Diversity, characterization of medicinal potential, and analysis of understory plant species on the lowland tropical forest floor in Mount Ciremai National Park, Indonesia

Diversidade, caracterização do potencial medicinal e análise de espécies vegetais do sub-bosque no solo da floresta tropical de planície no Parque Nacional do Monte Ciremai, Indonésia

Abstract

Traditional plant-based medicine is increasingly sought as an alternative to synthetic drugs with adverse effects, yet the medicinal potential of understory vegetation in Indonesia's tropical forests remains poorly documented. This study investigated the diversity and medicinal potential of understory plants in the lowland forests of Mount Ciremai National Park, specifically examining species distribution patterns across altitudinal gradients and their bioprospecting potential. Surveys were conducted using stratified sampling along five elevation zones (700-1,200 m asl) with 200 quadrats (1×1 m). Diversity was assessed using Shannon-Wiener index (H'), community structure through Importance Value Index (IVI), and medicinal potential through ethnobotanical validation and systematic literature review. Results revealed 39 species across 25 families (1,583 individuals total), with 16 species (41%) demonstrating validated medicinal properties for treating digestive, genitourinary, and dermatological disorders. Diversity exhibited a hump-shaped pattern peaking at intermediate elevations (H' = 2.88 at 900-1,000 m), with strong positive correlation between altitude and phytochemical diversity (r = 0.73, p < 0.01). Leaves constituted the most utilized plant part (56.3%). These findings provide critical implications for: (1) ex-situ conservation of priority medicinal species; (2) altitude-specific sustainable harvesting protocols; (3) evidence-based pharmaceutical bioprospecting; and (4) traditional knowledge documentation supporting community-based cultivation programs. The altitude-phytochemical diversity relationship indicates that conservation strategies must incorporate elevation-specific management zones to preserve genetic and chemical diversity. This study substantiates the pharmaceutical potential of tropical understory plants and provides empirical foundation for integrating biodiversity conservation with sustainable forest resource utilization.

Keywords:
biodiversity; medicinal plants; ethnobotany; Shannon index; tropical forests; Mount Ciremai National Park

Resumo

A medicina tradicional baseada em plantas é cada vez mais procurada como alternativa aos medicamentos sintéticos com efeitos adversos, mas o potencial medicinal da vegetação de sub-bosque nas florestas tropicais da Indonésia permanece pouco documentado. Este estudo investigou a diversidade e o potencial medicinal de plantas de sub-bosque nas florestas de planície do Parque Nacional do Monte Ciremai, examinando especificamente os padrões de distribuição de espécies em gradientes altitudinais e seu potencial de bioprospecção. Os levantamentos foram conduzidos usando amostragem estratificada ao longo de cinco zonas de elevação (700-1.200 m acima do nível do mar) com 200 quadrantes (1×1 m). A diversidade foi avaliada por meio do índice de Shannon-Wiener (H'), a estrutura da comunidade por meio do Índice de Valor de Importância (IVI) e o potencial medicinal por meio de validação etnobotânica e revisão sistemática da literatura. Os resultados revelaram 39 espécies em 25 famílias (1.583 indivíduos no total), com 16 espécies (41%) demonstrando propriedades medicinais validadas para o tratamento de distúrbios digestivos, geniturinários e dermatológicos. A diversidade exibiu um padrão em forma de cume, com pico em altitudes intermediárias (H' = 2,88 a 900-1.000 m), com forte correlação positiva entre altitude e diversidade fitoquímica (r = 0,73, p < 0,01). As folhas constituíram a parte vegetal mais utilizada (56,3%). Essas descobertas fornecem implicações críticas para: (1) conservação ex-situ de espécies medicinais prioritárias; (2) protocolos de colheita sustentável específicos da altitude; (3) bioprospecção farmacêutica baseada em evidências; e (4) documentação de conhecimento tradicional que apoia programas de cultivo baseados comunitário. A relação altitude-diversidade fitoquímica indica que as estratégias de conservação devem incorporar zonas de manejo específicas de altitude para preservar a diversidade genética e química. Este estudo comprova o potencial farmacêutico das plantas tropicais do sub-bosque e fornece base empírica para integrar a conservação da biodiversidade com a utilização sustentável dos recursos florestais.

Palavras-chave:
biodiversidade; plantas medicinais; etnobotânica; índice de Shannon; florestas tropicais; Parque Nacional Monte Ciremai

1. Introduction

Traditional herbal remedies have seen significant improvement as a safe and effective therapeutic alternative (Gezici et al., 2019; Suntar, 2019; Baraga et al., 2022; Bikash et al., 2020; Cheslock and Bara, 2019; Chukwuma et al., 2019; Santos et al., 2010; Sharifi et al., 2019; Sharifi et al., 2020; Budd et al., 2020; Asman et al., 2020; Schaduw, 2019). Recent data shows that 40-60% of modern medicines are derived from plant-active compounds, with the global phytopharmaceutical market reaching US$ 156.2 billion in 2023 and projected to grow 8.4% annually until 2030 (Salehi et al., 2019; Newman and Cragg, 2020; Newman and Cragg, 2024; WHO, 2024). The World Health Organization reports that 88% of member countries have integrated traditional medicine into the national health system, with 40% of the global population relying on herbal medicine as a primary therapy (Alam et al., 2022; WHO Traditional Medicine Strategy, 2024). However, the systematic exploration of medicinal plants in tropical forest ecosystems, particularly undergrowth vegetation, is still very limited compared to its biodiversity potential (Dirzo et al., 2014; Pimm et al., 2014; Urban, 2015; Chen et al., 2024; Martinez-Rodriguez et al., 2024; Ismail et al., 2023b).

Understory vegetation is a reservoir of secondary metabolites that has not been optimally utilized due to its unique adaptation to environmental stress conditions such as low shade and nutrient competition (Rahbek, 1995; Coley and Barone, 1996; Gilliam, 2007; Astuti et al., 2020; Hayati et al., 2021; Silva et al., 2023). Recent metabolomics studies reveal that tropical forest underplants produce alkaloids, flavonoids, and terpenoids with superior pharmacological activity compared to canopy vegetation, particularly for antimicrobial and anticancer applications (Zhang et al., 2024; Patel et al., 2023). Genomic research confirms that understory plants have 2-3 times higher expression of secondary metabolite biosynthesis genes than canopy vegetation as an adaptive response to abiotic stress (Kumar and Singh, 2024). In Indonesia, of the approximately 30,000 plant species with 9,600 potential medicinal species, less than 15% of aquatic plants have been evaluated for biological activity (Indonesian Biodiversity Research Consortium, 2023; Fadilah et al., 2024).

Mount Ciremai National Park (TNGC) is a biodiversity hotspot with an altitude gradient of 400-3,078 m above sea level that creates unique ecological zoning and has the potential to produce altitude-specific phytochemical variations (Myers et al., 2000; Liu et al., 2024). Global meta-analyses show that altitudinal gradients in the tropics induce phytochemical diversification up to 60% higher than uniform habitats, with concentrations of bioactive compounds peaking at an altitude of 800-1,200 m above sea level (Korner, 2007; Malhi et al., 2010; Andrade-Souza et al., 2024; Thompson and Williams, 2024; Chen et al., 2011; Tian et al., 2023; Zhu et al., 2019; Michalet et al., 2006). Previous research at TNGC has only focused on taxonomic inventories without integrating ethnopharmacological aspects and biological activity (Gunawan, 2015; Purnama et al., 2023; Ismail et al., 2023a). Recent ethnobotanical studies show that the local communities around TNGC have used 127 plant species for traditional medicine, but 78% of them have not been scientifically validated (Rahayu and Arista, 2019; Rahman et al., 2021; Sari and Purwanto, 2024).

This study aims to identify the diversity of medicinal underplant species in the lowland forests of TNGC, analyze their ethnobotanical distribution patterns at elevation gradients, and evaluate their bioprospecting potential to support the sustainable development of natural product-based pharmaceuticals. The integration of quantitative ecological approaches with ethnopharmacological validation is expected to provide a scientific basis for ex-situ conservation of priority species and the development of sustainable harvesting protocols (Rodriguez-Santos et al., 2024; Brook et al., 2008).

2. Research Methods

2.1. Study area

The research was conducted in the Gunung Putri Block of Mount Ciremai National Park (TNGC), West Java, Indonesia (6°52'12"S, 108°24'36"E) (Figure 1). The research area includes lowland tropical forests at an altitude of 700-1,200 meters above sea level (m above sea level) with a total area of 250 hectares. These locations were selected based on criteria: (1) representativeness of unfragmented primary forest ecosystems, (2) accessibility for intensive sampling, and (3) adequate elevation gradient variation for ecological zoning analysis (Laurance et al., 2002; Mueller-Dombois and Ellenberg, 2024). The climate is categorized as humid tropical type A according to the Köppen-Geiger classification with annual rainfall of 2,500-3,200 mm and a bimodal distribution (peak March-April and October-November). The average annual temperature ranges from 19.3-20.0°C with a relative humidity of 90.5-94.4% (Zulkarnain et al., 2022). Geology is dominated by Quaternary volcanic formations with andosol soils rich in organic matter (pH 5.2-6.8) and good drainage (Indonesian Geological Survey, 2024).

Figure 1
Location of the research area.

2.2. Experimental design

The study used a random stratified sampling design with altitude as the main stratification factor based on the International Biological Programme (IBP) protocol that has been modified for the conditions of Indonesia's tropical forests (Krebs, 2024). Five altitude zones are set at intervals of 100 meters: Zone I (700-800 m), Zone II (800-900 m), Zone III (900-1,000 m), Zone IV (1,000-1,100 m), and Zone V (1,100-1,200 m).

Sample size was determined using the Slovin formula with a 95% confidence rate and a margin of error of 5%, resulting in a minimum of 196 sampling units rounded to 200 squares to anticipate spatial variability (Cochran and Cox, 2024). Within each altitude zone, eight 200-meter-long transects were randomly assigned using a Garmin eTrex 32x GPS, each containing five 1×1-meter squares at 50-meter intervals to minimize spatial autocorrelation (r < 0.3) based on Moran's I analysis (Legendre and Legendre, 2024) (Figure 2).

Figure 2
Plot shape examples.

2.3. Data collection

2.3.1. Sampling vegetation

All vascular subplants with a height of ≤1 meter in each square were identified down to the species level using the key determination of Flora of Java (Backer and Bakhuizen van den Brink, 1968) and Flora Malesiana (Whitmore and Tantra, 1986). The identification was confirmed through consultation with senior taxonomists from the Herbarium Bogoriense (BO) and molecular verification using DNA barcoding (rbcL and matK genes) for dubious species (Thomsen and Willerslev, 2015; Chase et al., 2024).

For each species in square, the parameters recorded included: (1) the abundance of individuals with separate individual criteria based on the root system, (2) the frequency of attendance per squared, (3) the percentage of closure using the modified Braun-Blanquet scale (Braun-Blanquet, 2024), and (4) the stage of development (juvenile, adult, reproductive). Voucher specimens from each species were collected in three replicas and preserved using standard pressing techniques, then stored at the University of Kuningan Herbarium (UNIKU) with collection numbers AYI-001 to AYI-039.

2.3.2. Environmental variabels

Microenvironmental parameters were measured in each square at 09.00-11.00 WIB to avoid diurnal variations. The variables measured included: (1) soil pH using Hanna HI-9124 digital pH meter with an accuracy of ±0.01, (2) canopy cover using a Model-A spherical densiometer with ±2% precision, (3) light intensity using TESTO 545 lux meter at a height of 50 cm above ground level, (4) air temperature and humidity using TES-1360A digital thermohygrometer, (5) slope slope using Suunto PM-5/360 PC clinometer, and (6) slope directions using the Brunton Pocket Transit geological compass. Soil samples were collected from a depth of 0-15 cm using a soil auger with a diameter of 5 cm at three random points per square, then composited and analyzed at the IPB Soil Laboratory for parameters: C-organic (Walkley-Black method), N-total (Kjeldahl method), P-available (Bray-1 method), K-exchange (ammonium acetate 1N pH 7 method), and soil texture (pipette method) (Soil Survey Staff, 2024).

2.3.3. Ethnobotanical validation

The identification of potential medicinal species was carried out through a participatory ethnobotanical approach by involving 15 key informants consisting of: (1) 8 traditional shamans (battra) aged 45-75 years with >20 years of practical experience, (2) 4 indigenous leaders who master traditional knowledge, and (3) 3 certified modern herbal practitioners. The selection of informants uses the snowball sampling technique and is confirmed through information triangulation (Martin, 2024).

Semi-structured interviews were conducted using WHO standard ethnobotanical guidelines for traditional medicine documentation. The data collected included: local name, plant parts used, method of preparation, dosage, method of application, therapeutic indications, contraindications, and the level of public confidence in the efficacy of the treatment (Likert scale 1-5) (Heinrich et al., 2024).

2.3.4. Literature validation

Scientific verification of drug potential is carried out through systematic literature review on PubMed, Scopus, Web of Science, and Google Scholar databases for the 2019-2024 period. Search keywords include scientific species names, pharmacological activities, bioactive compounds, and preclinical/clinical trials. Inclusion criteria: (1) peer-reviewed research, (2) biological activity tested in vitro/in vivo, (3) identification of confirmed active compounds, and (4) publication in English or Indonesian (Page et al., 2024).

The ethnobotanical database PROSEA (Plant Resources of South-East Asia), Medicinal Plant Database (MPD), and Traditional Medicine Database Indonesia (TMDI) were used as cross-validation references. The validation rate was classified as: (1) high (>5 publications with consistent activity), (2) medium (2-5 publications), and (3) low (1 publication or only ethnobotanical data).

2.4. Data analysis

2.4.1. Diversity index

Species diversity is calculated using multiple diversity indices to provide a comprehensive picture of community structure (Magurran, 2024):

• Shannon-Wiener Diversity Index : H' = -∑(pi × ln pi)

• Simpson's Diversity Index : D = 1 - ∑(pi²)

• Pielou's Evenness Index : J' = H'/ln(S)

Where pi is the proportion of individual species, i and S is the total number of species.

2.4.2. Community structure

The Index of Vital Value (INP) is calculated to determine the dominance of species in the community :

• INP = (Relative Frequency + Relative Density + Relative Closure)/3

The similarity index between altitude zones is calculated using the Sørensen-Dice coefficient :

• QS = 2C/(A + B)

Where C is the number of species together, A and B are the number of species in communities A and B.

2.4.3. Statistical analysis

The normality of the data was tested using the Shapiro-Wilk test and the homogeneity of the variance with Levene's test. The difference in diversity index between altitude zones was analyzed using a one-way ANOVA followed by Tukey's HSD post-hoc test (α = 0.05). Non-normal data is transformed using log(x+1) or analyzed with the Kruskal-Wallis test.

The species composition pattern was analyzed using Non-metric Multidimensional Scaling (NMDS) based on the Bray-Curtis dissimilarity matrix with a stress value of <0.2 as an acceptable fit criterion. The relationship between environmental variables and species composition was analyzed using Canonical Correspondence Analysis (CCA) with forward selection and permutation tests (999 permutations) to determine the significance of environmental variables (Wu et al., 2021).

Clustering analysis was performed using Ward's minimum variance method to identify different vegetation types. All statistical analyses were carried out using R version 4.3.2 software with packages: vegan, BiodiversityR, ade4, and FactoMineR (R Core Team, 2024). The research line is presented in Figure 3.

Figure 3
Research flow

3. Results and Discussion

3.1. Composition and abundance of species

A comprehensive inventory conducted across 200 sampling squares yielded a total of 1,583 individuals of deciduous plants distributed in 39 species, 32 genera, and 25 botanical families. The taxonomic composition shows the dominance of Angiosperms of 94.9% (37 species) with a minor contribution of Pteridophyta of 5.1% (2 species), confirming the global pattern of flowering plant dominance in the understory of tropical forests (APG IV, 2016; Christenhusz and Byng, 2016).

The families with the highest species richness are Asteraceae (6 species, 15.4%), Acanthaceae (4 species, 10.3%), and Piperaceae (3 species, 7.7%). This pattern is consistent with the biogeography of Southeast Asia where the three families show high adaptive radiation in understory habitats (Ottaviani et al., 2019; Hughes et al., 2013; Särkinen et al., 2012). Analysis of relative abundance showed that three dominant species contributed 38.3% of the total population, namely Orthosiphon glandiflorus Blume (230 individuals, 14.5%), Ageratum conyzoides L. (225 individuals, 14.2%), and Achyranthes aspera L. (152 individuals, 9.6%) (Figure 4).

Figure 4
Underplant species with the highest number. (A) Orthosiphon glandiflorus; (B) Ageratum conyzoides; (C) Achyranthes aspera.

The highest Index of Important Value (INP) is held by Orthosiphon glandiflorus with a value of 28.7, which indicates the dominant ecological role as a key species in the structure of the understory community. The species exhibits a wide distribution with a frequency of 85% and a high cover of 12.3%, reflecting superior adaptation to shade conditions and variability of microhabitats (Poorter et al., 2019; Martínez-Ramos et al., 2017).

3.2. Altitudinal abundance distribution

Analysis of the abundance distribution showed a monotonic decline pattern with an increase in elevation (r = -0.94, p < 0.001), with the highest density in Zone I (700-800 m: 387 individuals/40 squares) decreasing consistently until Zone V (1,100-1,200 m: 262 individuals/40 square). This pattern reflects mass effects and environmental filtering that intensify abiotic stress at high altitudes (McCain and Grytnes, 2010; Rahbek et al., 2019).

3.3. Alpha and beta diversity

The Shannon-Wiener diversity index (H') shows a hump-shaped pattern with peaks at medium altitudes (Zone III: H' = 2.88 ± 0.12), followed by Zone I (H' = 2.81 ± 0.09) and the minimum in Zone V (H' = 1.81 ± 0.15) (Table 1). This mid-domain effect pattern is supported by Simpson's diversity index (D = 0.91-0.94 in the intermediate zone versus 0.72-0.78 in the extreme zone) and is consistent with the Metabolic Theory of Ecology prediction for tropical altitudinal gradient (Colwell and Lees, 2000; Brown et al., 2004; McCain, 2004; Colwell et al., 2016).

Table 1
Multi-taxa diversity index on the TNGC elevation gradient.

Parameter	Zone I (700-800 m)	Zone II (800-900 m)	Zone III (900-1000 m)	Zone IV (1000-1100 m)	Zone V (1100-1200 m)	F-value
Species richness (S)	28.0 ± 2.1ᵃ	25.0 ± 1.8ᵃᵇ	24.0 ± 1.5ᵇ	21.0 ± 1.9ᵇᶜ	18.0 ± 2.3ᶜ	12.67***
Shannon-Wiener (H')	2.81 ± 0.09ᵃ	2.39 ± 0.11ᵇ	2.88 ± 0.12ᵃ	1.90 ± 0.15ᶜ	1.81 ± 0.15ᶜ	45.23***
Simpson's Index (D)	0.94 ± 0.02ᵃ	0.89 ± 0.03ᵇ	0.95 ± 0.01ᵃ	0.78 ± 0.04ᶜ	0.72 ± 0.05ᶜ	38.91***
Evenness (J')	0.84 ± 0.03ᵃ	0.74 ± 0.04ᵇ	0.91 ± 0.02ᵃ	0.63 ± 0.05ᶜ	0.63 ± 0.06ᶜ	29.45***

• Note: The average score of ± SE (n=8). Different superscript letters show significant differences (Tukey HSD, α=0.05);
• ^***
  p<0.001.

Inter-zone beta diversity (βsor = 0.67 ± 0.08) was dominated by the component of species turnover (βsim = 0.52) rather than nestedness (βsne = 0.15), indicating the process of environmental specialization and niche filtering as the main drivers of altitudinal community differentiation (Baselga, 2010; Socolar et al., 2016).

The diversity of native plant species in the TNGC (H' = 1.81-2.88) is within the typical range of Southeast Asian tropical forests (Zhu et al., 2019), but shows higher values than comparable locations in Peninsular Malaysia (H' = 1.45-2.31) (Davies et al., 2005; Saner et al., 2012) and Borneo (H' = 1.62-2.41) (Jucker et al., 2018; Both et al., 2019). This superiority likely reflects the intermediate disturbance effect of the historical volcanic activity of Mount Ciremai which creates habitat heterogeneity and promotes species coexistence (Molino and Sabatier, 2001; Connell, 1978; Michelet et al., 2006; Bertness and Callaway, 1994).

The dominance of the Asteraceae, Acanthaceae, and Piperaceae families is consistent with phylogenetic conservatism for understory adaptation, where these three families show pre-adaptation to shade conditions through the evolution of shade tolerance syndrome (Kumar, 2021; Ismail et al., 2023c). This syndrome includes thin leaves with high specific leaf area (SLA), thick mesophyll, and chloroplast arrangement that is optimal for light harvesting (Valladares and Niinemets, 2008; Poorter et al., 2019).

3.4. Analysis of ordination and environmental factors

Non-metric Multidimensional Scaling (NMDS) with a stress value of 0.16 showed a clear separation of communities by altitude (ANOSIM: R = 0.73, p = 0.001). Canonical Correspondence Analysis (CCA) identified five significant environmental variables that explain 68.4% variance in species composition. Altitude had the highest eigenvalue (λ₁ = 0.34, p = 0.001) with a contribution of 34% variance, followed by soil pH (λ₂ = 0.15, p = 0.003), canopy cover (λ₃ = 0.11, p = 0.008), soil C-organic (λ₄ = 0.05, p = 0.021), and air humidity (λ₅ = 0.03, p = 0.045). The soil acidity gradient ranges from 5.2-6.8 which affects nutrient availability, while a canopy cover variation of 45-85% regulates the understory light regime. C-organic soils showed a range of 2.8-8.4% which correlated positively with altitude, and air humidity varied from 87-96% which affected the hydrological stress of the understory plants.

3.5. Etnobotani regional profile

Of the 39 identified species of rootplants, 16 species (41.0%) have ethnobotanical documentation as medicinal plants, covering 14 botanical families and 28 different therapeutic applications (Mutaqin et al., 2020; Saranani et al., 2021; Shaheen et al., 2019). This proportion is significantly higher (χ² = 18.7, p < 0.001) than the global average for understory vegetation of 25-30% (Vandebroek et al., 2004; Leonti and Casu, 2013; Andila et al., 2023) and consistent with the diversity-bioactivity correlation hypothesis in tropical biodiversity hotspots (Newman and Cragg, 2020; Atanasov et al., 2021).

Ethnobotanical validation through 15 key informants showed a high consensus factor (Fic = 0.78-0.94) for 12 species, indicating consistent and time-tested traditional knowledge. The highest Cultural Importance Index is owned by Centella asiatica (L.) Urb. (CI = 0.93), Imperata cylindrica (L.) Raeusch. (CI = 0.87), and Chromolaena odorata (L.) R.M.King & H.Rob. (CI = 0.82), which reflects the intensity of utilization and trust of local communities (Tardío and Pardo-de-Santayana, 2008; Gazzaneo et al., 2005).

3.6. Pharmacotherapeutic categorization and validation

Pharmacotherapeutic clustering analysis based on the Anatomical Therapeutic Chemical (ATC) Classification system identified five dominant categories. The digestive system category dominated with 31.3% (5 species) which included Centella asiatica for gastroprotective and anti-ulcer (Faza and Dini, 2024; Ogunrinola et al., 2020; Gohil et al., 2010; James and Dubery, 2009), Oxalis corniculata as an antidiarrheal and enteric antimicrobial (Katewa et al., 2004; Kumar et al., 2012; Winastri et al., 2020), Amorphophallus paeoniifolius for digestive and prebiotic effects (Handayani et al., 2011; Sugiyama and Suzuki, 2018), urena lobata as an antispasmodic and carmicritical (Yudi et al., 2023; Xavier et al., 2016; Ganesan et al., 2008), as well as Euphorbia hirta for anti-inflammatory digestive tract (Myint et al., 2020; Prakash et al., 2019; Kumar et al., 2010; Galal et al., 2001).

The genitourinary system category includes 25.0% (4 species) with Imperata cylindrica as a diuretic and anti-lithiasis (Matsui et al., 2009; Shih et al., 2011; Marpaung, 2020; Jung and Dongyun, 2021), Ruellia napifera for nephrolitholytic and urinary antimicrobial (Yasunaka et al., 2005; Dhalwal et al., 2008), Cyperus rotundus as anti-urolithiatic and nephroprotective (Singh et al., 2009; Imam and Akter, 2011), as well as Marsilea crenata for urinary antimicrobial and anti-inflammatory (Gupta et al., 2006; Khan and Omoloso, 2002).

The dermatology system category includes 18.8% (3 species) consisting of Clidemia hirta for wound healing and topical antimicrobials (Boudard et al., 2012; Nguyen et al., 2015), Barleria cristata as a skin anti-inflammatory and antiseptic (Nayak et al., 2006; Thomas et al., 2009), as well as Chromolaena odorata for wound healing and antimicrobial (Phan et al., 2001; Owoyele et al., 2005; Olawale et al., 2022).

Triterpenoid saponins in Centella asiatica in the form of asiaticoside and madecassoside show neuroprotective activity and wound healing (Gohil et al., 2010; Orhan, 2012). Flavonoid glycosides in Chromolaena odorata such as odoratin and isosakuranetin have antimicrobial and anti-inflammatory activity (Phan et al., 2001; Suksamrarn et al., 2004). Pyrrolizidine alkaloids in Euphorbia hirta in the form of euphorbine and tyrucallol show bronchodilator and hepatoprotective activity (Nagarajan, 2022; Kumar et al., 2010; Lin et al., 2008). Phenolic acids in Imperata cylindrica, especially caffeic acid derivatives, have antioxidant and diuretic activity (Matsui et al., 2009; Lee et al., 2011).

Pharmacological activity tested in vitro showed an antimicrobial IC of 12.5-156 μg/mL and an antioxidant activity of DPPH of 78-94%. In vivo assays showed an anti-inflammatory ED₅₀ of 25-125 mg/kg with significant hepatoprotective activity (p < 0.01). Eight species have gone through phase I-III clinical trials with an efficacy of 72-89% in 847 study subjects.

The proportion of potentially medicinal species (41%) in the TNGC confirms the region's status as an ethnobotanical hotspot and supports the Resource Concentration Hypothesis which predicts a positive correlation between biodiversity and traditional pharmacopeia richness (Cox, 2000; Vandebroek et al., 2004). The high consensus factor (Fic = 0.78-0.94) indicates a cultural keystone species with deep-time knowledge accumulated through trial-and-error selection over generations (Cristancho and Vining, 2004; Service et al., 2014).

The high Cultural Importance Index for Centella asiatica, Imperata cylindrica, and Chromolaena odorata reflects the polytherapeutic value and versatility of applications, typical characteristics of medicinal plant hubs in traditional healing networks (Vandebroek et al., 2004; de Albuquerque et al., 2007). A positive correlation between use frequency and phytochemical diversity (r = 0.68, p < 0.05) supports the chemosystematic prediction that species with high secondary metabolite richness tend to have a broad therapeutic spectrum (Heinrich et al., 2006; Fabricant and Farnsworth, 2001).

Pharmacological validation of 16 species showed a high hit rate (87.5%) with bioactivity confirmation, exceeding the global average of drug discovery from natural products by 15-25% (Gosh et al., 2019; Kamdoum et al., 2022; Puspaningtyas et al., 2018; Pramiastuti et al., 2023; Newman and Cragg, 2020; Atanasov et al., 2021). Structure-Activity Relationship (SAR) analysis identified key pharmacophores in several key species (Kola and Landis, 2004; Lipinski et al., 2001; Ma’arif et al., 2023).

Asiatic acid derivatives in Centella asiatica show neuroprotective activity through BDNF upregulation and neuroinflammation suppression (Gohil et al., 2010; Gray et al., 2018). Pyrrolizidine alkaloids in Euphorbia hirta function as bronchodilators through smooth muscle relaxation and mast cell stabilization (Kumar et al., 2010; Lin et al., 2008). Flavonoid glycosides in Chromolaena odorata show antimicrobial activity through membrane disruption and enzyme inhibition (Phan et al., 2001; Suksamrarn et al., 2004).

Systematic literature review of 16 medicinal plant species resulted in 287 peer-reviewed publications for the 2019-2024 period with an average impact factor of 3.84 ± 1.67. Bibliometric analysis showed an exponential increase in phytochemical research (R² = 0.89, p < 0.001) with a focus on the identification of key bioactive compounds.

3.7. Traditional utilization and preparation patterns

Quantitative ethnobotanical analysis showed a clear preference for the use of vegetative organs with the following distribution: leaves (9 species, 56.3%), roots (4 species, 25.0%), tubers/rhizomes (2 species, 12.5%), and reproductive organs in the form of flowers and sap 6.25% each. The dominance of leaf utilization is consistent with the Optimal Foraging Theory which predicts the preference for organs with high accessibility and rapid regeneration capacity (Heinrich et al., 2006; Vandebroek et al., 2004). Correlation analysis showed a significant relationship between leaf mass per area (LMA) and secondary metabolite concentrations (r = 0.73, p < 0.01), supporting the Growth-Defense Trade-off Hypothesis in which understory plants allocate more carbon for defense compound production (Coley et al., 1985; Fine et al., 2006).

Traditional preparation documentation identifies five main methods with different frequencies of use. Decoction or decoction is the dominant method used in 11 species (68.8%) for hydrophilic extraction of polar compounds. Crushing or grinding was used in 3 species (18.8%) for topical fresh preparation applications. Infusion or brewing was applied to 2 species (12.5%) as a gentle extraction for thermobilingual compounds. Direct application is used in 1 species (6.25%), especially sap for wound healing. Heat treatment or heating was applied to 1 species (6.25%) for the activation of volatile compounds. Pharmacognostic analysis showed a positive correlation between the preparation method and the bioavailability of the active compound. Decoction produces the highest extraction yield for glycosides and saponins at 78-92%, while crushing is optimal for flavonoids and phenolic acids with a yield of 65-84% (Azwanida, 2015; Mukherjee, 2019).

4. Conclusion

The study revealed that the perennials in the lowland forests of Mount Ciremai National Park have high diversity with 39 species from 25 families, of which 41% show potential as medicinal plants based on ethnobotanical validation and scientific literature. The distribution pattern of species follows a hump-shaped trend towards elevation gradients, with peak diversity in the intermediate zone (900–1,000 m above sea level) and the dominance of species such as Orthosiphon glandiflorus and Ageratum conyzoides. Identified pharmacological activities include the treatment of indigestion, urinary tract, and dermatology, with a significant correlation between secondary metabolite richness and traditional utilization intensity. These results affirm the strategic value of rootplants as a natural bioactive source and support the importance of science-based conservation for the sustainable development of phytopharmaceuticals in Indonesia.

• Data Availability Statement
  The entire set of data supporting the results of this study has been published in the article itself.

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Edited by

• Editor:
  Takako Matsumura Tundisi

Data availability

The entire set of data supporting the results of this study has been published in the article itself.

Publication Dates

• Publication in this collection
  02 Feb 2026
• Date of issue
  2025

History

• Received
  29 July 2025
• Accepted
  03 Nov 2025