Journal of Threatened Taxa | www.threatenedtaxa.org | 26 December 2023 | 15(12): 24409–24419

ISSN 0974-7907 (Online) | ISSN 0974-7893 (Print)

https://doi.org/10.11609/jott.8696.15.12.24409-24419

#8696 | Received 18 August 2023 | Final received 21 November 2023 | Finally accepted 11 December 2023

Comparative morphological and ethnobotanical assessment of certain taxa of genus Glochidion (Phyllanthaceae) from Assam, India

Priyanka Brahma ¹ & Sanjib Baruah ²

^1,2 Department of Botany, Bodoland University, Kokrajhar, Assam 783370, India.

¹ priyabrahma659@gmail.com (corresponding author), ² sanjibbaruah9@gmail.com

Editor: Kannan C.S. Warrier, ICFRE - Institute of Forest Genetics and Tree Breeding, Coimbatore, India. Date of publication: 26 December 2023 (online & print)

Citation: Brahma, P. & S. Baruah (2023). Comparative morphological and ethnobotanical assessment of certain taxa of genus Glochidion (Phyllanthaceae) from Assam, India. Journal of Threatened Taxa 15(12): 24409–24419. https://doi.org/10.11609/jott.8696.15.12.24409-24419

Copyright: © Brahma & Baruah 2023. Creative Commons Attribution 4.0 International License. JoTT allows unrestricted use, reproduction, and distribution of this article in any medium by providing adequate credit to the author(s) and the source of publication.

Funding: None.

Competing interests: The authors declare no competing interests.

Author details: Dr. S. Baruah is working as an assistant professor at the Department of Botany, Bodoland University, Kokrajhar, Assam. SB has 15 years of research and teaching experience on the plant taxonomy and conservation of threatened plants in north-east India. Priyanka Brahma is a bona fide Ph.D. research scholar in the Department of Botany at Bodoland University. PB is pursuing her PhD on the taxonomy and phytochemistry of Glochidion in Assam.

Author contributions: PB has collected, done photography, identified, carried out the morphological analysis of the specimen, performed the PCA and cluster analysis and drafted the manuscript. SB contributed to the present study’s design, supervised the work and revised the manuscript. The final manuscript was examined and approved by both the authors.

Acknowledgements: The authors are grateful to the Assam State Biodiversity Board (ASBB) and PCCF (Chief Wildlife Warden), Panjabari, Assam for granting permission to collect the specimen in protected areas of Assam. The authors are thankful to the authorities of BSI, Shillong, Meghalaya for providing the accession number of the deposited specimen. We also acknowledge the local villagers and traditional healers for sharing their knowledge on the collected specimens. The first author is thankful to the University Grants Commission for the National Fellowship and Scholarship for Higher Education of ST Students (NFST) scheme (Award No. 202021-NFST-ASS-01128) and the Government of India, Ministry of Tribal Affairs, Scholarship Division for financial assistance with her Ph.D.

Abstract: The genus Glochidion, a member of the family Phyllanthaceae, primarily comprises shrub or tree species. It stands out from other genera due to distinctive reproductive features, including prolonged styles in flowers and fruits, apiculate anthers, and lobed and unlobed capsules. This study aimed to compare the morphological characteristics of nine taxa in Assam to facilitate identification and assess ethnobotanical knowledge. Ethnobotanical information was gathered by interviewing the local community, and a taxonomic key was provided for accurate identification. Morphological data underwent principal component analysis (PCA) and cluster analysis using PAST for validation. The comparison revealed distinct characteristics in both vegetative and reproductive traits among Glochidion members. Reproductive features, such as inflorescence, style, ovary, and capsules, were key factors for differentiation and identification. PCA and cluster analysis demonstrated correlation and variation among the taxa, contributing significantly to their demarcation. Ethnobotanical studies indicated the genus’s potential medicinal properties, supported by both primary and secondary information.

Keywords: Angiosperms, cluster analysis, ethnobotany, PCA, Phyllanthoideae, taxonomy, UPGMA.

INTRODUCTION

Northeastern India, including Assam, has a wide variety of vegetation due to its topographic and climatic diversity. It is one of the most diverse regions in the world (Dutta & Dutta 2005; Mao & Roy 2016; Bhattacharyya et al. 2020). The people of Assam have employed a number of plants to treat a wide range of ailments since ancient times (Kanjilal et al. 1940; Bhattacharya et al. 1991). There are many medicinal plants in the area that are well-known to ethnic communities, and Assam is regarded as one of the ecological hot spots in the world (Myers et al. 2000; Asati & Yadav 2004; Saikia et al. 2006).

The species of Glochidion J.R.Forst. & G.Forst. have been used by local people in different places in the world and have immense value in the field of medicine (Lai et al. 2004; Xiao et al. 2008; Bajpai et al. 2016; Chakrabarty & Balakrishnan 2018). Some important biological and pharmacological activities, including the anticancer, antioxidant, and antimicrobial activities of a few members of Glochidion have been reported by many workers (Azam et al. 2012; Rathod & Rajurkar 2017). The people of India mainly rely on medicinal plants and are well-known for ethnobotanical knowledge (Maikhuri & Gangwar 1993; Prakash et al. 2008). Therefore, it was felt worth exploring the genus Glochidion in Assam for its current taxonomy and to assess its ethnomedicinal uses.

The genus Glochidion J.R.Forst. & G.Forst. is a member of the family Phyllanthaceae which is native to northern Australia, Polynesia, southern Asia, and tropical Asia (Chakrabarty & Balakrishnan 2018). The members of the genus are either shrubs or trees, monoecious, pubescent, or glabrous with drooping branches. They are mostly found in evergreen, moist deciduous, tropical, primary and secondary forests, sal forests, hilly areas, and some swampy areas. There are over 320 species worldwide; about 22 species, and eight varieties in India (Balakrishnan & Chakrabarty 2007; Balakrishnan et al. 2012; Chakrabarty & Balakrishnan 2018; Brahma & Baruah 2023). Kanjilal et al. (1940) designated 16 species from erstwhile Assam in ‘Flora of Assam’. At present 12 species and four varieties of the genus are found in Assam (Chakrabarty & Balakrishnan 2018). Traditionally, Glochidion was placed in Euphorbiaceae (Bentham & Hooker 1862–1883; Hutchinson 1973). Later, Hoffmann et al. (2006) discovered that the genus Glochidion sensu lato includes Breynia J.R.Frost & G.Forst., Flueggea Willd. and Margaritaria L.f., which are all allied to Phyllanthus as members of the tribe Phyllantheae and, therefore, belong to the segregate family Phyllanthaceae, and this was later ascertained by Chase et al. (2016) on the molecular basis. The absence of latex and the bi-ovulate ovary distinguish the family Phyllanthaceae from Euphorbiaceae (Chakrabarty & Balakrishnan 2018).

Earlier in some floras, G. ellipticum used to be referred to as G. assamicum, a synonym of G. ellipticum; G. velutinum, i.e., synonym of G. heyneanum; G. hirsutum or G. tomentosum, i.e., synonym of G. zeylanicum var. tomentosum; and G. arborescens, i.e., synonym of G. zeylanicum var. arborescens (Hooker 1890; Kanjilal et al. 1940). According to recent literature and taxonomy data, the taxa G. ellipticum, G. heyneanum, G. zeylanicum var. tomentosum, G. zeylanicum var. arborescens are the accepted names (Chakrabarty & Balakrishnan 2018; WFO 2023).

The present study aimed to resolve the taxonomic identity of certain members of the genus Glochidion based on their morphology. The principal component analysis (PCA) and cluster analysis were carried out to authenticate the morphological evaluation of the taxa studied. All the relevant ethnobotanical information about the Glochidion taxa collected from Assam was documented based on primary sources that could yield potential information in the field of medical research.

MATERIALS AND METHODS

Sample collection, Identification, and Ethnobotanical assessment

Field surveys were conducted in the diverse forest areas in Assam from December 2019 to January 2023. Before conducting the field survey, approval was taken from Assam State Biodiversity Board (ASBB) and PCCF Wildlife Warden, Panjabari, Assam. Glochidion specimens were collected randomly from various locations in Assam, India (Table 1). During the field, collected specimens were taken in an airtight poly bag for further morphological examination and photographs of the specimens were taken using a Realme XT 64 MP mobile camera phone. Garmin GPS etrex 10 was used to record and identify precise latitudes and longitudes of the area of the specimen. In the lab, both vegetative and reproductive characteristics of freshly collected specimens were examined carefully under a Biocraft 20X simple microscope and a Leica EZ4W stereo microscope.

After a critical analysis of the character, identification was made with the help of some authentic literature (Hooker 1890; Kanjilal et al. 1940; Borthakur et al. 2018; Chakrabarty & Balakrishnan 2018), online taxonomic databases (e-Floras 2008; The Plant List 2013; GBIF 2021; POWO 2023) and also with the help of India herbaria (ASSAM, ARUN, and CAL) and digital herbaria (A, MO, NY) (acronyms following Thiers 2018). After reviewing pertinent literature, the threat status of the collected plant taxa was also determined (IUCN 2022). During identification, accepted scientific names and the synonyms of the collected taxa were also checked and confirmed through online databases such as IPNI, POWO, and The WFO Plant List (IPNI 2023; POWO 2023; WFO 2023). The dominant characters that played a key role in the identification of the specimen were their reproductive characters. The list of the collected specimens with their locality, accession number, GPS coordinates, and distribution map were procured (Table 1; Figure 1). The distribution map was created with QGIS 3.26.3 version software.

Ethnobotanical information of all collected taxa was made by the scrutiny of literature as well as communication with some local people and traditional healers in the study area. In addition to documenting the traditional uses and parts utilized for the specimen, we recorded their vernacular names, mode of preparation, application, and route of administration, as outlined in Table 5.

Herbarium preparation and deposition

Herbarium preparations adhered to the established techniques outlined by Jain & Rao (1977), while poisoning procedures followed the methods specified by Clark (1986). Authenticated and verified herbarium specimens for each collected taxon were deposited at the Botanical Survey of India (BSI) in Shillong, Meghalaya.

PCA and Cluster analysis

Fifteen morphological characters (Table 3) were analyzed based on using principal component analysis (PCA) and cluster analysis (Hammer et al. 2001). Multivariate PCA and hierarchical cluster analysis were assessed using the software PAST 4.06b version.

RESULTS

Glochidion: Morphological diagnosis

Monoecious and rarely dioecious; primarily of shrubs or trees; pubescent or glabrous; droopingly branched. Leaves simple, alternate, usually asymmetrical at the base, entire, petiolate, stipulate. The inflorescence is usually axillary, supra-axillary, or pedunculate, with few to many flowers. Staminate flowers are mostly long pedicellate, with sepals 3–6, free, but no petals or disc. Anthers are present, 3–12, connective, pistillode absent. Pistillate flowers shortly pedicellate or sessile; sepals 3–6, free to connate; petals or disc absent; staminodes absent. Ovary 3–14 locular, biovulate locules; styles usually connate into a column, conical, or globose. Capsular pedicellate or sessile with a style column at the apex, depressed, subglobose, unlobed, and deeply or conspicuously lobed; pubescent or glabrous, green, white, or creamy to reddish. Seeds are usually 3–14, compressed, hemispherical with an arillate coat.

Review on the ethnobotanical knowledge of members of Glochidion in India

Some ethnobotanical uses of members of the genus Glochidion were mentioned by earlier workers. These are given below-

The paste of Glochidion tomentosum Dalz. is used externally in wounds by the tribes of Eastern Ghat, India (Reddy et al. 2006). The Chiru tribe of Manipur, India, consumed young leaves of G. multiloculare (Rottler ex. Willd.) Voigt and cooked them as an enjoyable curry and used them against stomach disorders (Rajkumari et al. 2013). The bark of G. multiloculare is used for skin diseases and wounds (Bajpai et al. 2016). Roots of G. multiloculare are used in snake bites (Brahma et al. 2002). The fruit and stem of G. heyneanum (Wight & Arn.) Wight is used in diabetes, fever, and bone fracture (Kumar et al. 2019). Roots of G. heyneanum are used in snake bites (Bajpai et al. 2016). Barks and leaves of G. zeylanicum (Gaertn.) A.Juss. are used in snake bites and stomach ulcers, and tender shoots are applied to itches (Das et al. 2013; Chakrabarty & Balakrishnan 2018; Kumar et al. 2019). Branches and leaves of G. sphaerogynum (Mull.Arg.) Kurz. are used in influenza and eczema (Lalrinkimi & Lallianthanga 2019). Fruits of G. daltonii (Mull.Arg.) Kurz. is used in cough and dysentery, and the bark of G. ellipticum Wight is used in inflammation (Bajpai et al. 2016). The paste made from the seeds of G. ellipticum is used as an antiallergic (Babu 1995). The bark of G. lanceolarium (Roxb.) Voigt is used in stomach diseases and is used as an anti-itch drug, oil made from seeds is also used as a source of light (Chanda et al. 2007; Bajpai et al. 2016; Chakrabarty & Balakrishnan 2018). The seeds of G. calocarpum Kuna are applied externally for skin diseases, and leaves are used orally to cure fever (Elanchezhian et al. 2007). According to Lalfakzuala et al. (2007), fruits of G. arborescens are used as wild edible fruits that are consumed by the local people of Mizoram. The fruits of G. khasicum (Mull.Arg.) Hook.f. are also edible and consumed by the tribal people of the Khasi hills in Meghalaya (Chakrabarty & Balakrishnan 2018).

DISCUSSION

The comparative morphological characteristics of certain species of the genus Glochidion showed many similarities and distinctive characteristics, which can be helpful for the identification and classification of the taxa (Table 2). Some of the major distinctive characteristics were leaf morphology, petiole length, and reproductive structures, i.e., inflorescences, male and female flowers, anthers, style, ovary, and capsule. The presence or absence of hairs on stems, leaves, inflorescence, and capsules also significantly differentiates the taxa. G. multiloculare var. pubescens an endemic variety of Assam showed nearly identical habit, vegetative and reproductive characteristics, with the exception of a glabrous plant body in G. multiloculare var. multiloculare (Chakrabarty & Balakrishnan 2018). The other major distinctive characteristics of both the taxa are that solitary or individual flowers and fruits occur in each axil in G. multiloculare var. pubescens while multiple flowers and fruits in each axil of the plant body have been observed in G. multiloculare var. multiloculare. The variety G. zeylanicum var. tomentosum presented almost the same character as G. zeylanicum var. zeylanicum, with the major difference being its hairy or tomentose character. G. heyneanum showed puberulous habits on the stems and leaves. In some taxa, leaves were asymmetric or symmetric at the base. 

The majority of taxa exhibited axillary inflorescence while G. zeylanicum var. zeylanicum, G. zeylanicum var. tomentosum and G. zeylanicum var. arborescens showed supra-axillary or pedunculate and rarely axillary inflorescence. Male flowers of the taxa revealed remarkably similar traits, but the number of anthers separated them. Female flowers presented different characteristics from male flowers. The peduncles of all the female flowers were shorter than the male flowers. The number of locules in the ovary varied by taxon, and style characters also played a key role. The shape, size, color, locules, and hairy habit of the capsule were distinguished among taxa, which showed taxonomic significance. Some taxa like G. zeylanicum var. zeylanicum and their varieties, were easily identified with their unlobed and ambiguously lobed capsule. G. multiloculare and G. sphaerogynum exhibited deeply or conspicuously lobed capsules while G. ellipticum presented a superficially lobed capsule. The capsule of G. lanceolarium was sessile, i.e., the fruit without the stalk or it lacked a pedicel, which distinguished it from other taxa. When dried, most of the leaves of the members were curled at the margin. While Glochidion and Epicephala moths were mutualists (Kato et al. 2003), most Glochidion leaves and drooping branches were found in insect-damaging conditions. As a result, some easily detectable characters for taxa identification could exist.

Based on morphological data, both PCA and cluster analysis were analyzed (Table 3–4 & Figure 2–3). The first PCA variance was 83.254% with an eigenvalue of 41.3263 followed by the second PCA variance of 8.5804% with an eigenvalue of 4.25921. The line connected to PC1 and PC2 makes up 91.8344% of the total variance and is a good sign of the variability of the initial data. PC1 represented the variation of the taxa based on the characters such as leaf shape (LSh), leaf apex and base (LAB), anther (AN), female flower (FF), sepal of the female flower (SF), ovary (OV), capsule color (CC), seed color (SC) and PC2 represented the characters such as leaf surface (LS), petiole length (PL), inflorescence (INF), male flower (MF), sepal of male flower (SM), style (ST), capsule (CP). In PC1, five taxa were observed i.e., G. ellipticum (GE), G. lanceolarium (GL), G. multiloculare var. pubescens (GMP), G. sphaerogynum (GS), and G. zeylanicum var. tomentosum (GZT). PC2 denoted a total of four taxa, viz., G. heyneanum (GH), G. multiloculare var. multiloculare (GM), G. zeylanicum var. arborescens (GZA), and G. zeylanicum var. zeylanicum (GZZ). From cluster analysis the tree revealed that G. multiloculare var. multiloculare (GM) and G. multiloculare var. pubescens (GMP) as cluster 1, G. heyneanum (GH) as cluster 2, G. sphaerogynum (GS) as cluster 3, G. ellipticum (GE) and G. lanceolarium (GL) as cluster 4, G. zeylanicum var. arborescens (GZA), and G. zeylanicum var. tomentosum (GZT) as cluster 5, G. zeylanicum var. zeylanicum (GZZ) as cluster 6. Taxa present in the same cluster specified more correlation than the taxa present in the different clusters.

The ethnobotanical study revealed that some members of the genus Glochidion were traditionally used to cure different diseases (Table 5) in Assam. Among the uses, the most frequent are skin diseases, fractures, body swelling, and snake bites.

CONCLUSION

The study revealed that there are similarities and differences among the members of the genus Glochidion which are more reliable for grouping and classifying the taxa. Documentation of ethnobotanical evidence signifies the importance of the genus. Both primary and secondary sources of the ethnobotanical knowledge showed the members have medicinally important properties and almost all parts, i.e., leaves, bark, and roots, have been used by the local people for the treatment of various diseases in India including Assam. This study summarized that both taxonomical study and conservation of ethnobotanical knowledge are of great significance, with the ability to stimulate subsequent biological investigation. Moreover, PCA and cluster analysis also validated the data on comparative morphological traits that showed correlation and variation among the analyzed species.

Table 1. List of recorded taxa in the studied area with their locality, accession number, and GPS coordinates.

Taxa	Locality	Accession No.	GPS coordinates
Glochidion ellipticum Wight	Kokrajhar District, Assam	98605	26.4947^0N, 90.4319^0E
G. heyneanum (Wight & Arn.) Wight	Kokrajhar District, Assam	98606	26.6236^0N, 90.4061^0E
G. lanceolarium (Roxb.) Voigt	Chakrashila Wildlife sanctuary, Kokrajhar District, Assam	98608	26.4236^0N, 90.4963^0E
G. multiloculare (Rottler ex Willd.) Voigt	Kokrajhar District, Assam	98604	26.7338^0N, 90.4308^0E
G. multiloculare var. pubescens Chakrab. & M.Gangop.	Orang National Park, Udalguri District, Assam	98610	26.7858^0N, 92.3305^0E
G. sphaerogynum (Mull.Arg.) Kurz	Chakrashila Wildlife Sanctuary, Kokrajhar District, Assam	98609	26.2902^0N, 90.3747^0E
G. zeylanicum var. arborescens (Blume) Chakrab. & M.Gangop.	Ultapani Forest Range, Kokrajhar District, Assam	98603	26.8002^0N, 90.3466^0E
G. zeylanicum var. tomentosum Trimen.	Ultapani Forest Range, Kokrajhar District, Assam	98607	26.7722^0N, 90.4158^0E
G. zeylanicum (Gaertn.) A.Juss	Nokpakghat, Karbi Anglong District, Assam	98611	26.3838^0N, 93.2061^0E

Table 2. Comparative morphological characters of certain species of genus Glochidion collected from different localities of Assam.

Characters	G. ellipticum	G. heyneanum	G. lanceolarium	G. multiloculare var. multiloculare	G. multiloculare var. pubescens	G. sphaerogynum	G. zeylanicum var. arborescens	G. zeylanicum var. tomentosum	G. zeylanicum var. zeylanicum
Leaf shape	Elliptic to lanceolate, oblong to obovate	Ovate to elliptic, obovate	Lanceolate to oblanceolate, elliptic	Oblong to lanceolate, elliptic to oblanceolate	Oblong to lanceolate, elliptic to oblanceolate	Oblong to elliptic, falcate	Ovate to elliptic	Ovate to elliptic, cordate	Ovate to elliptic, cordate
Leaf apex & and base	Apiculate, caudate, acuminate at apex, obtuse at base	Acute, apiculate at apex, obtuse or rounded at base	Apiculate, acuminate, or acute at apex, obtuse or rounded at base	Acute, apiculate, or retuse at apex, obtuse or rounded at base	Acute, apiculate, or retuse at apex, obtuse or rounded at base	Acuminate at apex, attenuate at base	Acute, acuminate at apex, obtuse or rounded at base	Obcordate, acute at apex, obtuse, truncate, asymmetric at base	Acute, apiculate at apex, cordate, asymmetric, truncate at base
Leaf surface	Glabrous on both surfaces	Pubescent on both surfaces and densely pubescent beneath	Glabrous on both surfaces	Glabrous on both surfaces at mature and pubescent at young	Pubescent on both surfaces and densely pubescent beneath	Glabrous on both surfaces	Densely pubescent on both surfaces	Densely pubescent on both surfaces	Glabrous on both surfaces
Petiole length	0.4–1 cm long	0.1–0.5 cm long	0.6–1 cm long	0.1–0.5 cm long	0.1–0.5 cm long	0.9–1 cm long	0.3–0.5 cm long	0.5–0.7 cm long	0.1–0.8 cm long
Inflorescence	Axillary	Axillary	Axillary	Axillary	Axillary	Axillary	Supra-axillary, pedunculate, rarely axillary	Supra-axillary, pedunculate, rarely axillary	Supra-axillary, pedunculate, rarely axillary
Male flower	Pedicellate, 0.5–1.7 cm long	Pedicellate, 0.5–1 cm long	Pedicillate, 0.9–2 cm long	Pedicellate, 0.5–1 cm long	Pedicellate, 0.5–1 cm long	Pedicellate, 0.5–1.8 cm long	Pedicellate, 0.5–0.7 cm long	Pedicellate, 0.7–1.8 cm long	Pedicellate, 0.5–1 cm long
Sepal	6	6	6	6	6	6	6	6	6
Anther	4–5	3–4	4–6	5–12	5–12	3–5	5–7	5–8	3–8
Female flower	Pedicellate, 0.1–0.5 cm long	Pedicellate, 0.1–0.6 cm long	Sessile, 0.06–0.09 cm long	Pedicellate, 0.3–0.5 cm long	Pedicellate, 0.3–0.5 cm long	Pedicellate, 0.3–0.5 cm long	Pedicellate, 0.4–0.6 cm long	Pedicellate, 0.4–1 cm long	Pedicellate, 0.1–0.9 cm long
Sepal	6	6	6	6–12	6–12	6	6	6	6
Style	Columnar, conical	Columnar	Columnar	Conical, subglobose	Conical, subglobose	Discoid	Columnar, persistent	Columnar, subconical	Free
Ovary	Subglobose, 3–6 locular	Depressed, subglobose, 4–5 locular	Depressed, subglobose, 5–8 locular	Depressed, 5-12 locular	Depressed, 5-12 locular	Depressed, 4–12 locular	Subglobose, 4–6 locular	Depressed, Subglobose, 5–8 locular	Depressed, Subglobose, 4–8 locular
Capsule	Pubescent, pedicellate, shallowly lobed	Pubescent, pedicellate, conspicuously lobed	Sparsely pubescent, sessile, shallowly lobed to deeply lobed	Pubescent, pedicellate, conspicuously lobed	Pubescent, pedicellate, conspicuously lobed	Glabrous, pedicellate, ambiguously lobed	Densely pubescent, pedicellate, obviously unlobed	Densely pubescent, pedicellate, ambiguously lobed	Glabrous, pedicellate, ambiguously lobed
Capsule color	Light green to creamy, whitish	Green to yellow-green	Light green, creamy to reddish	Green	Green	Green	Light green, yellow green to reddish	Light green, yellow green to reddish	Light green, yellow green to reddish
Seed color	Yellow green to red	Yellow green to orange	Yellow green to red	Green to red	Green to red	Green to orange	Yellow green to red	Yellow green to red	Yellow green to red

Table 3. Character state of morphological characteristics of different members of Glochidion used in cluster analysis.

Taxa	LSh	LAB	LS	PL	INF	MF	SM	AN	FF	SF	ST	OV	CP	CC	SC
E	1	1	2	1	0	1	0	1	1	1	1	1	1	1	1
H	4	4	1	0	0	0	0	4	3	1	2	4	2	3	3
L	2	2	2	2	0	2	0	2	2	1	2	2	2	2	1
M	0	0	0	0	0	0	0	0	0	0	0	0	0	0	0
MP	0	0	1	0	0	0	0	0	0	0	0	0	0	0	0
S	3	3	2	3	0	3	0	3	0	1	3	3	3	0	2
ZA	6	6	1	5	1	4	0	6	5	1	5	6	4	4	1
ZT	5	7	1	6	1	5	0	7	6	1	6	2	5	4	1
ZZ	5	5	2	4	1	0	0	5	4	1	4	5	3	4	1

E—G. ellipticum | H—G. heyneanum | L—G. lanceolarium | M—G. multiloculare var. multiloculare | MP—G. multiloculare var. pubescens | S—G. sphaerogynum | ZA—G. zeylanicum var. arborescens | ZT—G. zeylanicum var. tomentosum | ZZ—G. zeylanicum var. zeylanicum | LSh—Leaf shape | LAB—Leaf apex & base | LS—Leaf surface | PL—Petiole length | INF— Inflorescense | MF—Male flower | SM—Sepal of male flower | AN—Anther | FF—Female flower | SF—Sepal of female flower | ST—Style | OV—Ovary | CP—Capsule | CC—Capsule color | SC—Seed color

Character states: Leaf Shape: Oblong to lanceolate, elliptic to oblanceolate = 0; Elliptic to lanceolate, oblong to obovate = 1; Lanceolate to oblanceolate, elliptic = 2; Oblong to elliptic, falcate = 3; Ovate to elliptic, obovate = 4; Ovate to elliptic, cordate = 5; Ovate to elliptic = 6; Leaf apex & base: Acute, apiculate or retuse at apex, obtuse or rounded at base = 0; Apiculate, caudate, acuminate at apex, obtuse at base = 1; Apiculate, acuminate or acute at apex, obtuse or rounded at base = 2; Acuminate at apex, attenuate at base = 3; Acute, apiculate at apex, obtuse or rounded at base = 4; Acute, apiculate at apex, cordate, asymmetric, truncate at base = 5; Acute, acuminate at apex, obtuse or rounded at base = 6; Obcordate, acute at apex, obtuse, truncate, asymmetric at base = 7; Leaf surface: Glabrous on both surfaces at mature and pubescent at young = 0; Pubescent on both surfaces and densely pubescent beneath = 1; Glabrous on both surfaces = 2; Petiole length: 0.1–0.5 cm long = 0; 0.4–1 cm long = 1; 0.6–1 cm long = 2; 0.9–1 cm long = 3; 0.1–0.8 cm long = 4; 0.3–0.5 cm long = 5; 0.5–0.7 cm long = 6; Inflorescence: Axillary = 0; Supra-axillary, pedunculate, rarely axillary = 1; Male flower: Pedicellate, 0.5–1 cm long = 0; Pedicellate, 0.5–1.7 cm long = 1; Pedicillate, 0.9–2 cm long = 2; Pedicellate, 0.5–1.8 cm long = 3; Pedicellate, 0.5–0.7 cm long = 4; Pedicellate, 0.7–1.8 cm long = 5; Sepal of male flower: 6 = 0; Anther: 5–12 = 0; 4–5 = 1; 4–6 = 2; 3–5 = 3; 3–4 = 4; 3–8 = 5; 5–7 = 6; 5–8 = 7; Female flower: Pedicellate, 0.3–0.5 cm long = 0; Pedicellate, 0.1–0.5 cm long = 1; Sessile, 0.06–0.09 cm long = 2; Pedicellate, 0.1–0.6 cm long = 3; Pedicellate, 0.1–0.9 cm long = 4; Pedicellate, 0.4–0.6 cm long = 5; Pedicellate, 0.4–1 cm long = 6; Sepal of female flower: 6–12 = 0; 6 = 1; Style: Conical, subglobose = 0; Columnar, conical =1; Columnar = 2; Discoid = 3; Free = 4; Columnar, persistent = 5; Columnar, subconical = 6; Ovary: Depressed, 5–12 locular = 0; Subglobose, 3–6 locular = 1; Depressed, subglobose, 5–8 locular = 2; Depressed, 4–12 locular = 3; Depressed, subglobose, 4–5 locular = 4; Depressed, Subglobose, 4–8 locular = 5; Subglobose, 4–6 locular = 6; Capsule: Pubescent, pedicellate, conspicuously lobed = 0; Pubescent, pedicellate, shallowly lobed = 1; Sparsely pubescent, sessile, shallowly lobed to deeply lobed = 2; Glabrous, pedicellate, ambiguously lobed =3; Densely pubescent, pedicellate, obviously unlobed =4; Densely pubescent, pedicellate, ambiguously lobed; Capsule color: Green = 0; Light green to creamy, whitish = 1; Light green, creamy to reddish = 2; Green to yellow green = 3; Light green, yellow green to reddish = 4; Seed color: Green to red = 0; Yellow green to red = 1; Green to orange = 2; Yellow green to orange = 3

Table 4. Principal component analysis (PCA) based on morphological characteristics of different members of Glochidion.

PC	Eigenvalue	% variance
1	41.3263	83.254
2	4.25921	8.5804
3	2.02297	4.0754

Table 5. Enumeration of ethnobotanical knowledge of Glochidion in Assam.

Botanical name	Vernacular names	Parts used	Mode of preparation	Application	Route of administration
Glochidion multiloculare (Rottler ex Willd.) Voigt	Thakha Biphang or thakha mala (Bodo), Gorumora, Dolpoduli (Assamese)	Leaves, barks, Roots	- A small number of leaves are ground into a paste. - A small amount of bark is taken and ground into a paste. - 3–4 roots are taken to make a paste.	- Fracture and body swelling - Skin diseases and wounds - Snake bite	- External - External - External
G. ellipticum Wight	Thakha Biphang or thakha mala (Bodo), Panimadhuri (Assamese), Latimaowa (Nepali)	Bark, stem branches, and roots	- A small amount of bark is removed and ground into a paste, which is then administered to the diseased area. - A paste is made by crushing the stem and applying it to the swelling area. - Roots are ground into a paste.	- Body swelling, Skin problem - Body swelling - Snake bite	- External - External - External
G. sphaerogynum (Mull.Arg.) Kurz.	Thakha Biphang or thakha mala (Bodo), Panimadhuri (Assamese), Boljakru (Garo)	Young branches and leaves	Young branches and leaves are ground into a paste and blended with a small amount of water.	- Skin diseases - Branches are used as firewood also.	- External

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