A NUMERICAL TAXONOMY OF Sida spp. (MALVACEAE) IN JAVA AND BALI ISLAND, INDONESIA

Abstract

Sida is a pantropical weed that can be annual or perennial. Several species have been identified in Indonesia. Sida cordifolia, S. rhombifolia, S. alnifolia, S. rhomboidea, S. acuta, S. glutinosa, and S. mysorensis were chosen for this study because they are representative of the species that are extensively spread in Indonesia. This study aims to update taxonomic information on the Sida genus in Indonesia. A phenogram was created using the data that was derived from the scoring of 70 morphological features. At PAST, the phenogram was created using cluster analysis, and Principal Cluster Analysis was used to determine the reason for clustering. The first cluster shows the separation of Sida acuta from all OTUs at a distance of 0.30. The second cluster shows the separation of OTU Sida cordifolia from OTU Sida glutinosa and OTU Sida mysorensis cluster at a distance between 0.20 until 0.25. The third cluster shows the separation of OTU Sida rhomboidea from OTU Sida rhombifolia and OTU Sida alnifolia cluster at a distance that was close to 0.20. The biggest value of PCA results is -0.45 and belongs to awn tip which is a character of mericarps. It demonstrates how crucial the mericarp trait is to identifying and appreciating the diversity of the genus Sida.

Headnote

ABSTRACT

Sida is a pantropical weed that can be annual or perennial. Several species have been identified in Indonesia. Sida cordifolia, S. rhombifolia, S. alnifolia, S. rhomboidea, S. acuta, S. glutinosa, and S. mysorensis were chosen for this study because they are representative of the species that are extensively spread in Indonesia. This study aims to update taxonomic information on the Sida genus in Indonesia. A phenogram was created using the data that was derived from the scoring of 70 morphological features. At PAST, the phenogram was created using cluster analysis, and Principal Cluster Analysis was used to determine the reason for clustering. The first cluster shows the separation of Sida acuta from all OTUs at a distance of 0.30. The second cluster shows the separation of OTU Sida cordifolia from OTU Sida glutinosa and OTU Sida mysorensis cluster at a distance between 0.20 until 0.25. The third cluster shows the separation of OTU Sida rhomboidea from OTU Sida rhombifolia and OTU Sida alnifolia cluster at a distance that was close to 0.20. The biggest value of PCA results is -0.45 and belongs to awn tip which is a character of mericarps. It demonstrates how crucial the mericarp trait is to identifying and appreciating the diversity of the genus Sida.

Keywords: biodiversity, biosystematic, malvaceae, taxonomical data. running title: A numerical taxonomy of sida in Indonesia

INTRODUCTION

Sida is one of heterogeneous genera with highest number of species in Malveae (8, 14; 27). Sida refers to a genus in family Malvaceae, subfamily Malvoideae, tribe Malveae and subtribe Abutilinae. This genus consists of several sections, including Cordifoliae, Distichifoliae, Ellipticifoliae, Hookerianae, Malacroideae, Muticae, Nelavagae, Oligandrae, Sidae, Spinosae and Stenindae (1, 29). Almost all Sida species knew as annual or perennial common weeds in highland and lowland areas. Mabberley (18) recorded 250 species of Sida spread across the tropics and subtropics. In Indonesia, Sida occurs in waste places, teak forests, coconut plantations, along roadsides, and near the coast. Taxonomical research of Sida in Indonesia has recorded the presence of several species. Baker & Bakhuizen (5) and in Flora of Java and Malesian Malvaceae Revised noted that there are twelve species in Java, which include S. acuta, S. balica, S. cordifolia, S.elongata, S. glutinosa, S. javensis, S. mysorensis, S. retusa, S. rhombifolia, S. spinosa, S. subcordata, and S. veronicaefolia. While Steenis (25) only recorded S. rhombifolia and S. acuta on the island of Java. Generally, Sida is recognized as a wild shrub with serrated leaf margin, yellow flowers, and brownish schizocarpic fruit. However, Sida has distinctive features from related genera in tribe Malveae. According to Brandao et al. (8) and Aguilar et al. (1), the calyx of Sida is 10ribbed and the schizocarpic fruit is polycoccus type, uniseminate and consist 5-14 mericarps which is trigonous shaped and muticous to prominently aristate. Commonly used as ethnomedical medicine in several countries such as Brazil, Colombia, Nigeria, Philippines, Central America, and India. Sida has benefits to treat rheumatism, asthma, facial paralysis, gonorrhea, diarrhea, snake bites, and fever. This plant is also used for the treatment of wounds, anti-inflammatory, and skin diseases (22). According to Bai et al. (5), Sida are used in various Ayurvedic and ethnic medicines. Phytochemical studies aimed by Aminah et al. (2)revealed Sida has several secondary metabolisms such as alkaloid, flavonoid, coumarins, ecdysteroids, triterpenes, and tocopherols. In Indonesia, S. rhombifolia itself is the one of Indonesian herbal medicine list from the Indonesian Ministry of Health scientific program (17). According to Choudhuri (10), numerical taxonomy refers to phenetic that attempts to group species into higher taxa based on overall similarity, usually in morphology. Phenetic is also one of major approach to biological classification beside phyletic and cladistic (26). Because morphology-based grouping is always obvious and understandable, numerical taxonomy can serve as a basic classification. It can be seen which characters are different between taxon (OTU) and also know the characters that affect taxon grouping. This study is usually carried out on non-cultivated and cultivated plants. For cultivated plants such as Begonia L. (24), it can be a reference for hybridization to obtain new hybrids. For non-cultivated plants such as wild Datura L. (9) and Abutilon author? (2), it provides data on the distribution of the taxon and data on taxon grouping which can be different or the same as taxon's original classification. The goal of this study was to identify relationships based on morphological characteristics. Ancient classification systems, naming, identification, and descriptions of plants were all based on morphological characteristics. Almost all taxonomic issues are still resolved using morphological features, which may be applied at all taxonomic levels, from the variety to the division (26). The capacity to observe plant diversity more easily is another advantage morphological characteristics have over other traits (21). The resulting grouping shows the most frequent morphological similarities across taxa, which may different if researched using other bases. The study was carried out in an effort to advance knowledge of Indonesian biodiversity, particularly in genus Sida (Malvaceae). Circumscription of Malvaceae has changed as the result of recent scientific advancements, making it crucial to understand the variety of morphological characteristics among its members. The gathered information will reveal what morphological traits exist in Sida and how they influence the relationships between Sida species.

MATERIALS AND METHODS

The research was conducted at the Biosystematics Laboratory, Department of Biology, Faculty of Science and Technology, Airlangga University, East Java, Indonesia. This research was conducted from January to June 2020. Plant materials were collected from living specimen ad literature of plant description (Table 1). S. cordifolia and S. acuta were found in the West Bali National Park. Specimens of S. rhombifolia, S. alnifolia and S. rhomboidea were found in Kediri (East Java). Two specimens namely S. glutinosa and S. mysorensis were obtained from papers or scientific publications entitled Malesian Malvaceae Revised. Malesian Malvaceae Revised, a scientific journal, provided the species description, and morphological characteristics were derived by observing the sample. The morphological characters used in this study were 70 characters which included 20 leaf characters, 17 flower characters, 10 fruit characters, 10 mericarp characters, 6 seeds characters and 7 stem characters which 24 quantitative characters and 46 qualitative characters (Table 2). The numbered morphological data (Table 3) exacted to cluster analysis and principal component analysis were performed by PAST 4.3. According to Permata & Susandarini (2022), cluster analysis using the UPGMA clustering method and Gower coefficient of similarity produces a phenogram that shows the relationship between species. Meanwhile, principal componen analysis shows the large character effect on grouping.

RESULTS AND DISCUSSION

Numerical taxonomy studies among OTUs (seven species of Sida) presented in the phenogram (Figure 1). There are three main clusters that appear on the phenogram. The first cluster shows the separation of OTU S. acuta from all OTUs at a distance of 0.30. The second cluster shows the separation of OTU S. cordifolia from OTU S. glutinosa and OTU S. mysorensis cluster at a distance between 0.20 until 0.25. The third cluster shows the separation of OTU S. rhomboidea from OTU S. rhombifolia and OTU S. alnifolia cluster at a distance that was close to 0.20. The first cluster shows that OTU S. acuta is separated from remaining species at a distance of 0.30 (Figure 1). This is because it has a distinctive morphological character. This plant has an acute leaf apex, oblong leaf circumscription, lawn green color in the petiole and the lateral stipule, burlywood color in the pedicel, and the calyx that covers the fruit and green almost tan color in stem. The separation between the second (sect. Cordifoliae and Nelavaga) and third (sect. Sida) clusters is at a distance that is close to 0.25. This separation is caused by twelve morphological characters including leaf base, leaf circumscription, leaf width near apex, petiole length, petiole color, lateral stipule color, pedicel color, pedicel length, color of calyx that covers the fruit, fruit pedicel color, awn tip , and stem color. The second cluster consists of OTU S. glutinosa and OTU S. mysorensis which are members of the section Nelavaga and OTU S. cordifolia which are members of the section Cordifoliae. OTU S. cordifolia is separated from the members of the section Nelavaga at a distance between 0.20 and 0.25 due to several morphological characters, such as leaf apex, leaf circumscription, leaf length near the base, leaf width near the base, lateral stipule length, corolla length, corolla width, color of corolla, fruit pedicel length, presence of jointed in fruit pedicel, number of mericarp on fruit, and the presence of veins on the right and left radial mericarp. Meanwhile, all members of the third cluster consist of members of the section Sida, including OTU S. rhombifolia, OTU S. alnifolia, and OTU S. rhomboidea. In this observation, seven Sida species have several same morphological characters such as; the abaxial and adaxial leaf surface is pubescent, serrate leaf margin, folium simplex leaf, pubescent petiole surface, pair lateral stipules, the calyx is five numbered, pubescent calyx surface, calyx apex is acute, the calyx is lawngreen coloured, epicalyx is absent, axillar phyllotaxis, anther is yellow coloured, the corolla is five numbered and valvate aestivation, monoadelph stamen, polycoccus fruit, the calyx is five numbered and pubescent, an axillary spot of fruit pedicel, mericarp is a trigonous shape, pair awn, pubescent awn surface, single seed, glabrous seed surface, seed colour is black saddle brown, hilus spot in apex, shrubs, teres stem, and cambium is present. Among 70 morphological characters, there are 6 morphological characters that greatly affect the clustering of OTUs (the seven species of Sida). There are values greater than 0.20 or less than -0.20. The six morphological characters such as petiole color (PC 0.30), lateral stipule color (LSCR 0.30), color of the calyx that covers the fruit (CCF 0.30), color of fruit pedicel (CFP 0.30), stem color (SCR 0.30) and awn tip (TAW -0.45). Then, leaf circumscription (LC) and presence of epicalyx (PEC) which has an exact value of 0.20 indicates that both morphological characters are quite influential on the clustering of OTUs. The morphological character approach, which considers the characteristics of leaves, flowers, fruits, mericarps, seeds, and stems, was used to conduct the numerical taxonomy analysis of the seven Sida species. All of plant organs have an important role to distinguish between species but to determine Sida from remaining members of the Malvaceae, floral and seed features are the principal morphological traits. According , all of Sida species in the Malesia region have a yellow corolla. The problem is they appear to be very similar to one another at first glance. Therefore, to differentiate between species, the majority of taxonomists rely on the traits of the leaves and mericarp. Foliar researches such as Saibaba & Rao (22), Aworinde et al. (4), and also Khunnur & Kotresha (15) studied leaf venation and various leaf circumscription of several Sida species. The researches show the differences in leaf venation and circumscription between species and can be used as an identification reference. Mericarp or mericarpium is often used by taxonomists of the genus Sida in identifying a species, delimiting related taxa such as sections and species, or discovering to a new species or variety. Several studies such as; Yoshikawa et al. (29) released the species Sida uniaristata sp. nov. using the singular mericarpium awn character, Verdcourt (28) published the reviews the mericarpium of each varieties of Sida rhombifolia in East Africa, and Bharati (6) was published the reviews characteristics of various Sida species that are identified by mericarpium. Other traits also are seen in the color of the lateral stipule, petiole, and stem. All section Sida members (S. rhombifolia, S. rhomboidea, S. alnifolia) have a maroon or reddish color on lateral stipule, petiole, and stem. Petiole anatomy research of Sida by Olatunji and Bakare (8) shows that tanniniferous substances are responsible for red coloration. Regularly, color characteristics are important to be used in the identification of all plant species. Genus Sida has several sections including Sida, Nelavaga, Cordifoliae and Distichifolia. In 1966, S. acuta, S. cordifolia, S. glutinosa and S. rhombifolia were in section Sida. This has been published in Malesian Malvaceae Revised which was written by Borssum Waalkes. This shows that almost all species were formerly members of the section Sida before switching sections. S. acuta has been a member of the section Distichifolia since 2003 (Krapovickas, 2003). Meanwhile, according to Fryxell (12). S. cordifolia is in the section Cordifoliae, and S. glutinosa is a synonym of S. glabra which this species belongs to the section Nelavagae. Section Sida contains three species of which two of them are considered by many taxonomists to be synonyms of S. rhombifolia itself. Sivarajan & Pradeep (23) in their publication described S. alnifolia and S. rhomboidea. The two species are not a synonym but closely related to S. rhombifolia. In this study, all three show differences in the own tip. S. rhombifolia has a retuse awn tip, S. alnifolia has a curved awn tip and S. rhomboidea has an obtuse awn tip. his own tip character is a differentiator that has a high value in principal component analysis. The different morphological characters were analyzed using PCA to find out the contribution value in the grouping of seven Sida species. The morphological character with the largest contribution or the most important role has a value greater than 0.20 or less than -0.20. The result shows in Figure 2. The greater value belongs to the awn tip that has -0,45. It shows that the character of mericarp is an important character to identify and see the diversity of the genus Sida. This is proven by two studies such as the identification of Sida species based on the mericarp such as Identification of Indian Sida through mericarp written by Bharati in 2015 and the discovery of S. uniaristata in 2019 by Yoshikawa et al. which is based on the one awn of a mericarp. As a heterogeneous and polyphyletic genus, Sida has undergone several reductions and resulted in 10 separate genera (1; 13; 8). Furthermore, there has been an infrageneric classification change in Sida species recently. Nomenclaturally, many species in this genus have synonyms and it cause confusion about the number of accepted species. Seeing this, taxonomic studies on the genus Sida continue to be carried out both phylogenetic and phenetic. This morphological data collection is expected to allow genus Sida to achieve the monophyletic status. In conclusion, Sida is a heterogeneous genus in the Malvaceae family of which there are seven species in Indonesia. S. acuta, S. rhombifolia, S. alnifolia, S. rhomboidea, S. cordifolia, S. glutinosa and S. mysorensis have many similarities and differences in morphological characteristics. These seven species are the member of section Sida, Distichifolia, Cordifoliae, and Nelavaga are divided into 3 groups, with mericarp as the main distinguishing tools which also often function as identification tools in this plant. Finally, this research is an attempt to provide numerical taxonomy data related to Sida spp. based on morphological characters which are expected to be useful for biosystematics and taxonomy in Indonesia.

ACKNOWLEDGEMENTS

Thank you to the Department of Biology at Airlangga University for conducting training and field observations in the West Bali National Park. Authors also thank staff of West Bali National Park for all kinds of relief during the observation. We are very grateful to Dr. Hamidah, Dr. Junairiah, and Almando Geraldi, Ph.D. from Airlangga University who has provided meaningful assistance to the authors.

Sidebar

Received:22/3/2021, Accepted:27/7/2021

References

REFERENCES

1. Aguilar J.F, Fryxell P.A, and R. K. Jansen 2003. Phylogenetic Relationship and Clasification of the Sida Generic Alliance (Malvaceae) Based on nrDNA ITS Evidence. Syst. Bot. 28 (2): 352-364. DOI: 10.1043/0363-6445-28.2.352

2. Alzahrani DA, Albokhari EJ, Khoj A. 2021. Taxonomic Studies on Some Members of the Genus Abutilón Mill. (Malvaceae). AJPS. 12: 199-220. DOI: 10.4236/ajps.2021.122012

3. Aminah NS, Laili ER, Rafi M, Rochman A, Insanu M, Tun K.N.W. 2021. Secondary Metabolite compounds from Sida Genus and their Bioactivity. Heliyon. 7 (4). DOI: 10.1016/j.heliyon.2021.e06682

4. Aworinde D.O, Ogundairo B.O, Erinoso 5.M, A.O. Olanloye 2012 Foliar Epiderml Studies of Some Nigerian Species of Sida Linn. (Malvaceae). J. Agric. Sci. 2 (2): 18-22. www.scholarly-journals.com

5. Bai MDA, Rani SPS, Balachandran S, Jayakumar G. 2012. The Use of Sida Plant in the Preparation of Nayapayam Kashayam. IJRAP. 3 (1): 99-104. www.ijrap.net

6. Bharati K. A. 2016. Identification of Indian Sida through mericarp. Pharmacogn. J. 8 (5): 490-496. DOI: 10.5530/pj.2016.5.14

7. Borssum Waalkes, Jan van. 1966. Malesian Malvaceae revised. Blumea. 14 (1): 1-213.

8. Brandao J.S, Baracho G.S, De Sales M.F, and M.P.V. Filho 2017. Synopsis of Sida (Malvaceae, Malvoideae, Malveae) in the State of Pernambuco, Brazil. Phytotaxa. 307 (3): 205-227. DOI: 10.11646/phytotaxa.307.3.5

9. Cavazos M.L. 2000. Phenetic analysis of Datura section Dutra (Solanaceae) in Mexico. Bot. J. Linn. Soc. 133: 493-507. DOI: 10.1006/bojl.2000.0337

10. Choudhuri S. 2014. Fundamentals of Molecular Evolution. In: Bioinformatics for Beginners. Elsevier, Amsterdam

11. Dorly, Ningrum R.K, Suryantari N.K, and F.L.R. Anindita 2016. Leaf Anatomical Study of Three Members of Malvaceae Family in Jatiluhur Resevoir Area. Proceeding Biology Education conference. Sebelas Maret University, Surakarta, 2016. [Indonesian]

12. Fryxell P.A. 1985. Sidus Sidarum - V. The North and Central American Species of Sida. SIDA, Contributions to Botany . 11 (1): 62-91.

13. Fryxell P. A. 1997. The American Genera of Malvaceae-II. Brittonia. 49 (2): 204-269

14. Fryxell P. A. 2002. An Abutilón Nomenclator (Malvaceae). Lundellia. 5: 79-118. DOI: 10.25224/1097-993X-5.1.79

15. Khunnur SB, Kotresha SK. 2012. Systematics of Flowering Plants 73 Foliar Studies in Some Species of Sida L. (Malvaceae). Conference: Multidisciplinary Approaches in Angiosperm Systematics Volume 1-2. West Bengal. [Indian]

16. Krapovickas A. 2003. Sida Seccion Distichifolia (Monteiro) Krapov. comb. nov. stat. (Malvaceae-Malveae). Bonplandia. 12 (1-4): 83-121. DOI: 10.30972/bon.121-41407

17. Laili E.R, Aminah N. S, Kristanti A.N, Wardana A.P, Rafi M, Rohman A, Insanu M, and K.N.W. Tun 2022. Comparative Study of Sida rhombifolia from Two Different Locations. Rasayan J. Chem. 15 (1): 642-650. DOI: 10.31788/RJC.2022. 1516588

18. Mabberley, D. J. 2017. Mabberley's Plant-book: A Portable Dictionary of Plants, Their Classification and Uses. Cambridge University Press, Cambridge

19. Olatunji O. A, and O.A. Bakare 1993. Taxonomic Value of The Petiole Anatomy in The Genus Sida L. (Malvaceae) in Nigeria. Feddes Repert. 104: 35-39

20. Permata D.A, and R. Susandarini 2022 Morphological Diversity and Phenetic Relationship of Wild and Cultivated Begonia based on Morphology and Leaf Venation. Biodiversitas. 23 (2): 928-937. DOI: 10.13057/biodiv/d230235

21. Rahayu S. E., Chikmawati T, Kartawinata K, and A. Hartana 2012. Morphology vs Taxonomy in the Family Pandanaceae: A Case Study in the Javanese Species. Reinwardtia. 13 (4): 317-330. DOI: 10.14203/reinwardtia.v13i4.435

22. Rodrigues F. C, and A.F.M. de Oliveira 2020. The genus Sida L. (Malvaceae): An Update of its Ethnomedicinal Use, Pharmacology, and Phytochemistry. S. Afr. J. Bot. 132: 432-462. DOI: 10.1016/j.sajb.2020.04.030

23. Saibaba A.M, and R.S. Rao 1990. Leaf Venation Studies in Indian Sida. SIDA, Contributions to Botany. 14 (2): 215-222

24. Sivarajan V.V, and A.K. Pradeep 1994. Taxonomy of the Sida rhombifolia (Malvaceae) Complex in India. SIDA, Contributions to Botany. 16 (1): 63-78

25. Steenis C.G.G.J. 2008. Flora: Untuk Sekolah di Indonesia. Pradnya Paramita, Jakarta

26. Stuessy TS. 2009. Plant Taxonomy: The Systematic Evaluation of Comparative Data Second edition. Columbia University Press, New York

27. Tate J.A, Aguilar J.F, Wagstaff S.J, Duke J.C.L, Slota T.A.B, and B.B. Simpson 2005. Phylogenetic Relationship within the Tribe Malveae (Malvaceae, subfamily Malvoideae) as Inferred from ITS Sequence Data. Am. J. Bot. 92 (4): 584-602. DOI: 10.3732/ajb.92.4.584

28. Verdcourt B. 2004. The variation of Sida rhombifolia L. (Malvaceae) in East Africa. Kew Bull. 59: 233-239. DOI: 10.2307/4115855

29. Yoshikawa V.N, Duarte M.C, and V.M. Goncales 2019. Sida uniaristata, A New Species of Sida sect. Sidae L. (Malvaceae, Malvoideae) from Brazil. Syst. Bot. 44 (1): 184-188. DOI: 10.1600/036364419X698001