11 1.55 18.74 25 2.32 22.68 16 0.78 1.84 M.NH
5 Castanopsis
tungurut (Blume)
A.DC.
Fag. 12 0.59 13.50 4 0.26 3.33 11 0.34 7.46 M.NH
6 Neoscortechinia
kingii (Hook.f.)
Pax & K.Hoffm
Euph. 10 0.27 9.55 17 0.72 11.38 4 0.10 2.81 M.NSR.N
H
7 Schima wallichii
(DC.) Korth.
Thea. 6 0.64 9.03 11 2.72 16.38 20 1.43 16.77 L.M.NH
8 Pometia pinnata
J.R.Forst. &
G.Forst.
Sapin. 7 0.34 8.09 8 0.22 5.40 7 0.41 6.09 L.M.NSR.
NH
9 Macaranga
denticulata
(Blume)
Müll.Arg.
Euph. 9 0.17 7.56 12 0.27 7.01 12 0.23 6.88 M.NSR.N
H
10 Nephelium
juglandifolium
Blume
Sapin. 6 0.29 6.90 25 1.53 18.58 4 0.22 3.07 M.NSR.N
H.
11 Ficus
padana Burm.f.
Mor. 7 0.15 6.25 1 0.02 0.66 1 0.10 1.07 L.M.NSR.
NH
12 Strobocalyx
arborea (Buch.-
Ham.) Sch.Bip.
Astera. 3 0.24 4.17 1 0.01 0.61 2 0.04 1.39 M.NH
13 Knema
cinerea (Poir.)
Warb.
Myrist. 4 0.08 3.80 2 0.21 1.93 6 0.33 5.13 L.M.NSR.
NH
14 Melicope latifo-
lia (DC.)
T.G.Hartley
Rut. 4 0.05 3.60 3 0.18 2.44 4 0.25 3.55 M.NSR
SADILI et al.: Variation composition and structure of natural lowland forest
2023] 21
15 Sandoricum
koetjape
(Burm.f.) Merr.
Melia. 1 0.36 3.42 12 0.36 8.01 8 0.47 6.68 L.M.NSR.
NH
16 Gynotroches
axillaris Blume
Rhiz. 3 0.10 3.17 7 0.35 5.08 4 0.09 2.80 M
17 Persea venosa
Nees & Mart.
Laur. 2 0.15 2.68 1 0.03 0.70 1 0.12 1.16 M.NH
18 Cecropia
pachystachya
Trécul
Urt. 3 0.03 2.65 4 0.04 1.53 1 0.01 5.44 L.A.NH
19 Dalrympelea
sphaerocarpa
(Hassk.) Nor-
Ezzaw.
Staph 3 0.08 2.57 8 0.45 6.08 1 0.01 0.64 M
20 Myristica sp. Myrist. 2 0.12 2.48 9 0.16 5.78 1 0.02 0.66 M.NH
21 Aglaia elliptica
(C.DC.) Blume
Melia. 2 0.09 2.25 2 0.22 2.02 5 0.54 5.56 M.NSR.N
H
22 Litsea resinosa
Blume
Laur. 2 0.04 1.87 7 0.24 4.98 6 0.11 4.05 M.NH
23 Mallotus
paniculatus
(Lam.) Müll.Arg.
Euph. 2 0.03 1.85 1 0.04 0.75 3 0.14 2.45 M.NSR
24 Symplocos
acuminata
(Blume) Miq.
Sympl. 2 0.02 1.77 2 0.12 1.61 6 0.08 3.28 M.NH
25 Dendrocnide
stimulans (L.f.)
Chew
Urt. 2 0.02 1.30 9 0.17 5.82 5 0.05 3.17 M.NH
26 Quercus
gemelliflora
Blume
Fag. 1 0.06 1.25 1 0.03 0.70 1 0.01 0.66 M.NSR.N
H
27 Antidesma
velutinosum
Blume
Euph. 1 0.02 0.97 1 0.09 0.93 1 0.01 0.64 L.NGPF
28 Castanopsis
argentea (Blume)
A.DC.
Fag. 1 0.02 0.96 4 0.15 2.27 2 0.42 3.22 M.NH
29 Oreocnide
rubescens
(Blume) Miq.
Urt. 1 0.01 0.88 9 0.13 4.73 7 0.12 4.06 L.M
GROUP 2
30 Beilschmiedia
madang (Blume)
Blume
Laur. 7 0.31 7.42 6 0.22 4.33 M.NSR
31 Syzygium
koghianum
Petitm. & Bonati
Myrt. 6 0.21 6.35 4 0.13 2.81 M.NH
32 Astronia
macrophylla
Blume
Melast. 3 0.19 3.81 1 0.07 0.87 M.NSR
33 Lithocarpus
pallidus (Blume)
Rehder
Fag. 2 0.10 2.35 5 0.69 5.32 M.NH
REINWARDTIA 22 [VOL.22
34 Glochidion
rubrum Blume
var. rubrum
Euph. 2 0.05 1.96 3 0.04 1.89 M.NSR.N
H
35 Unidentified Uni-
dent.
2 0.04 1.90 2 0.02 0.63 M.NH
36 Litsea ligustrina
(Nees) Fern.-Vill.
Laur. 1 0.08 1.40 1 0.16 1.20 M.NH
37 Ficus ribes
Reinw. ex Blume
Mor. 1 0.01 0.87 3 0.03 1.82 L.M
38 Urophyllum
arboreum
(Reinw. ex
Blume) Korth.
Rub. 1 0.01 0.87 3 0.03 1.84 M.NSR
GROUP 3
39 Prasoxylon
alliaceum
(Blume) M.
Roem.
Melia. 6 0.25 6.60 1 0.02 0.66 M.NH
40 Polyspora
excelsa (Blume)
Orel, Peter G.
Wilson, Curry &
Luu
Thea. 3 0.06 2.83 2 0.03 1.30 M
41 Barringtonia
racemosa (L.)
Spreng
Lecyth. 3 0.04 2.27 1 0.03 0.71 M.NSR.N
H
42 Machilus rimosa
Blume
Euph. 2 0.05 1.98 3 0.05 2.01 M.NH
43 Lannea
coromandelica
(Houtt.) Merr.
Anac. 1 0.13 1.75 3 0.12 2.36 M.NSR.N
H
44 Cinnamomum sp. Laur. 1 0.11 1.57 2 0.03 1.00 M.NH
45 Litsea noronhae
Blume
Laur. 1 0.02 0.97 1 0.08 0.97 M.NH
GROUP 4
46 Chisocheton
ceramicus Miq.
Melia. 2 0.97 5.05 3 0.07 1.80 M.NSR.N
H
47 Didymocheton
nutans Blume
Melia. 14 0.79 9.94 7 0.42 5.83 M
48 Syzygium
antisepticum
(Blume) Merr. &
L.M.Perry
Myrt. 13 0.37 7.67 31 1.24 20.05 M. NH
49 Symplocos
acuminata
(Blume) Miq.
Sympl. 8 0.18 5.29 9 0.12 5.23 M.NSR.N
H
50 Myrsine hasseltii
Blume ex. Scheff.
Myrs. 7 0.11 4.44 1 0.11 1.13 M.NH
51 Elaeocarpus
angustifolius
Blume
Elaeoc. 2 0.71 4.00 2 0.06 1.48 M.NSR.N
H
SADILI et al.: Variation composition and structure of natural lowland forest
2023] 23
52 Lithocarpus
korthalsii (Endl.)
Soepadmo
Fag. 3 0.52 3.82 1 0.02 0.70 M.NH
53 Knema laurina
(Blume) Warb.
Myrist. 3 0.23 2.35 1 0.03 0.72 L.NSR.N
H
54 Nauclea subdita
(Korth.) Steud.
Rub. 3 0.14 2.28 3 0.30 3.20 M.NSR.N
H
55 Cinnamomum
parthenoxylon
(Jack) Meisn.
Laur. 3 0.11 2.15 2 0.19 2.08 M.NSR.N
H
56 Ostodes
paniculata Blume
Euph. 1 0.08 0.89 1 0.06 0.87 M.NH
57 Ficus tinctoria
subsp. gibbosa
(Blume) Corner
Mor. 1 0.04 0.73 3 0.03 1.57 M.NSR.N
H
58 Ficus fistulosa
Reinw. ex Blume
Mor. 1 0.03 0.70 3 0.03 1.93 L.M
59 Mangifera
laurina Blume
Anac. 1 0.01 0.61 1 0.06 0.90 M.NSR.N
H
60 Mangifera sp. Anac. 1 0.01 0.61 1 0.17 1.40 M.NH
GROUP 5
61 Neonauclea
lanceolata
(Blume) Merr.
Rub. 7 0.32 7.98 M
62 Myrsine affinis
A.DC.
Myrs. 5 0.11 4.39 M
63 Donella
lanceolata
(Blume) Aubrév
Sapot. 1 0.37 3.50 M.NSR.N
H
64 Litsea
brachystachya
(Blumea) Fern.-
Vill.
Laur. 3 0.10 3.16 M.NSR.N
H
65 Liquidambar
excelsa
(Noronha) Oken
Ham. 3 0.07 2.89 M
66 Actinodaphne
glomerata
(Blume) Nees
Laur. 2 0.07 2.10 M.NSR.N
H
67 Semecarpus
heterophylla
Blume
Anac. 2 0.07 1.87 L.NSR.N
H
68 Adina trichotoma
(Zoll. & Moritzi)
Benth. & Hook.f.
ex B.D.Jacks
Rub. 1 0.04 1.52 L.NSR.N
H
69 Litsea glutinosa
(Lour.) C.B.Rob.
Laur. 1 0.04 1.11 M.NSR.N
H
70 Litsea sp. Laur. 1 0.04 0.90 M.NH
71 Gironniera
subaequalis
Planch.
Cann. 1 0.04 0.89 M.NSR.N
H
REINWARDTIA 24 [VOL.22
72 Saurauia
bracteosa DC.
Acthin. 1 0.03 0.89 M.NH
73 Symplocos
fasciculata Zoll.
Sympl. 1 0.01 0.88 M
74 Ficus
grossularioides
Burm.f.
Mor. 1 0.01 0.87 M.NH
GROUP 6
75 Spondias
pinnata (L.f.)
Kurz
Anac. 3 0.98 5.65 M.NSR.N
H
76 Flacourtia sp. Flacour
.
5 0.13 3.38 M.NH
77 Ficus
glandulifera
(Miq.) Wall. ex
King
Mor. 2 0.32 2.45 M.NSR.N
H
78 Sterculia
chrysodasys
Miq.
Ster. 2 0.03 1.26 M.NSR.N
H
79 Diospyros
frutescens
Blume
Ebena. 1 0.09 0.94 L.NGPF
80 Artocarpus
elasticus Reinw.
ex Blume
Mor. 1 0.05 0.94 L.M.NSR.
NH
81 Podocarpus
neriifolius
D.Don
Pod. 1 0.05 0.76 M.NH
82 Magnolia
macklottii
(Korth.) Dandy
Mag. 1 0.04 0.73 M.NSR.N
H
83 Litsea
sphaerocarpa
Blume
Laur. 1 0.02 0.66 M.NSR
84 Leptospermum
polygalifolium
Salisb.
Laur. 1 0.02 0.64 M.NSR.N
H
GROUP 7
85 Calliandra
houstoniana
var. calothyrsus
(Meisn.) Barne-
by
Leg. 28 0.33 14.18 L.A.NH
86 Decaspermum
fruticosum
J.R.Forst. &
G.Forst.
Myrt. 3 0.40 3.72 M.NH
87 Swietenia
mahagoni (L.)
Jacq.
Melia. 4 0.31 3.55 L.A.NH
SADILI et al.: Variation composition and structure of natural lowland forest
2023] 25
88 Falcataria falca-
ta (L.) Greuter &
R.Rankin
Leg. 3 0.35 3.45 L.A.NH
89 Pinus merkusii
Jungh. & de
Vriese
Pin. 4 0.11 2.90 L.A.NH
90 Ficus variegata
Blume
Mor. 4 0.08 2.39 L.N.H
91 Cryptocarya
ferrea Blume
Laur. 3 0.08 2.14 L.N.H
92 Bellucia pentam-
era Naudin
Melast. 3 0.05 1.99 M.A.NH
93 Quercus lineata
Blume
Fag. 2 0.13 1.80 M.NSR.N
H
94 Artocarpus
heterophyllus
Lam.
Mor. 2 0.09 1.62 L.A.NH
95 Garcinia
cambogioides
(Murray)
Headland var.
cambogioides
Clusia. 2 0.06 1.45 M.NSR.N
H
96 Syzygium sp. Myrt. 1 0.15 1.32 M.NH
97 Blumeodendron t
okbrai (Blume)
Kurz
Euph. 1 0.05 0.82 L.M.NSR.
NH
98 Dracaena
angustifolia
(Medik.) Roxb.
Aspar. 1 0.04 0.79 L.A.NH
99 Ficus virens
Aiton var. virens
Mor. 1 0.04 0.78 M.NSR.N
H
100 Magnolia
sumatrana var. gl
auca (Blume)
Figlar & Noot.
Mag. 1 0.03 0.72 M.NH
101 Aporosa
spaeridophora
Merr.
Euph. 1 0.02 0.71 M.NSR.N
H
102 Pavetta montana
Reinw. ex Blume
Rub. 1 0.01 0.65 M.NSR.N
H
103 Homalanthus
populneus (Geisel
er) Kuntze
Euph. 1 0.01 0.64 L.M.NH
104 Elaeocarpus
angustifolius Blu
me
Elaeoc. 1 0.01 0.63 M.NH
105 Sterculia sp. Ster. 1 0.01 0.63 M.NH
106 Magnolia
montana (Blume)
Figlar
Mag. 1 0.01 0.62 L.M.NH
107 Neesia
altissima (Blume)
Blume
Bomb. 1 0.01 0.62 M.NSR
REINWARDTIA 26 [VOL.22